MONOGRAPH OF THE OXYTRICHIDAE

(CILIOPHORA, HYPOTRICHIA)
 MONOGRAPHIAE
BIOLOGICAE
VOLUME 78

Series Editors

H. J. Dumont and M. J. A. Werger

The titles published in this series are listed at the end of this volume
Monograph of the Oxytrichidae
(Ciliophora, Hypotrichia)

by

HELMUT BERGER
Consulting Engineering Office for Ecology,
Salzburg, Austria

SPRINGER-SCIENCE+BUSINESS MEDIA, B.V.

A C L P . Catalogue record for this book is available from the Library of Congress.

I S B N 978-94-010-5958-9 I S B N 978-94-011-4637-1 (eBook)

D O I 10.1007/978-94-011-4637-1

Printed on acid-free paper

A l l Rights Reserved
© 1999 Springer Science+Business Media Dordrecht
Originally published by Kluwer Academic Publishers in 1999
Softcover reprint of the hardcover 1st edition 1999
No part of the material protected by this copyright notice may be reproduced or
utilized in any form or by any means, electronic or mechanical,
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retrieval system, without written permission from the copyright owner.
 Dedication
This book is dedicated to my family, Elisabeth, Magdalena, Eva, Helena Valentina and
my parents, Maria and Johann t Berger
Contents
Preface .............................................................. ix
Acknowledgements .................................................... xi
A General Section ................ . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. 1
1 Morphology, Biology, and Tenninology .............................. 1
1·1 Size and Shape ...... . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. 1
1·2 Nuclear Apparatus . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. 1
1·3 Contractile Vacuole and Cytopyge .............................. 8
1·4 Cytoplasm and Colouring ..................................... 8
1·5 Cortical Granules ..................................... . . . . .. 8
1·6 Movement ................................................ 12
1·7 Somatic Infraciliature and Ultrastructure ........................ 15
1·8 Oral Apparatus ............................................ 24
1·9 Silverline System ................. . . . . . . . . . . . . . . . . . . . . . . . .. 26
1·10 Life Cycle ................................................ 26
1·10·1 Cell Division ........................................ 28
1·10·2 Conjugation ......................................... 36
1·10·3 Cyst ............................................... 43
1·10·4 Reorganisation, Regeneration, Doublets ................... 49
2 Phylogeny ..................................................... 56
3 Classification .................................................. 78
4 Parasitism ..................................................... 81
5 Ecology, Occurrence, and Geographic Distribution .................... 86
6 Collecting, Culturing, Observing, and Staining of Oxytrichid Ciliates 88
6·1 Collecting and Culturing ..................................... 88
6·2 Observing Living Hypotrichs ................................. 89
6· 3 Staining Procedures ......................................... 90
6·3·1 Feulgen Nuclear Reaction ............................... 91
6·3·2 Supravital Staining with Methyl Green-Pyronin .............. 91
6·3·3 Protargol Methods ..................................... 92
6·4 Preparation for Scanning Electron Microscopy ................... 99
7 Summary of New Taxa Described in this Book and of Nomenclatural Acts 100
B Systematic Section ............................................... 102
Oxytrichidae ( 169 species) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. 102
Oxytrichinae (113 species) ...................................... 113
Oxytricha (54 species) ....................................... 115
Allotricha (2 species) ........................................ 261
Cyrtohymena (15 species) .................................... 279
Notohymena (4 species) ...................................... 324
Urosomoida (7 species) ...................................... 345
Gonostomum (4 species) ..................................... 367
Urosoma (10 species) ....................................... 396
Tachysoma (10 species) ...................................... 431
Australocirrus (1 species) .................................... 470
viii

Onychodromopsis (2 species) ................................. 475

Rubrioxytricha (2 species) .................................... 479
Parurosoma (1 species) ...................................... 491
Pseudostrombidium (1 species) ................................ 494
Stylonychinae (37 species) ...................................... 499
Stylonychia (11 species) ...................................... 501
Coniculostomum (2 species) .................................. 606
Steinia (3 species) .......................................... 624
Histriculus (6 species) ....................................... 645
Sterkiella (7 species) .... . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. 668
Parastylonychia (1 species) ................................... 696
Pleurotricha (2 species) ...................................... 698
Rigidocortex (1 species) ...................................... 717
Onychodromus (2 species) .................................... 722
Laurentiella (1 species) ...................................... 752
Pattersoniella (1 species) ..................................... 766
Taxa of Unknown Position within the Oxytrichidae ................... 777
Ancystropodium (1 species) ...................................... 777
Apoamphisiella (2 species) ....................................... 781
Gastrostyla (8 species) ......................................... 789
Kerona (1 species) ............................................. 825
Paraurostyla (4 species) ..... . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. 841
Parentocirrus (1 species) ........................................ 878
Territricha (1 species) .......................................... 884
Pseudouroleptus (1 species) ...................................... 888
Taxa not Considered ................................................ 893
Illustrations of Species Indeterminata and Insufficient Redescriptions ...... 895
Addenda ........................................................... 925
References ......................................................... 938
Systematic Index .... . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. 1049
Table Index ....................................................... 1080
Preface

I have written this book because there is, as in almost all supraspecific ciliate taxa, an ur-
gent need for an up-to-date revision of the oxytrichids, which are common in terrestrial,
limnetic, and marine biotopes. The last comprehensive, illustrated guides to this group of
hypotrichs were provided by KAHL (1932) and SmLER (1974b); however, as regards syn-
onymy and faunistics, these works are outdated and not as detailed as EHRENBERG'S out-
standing book from 1838. In KAHL's revision, the oxytrichids sensu stricto are described
on about 30 pages, whereas in the present book the systematic section comprises about
830 pages. This extraordinary increase in page number is mainly due to the following
points: (i) Species number increased from about SO in KAHL to about 170 in the present
book. (ii) KAHL usually provided only a single illustration of each species, whereas almost
all published illustrations on oxytrichid ciliates are included in my book. (iii) Modern,
phylogenetic systematics of oxytrichids without morphogenetic data is impossible; conse-
quentlY,·almost all detailed descriptions, usually dealing with the type species, have been
included. (iv) Synonymy is discussed, and not only mentioned, as is unfortunately usual.
(v) Modem descriptions of oxytrichids, and ciliates in general, are much more compre-
hensive than the descriptions by KAHL; note, however, that STEIN'S (IS59b) description
on, for example, Stylonychia mytilus is 14 (folio-sized) pages long and includes 34 in-
comparable live drawings on 3 plates! (vi) Almost all physiologica~ ecological, and fau-
nistic literature available on each species is mentioned; the present book is therefore not
only a "field guide", but also a reference book.
Most species treated in the present book have a highly characteristic IS frontal-
ventral-transverse cirral pattern strongly indicating that they are a monophylum. How-
ever, the sole apomorphy of the last common ancestor of all species revised is a different
character, namely the fragmentation of a dorsal kinety. This feature, which can be clearly
seen only during morphogenesis, was recognised rather late because it needs silver im-
pregnation. It is so curious that convergent evolution is very unlikely.
From 1758 to 1997 about 440 species were described in oxytrichid genera, however,
only 169 species are considered as valid in the present book. Five species were discov-
ered by o. F. MOLLER in the second half of the 18th century, 39 species were described in
the 19th century, and the majority was found after the turn of the century. The most pro-
ductive and prominent workers on oxytrichid taxonomy are, in chronological order,
EHRENBERG, STEIN, STOKES, KAHL, and ForssNER; however, many others provided important
contributions to the alpha-taxonomy of the oxytrichids. 82 species are synonyms, the syn-
onymy rate is thus about 4S %. Nine species - all of them are very common and known
for a long time - have more than two synonyms. However, 81 % of the valid species have
no synonym. More than 140 species cannot be identified from the original description,
that is, they are species indeterminata. 32 genera are considered as valid; 12 of them are
monotypic, five contain 10 or more species each. These comprehensive genera include
60 % of all species. Oxytricha, the most voluminous group, comprises 54 species; how-
x

ever, only a part of them is described by modem methods, making the identification of
these "inconspicuous" species rather difficult.
This revision was my "leisure pursuit" for many hundreds of hours distributed over a
period of 13 years. Originally it was planned as volume 6 (Hypotrichia) of the Protozoen-
fauna (Gustav Fischer Verlag), a detailed monograph of all freshwater protozoa. How-
ever, only few groups (Suctoria, Urceolaridae, Nackte Rhizopoda, Heliozoea, Colpodea)
were published before this series was suspended in 1996. Fortunately, Prof. Dr. H. J. Du-
mont (Ghent, series editor of the Monographiae Biologicae) and Kluwer Academic Pub-
lishers were interested in my manuscript.
Now, I hope that this revision, whatever its shortcomings, is of more or less great
help to all who deal with oxytrichids. Hopefully, they will be encouraged to tackle the
many remaining problems, great and small, that is, redescribe poorly defined species, elu-
cidate their morphogenesis, ultrastructure, physiology, molecular biology, ecology, and
their phylogenetic relationships. Furthermore, many species in freshwater, soil, and ma-
rine habitats in Europe and especially in the other continents are waiting to be discovered
and thoroughly described.

Salzburg, January 1999 Helmut Berger

Acknowledgements

I am indebted to several people for helping me prepare this book, in particular my

mentor, Prof Dr Wilhelm FOISSNER (University of Salzburg, Austria) for supplying many
original micrographs and faunistic data and for his help with the literature; to Prof Dr
Christian BARDELE (University of tuebingen, Germany), Dipl BioI Hubert BUTIERER
(Amt der Ober5sterreichischen Landesregierung, Linz, Austria), Dr Ulrich BUITKAMP
(Stawa Lippstadt, Germany), Mr Josef DIECKMANN t (Miinster-Wolbeck, Germany), Dr
KomalKAMRA (University of Delhi, India), Prof Dr Martin SCHLEGEL (University of Leip-
zig, Germany), Dr Oliver SKIBBE (University of Berlin, Germany), Dr Weibo SoNG
(Ocean University of Qingdao, China), Mr Hans-JOrgen Voss (Bottrop, Germany), and
Dr Norbert WILBERT (University of Bonn, Germany) for supplying micrographs and illus-
trations; to the staff at the Salzburg University Library for their patient assistance in locat-
ing the vast literature on hypotrichs; and to MSc Eric STROBL (Salzburg, Austria) for im-
proving the English. I also wish to acknowledge the generosity of the Kluwer Academic
Publishers (especially Ms Angela JONES, Bioscience Division, and Drs Rene Mus, Pub-
lishing Editor) and the Series Editor, Prof Dr Henri J DUMONT (The State University of
Ghent, Belgium), for printing this book.
The figures are either originals or reproductions from the literature of the past 220
years. I give sincere thanks to the following publishers who largely freely granted permis-
sion to use published drawings and photographs: Akademiai Kiad6, Budapest (Symposia
Biologica Hungarica); American Microscopical Society, Lawrence (Transactions of the
American Microscopical Society); Bayerisches Landesamt fUr Wasserwirtschaft, MUn-
chen (Informationsberichte des Bayerischen Landesamtes fUr Wasserwirtschaft); Biolo-
giezentrum des Ober5sterreichlschen Landesmuseums, Linz (Kataloge des 0.0. Landes-
museums; Stapfia); Biopress Limited, Bristol (progress in Protistology); Cambridge Uni-
versity Press, Cambridge (Biological Reviews; The Cell Cycle in Development and Dif-
ferentiation); CNRS Editions, Paris (Archives de Zoologie Experimentale et Generale;
Protistologica; charged); Company of Biologists Ltd., Essex (Journal of Embryology and
experimental Morphology); CSIRO Publishing, Collingwood (Invertebrate Taxonomy);
Debrecen Agricultural University, Debrecen (Debreceni Agrarrudomanyi Egyetem Tu-
domanyos K5zlemenyei); Duncker & Humblot GmbH, Berlin (Zoologische Beitrage);
Editions Elsevier, Paris (Annales des Sciences Naturelles - Zoologie); Erik Mauch
Verlag, Dinkelscherben (Lauterbornia); Gustav Fischer Verlag, Jena, Stuttgart (Archlv
fUr Protistenkunde; European Journal of Protistology; Limnologica; Zoologischer Anzei-
ger; Zoologisches Jahrbuch Systematik); Institut fUr landwirtschaftliche Zoologie und Bi-
enenkunde, Bonn (Arbeiten aus dem Institut fUr landwirtschaftliche Zoologie und Bie-
nenkunde); Instituto de Biologia UNAM, Mexico (Manual de ciliados psamofilos mari-
nos y salobres de Mexico; Anales del Instituto de biologia. Universidad de Mexico); Na-
turhistorischer Verein der Rheinlande und Westfalens, Bonn (Decheniana); Oxford Uni-
versity Press, Tokyo (Journal of Electron Microscopy); Polish Academy of Science,
xii

Nencki Institute of Experimental Biology, Warsaw (Acta Protozoologica); Society of

Protozoologists, Lawrence (Guide to the Protozoa; The Journal of Protozoology);
Springer-Verlag, Heidelberg, Wien (Biology and Fertility of Soils; Chromosoma; Oeco-
logia; Polar Biology; Results and Problems in Cell Differentiation; Sitzungsberichte der
Osterreichischen Akademie der Wissenschaften); Taylor & Francis Ltd., London (The
Journal of Natural History); The Norwegian Academy of Science and Letters, Oslo (Zoo-
logica Scripta); The Korean Society of Systematic Zoology, Jeonji (The Korean Journal
of Systematic Zoology); The Zoological Society of Korea, Seoul (Korean Journal of Zo-
ology); University of St. Petersburg, Zoological Institute, St. Petersburg (Zoosystematica
Rossica); Wiley-VCH, Weinheim (Internationale Revue der gesamten Hydrobiologie);
Wissenschaftliche Verlagsgesellschaft mbH, Stuttgart (Cytobiologie). I also ask under-
standing from those publishers whose permission was not obtained due to my oversight.
Specific acknowledgements are made in the figure legends: there are named the authors
of the papers in which the illustrations originally appeared.
Finally, a special acknowledgement should be made to my wife, Elisabeth, for her
understanding of this time-consuming and expensive pursuit, and to my daughters, Mag-
dalena, Eva and Helena, for giving up time that belonged to them.
A General Section

1 Morphology, Biology, and Terminology

In the following chapters the general morphology of oxytrichid hypotrichs and tenns spe-
cific to this group are described and explained. For explanation of other tenns, see
CORLISS (1979), CoRLISS & LaM (1985), and HAUSMANN & HOLSMANN (1996).

1·1 Size and Shape

The length of oxytrichid ciliates ranges from about 40--60 Jim (for example, Oxytricha
setigera) to 30~00 Jim (for example, Conicu/ostomum monilata, Sty/onychia mytilus
complex). Most oxytrichids are short to long elliptical in outline, ventrally flattened, and
dorsally distinctly vaulted (for example, Fig. 148k). The cortex is flexible (supple) and
sometimes slightly to distinctly contractile in the Oxytrichinae (for example, Oxytricha,
Tachysoma, Urosomoida) and in most other non-euplotid hypotrichs. Conversely, the
body of the Stylonychinae (for example, Sty/onychia, P/eurotricha) is rigid as in the
euplotids; this is, however, very likely a convergence. Most flexible oxytrichids have a
slender body, that is, width is <40 % of length, usually about 30 %, while width is often
~40 % of length in oxytrichids with a rigid body. Furthermore, tailed species occur only
in flexible oxytrichids, especially in Urosoma, Urosomoida, and Parurosoma. Ancystro-
podium even has a retractable tail-like process. The oral apparatus is in the left anterior
quadrant of the cell; its relative size is usually about 2~0 % of body length in the Oxy-
trichinae and ~40 % in the Stylonychinae. The biomass of oxytrichids ranges from 8 mg
(Oxytricha setigera) to about 1300 mg (P/eurotricha grandis) for 106 individuals.

1·2 Nuclear Apparatus

Oxytrichids have a homomerous, polyploid macronucleus like the vast majority of cili-
ates (RAIKov 1969, 1982). It consists of two nodules in most oxytrichid ciliates (Fig. Ib,
c). Species of Coniculostomum have a single, moniliform macronucleus shaped like a
question mark (Fig. la) and only some species have 4, about 8, or more (10-17 in Gono-
stomum kuehne/Ii) nodules (Fig. Id, e, Table I). Usually, they are arranged slightly left of
cell-median. The nodules of Sty/onychia mytilus and Gastrostyla steinii (and probably
that of other species, too) are bounded by a common macronuclear envelope with a nar-
row connecting isthmus between G1 interphase nuclei (Fig. 2b). The isthmus contains no
chromatin or nucleolar material, but a bundle of 8-9 nm filaments that often extends into
the adjoined macronuclear nodules (WALKER & GOODE 1976).
The macronucleus of hypotrichs contains only low molecular weight (gene-sized)
DNA molecules. They range in size from a few hundred base pairs to about 15 kilobase
2 GENERAL SECTION

Fig. la-e Diversity of nuclear apparatus in oxytrichid ciliates (Original. Macronucleus dotted, micronucleus
black). See Table I for further, very rare types. a) Monilifonn macronucleus and several micronuclei; for ex-
ample, in Coniculostomum spp. and Stylonychia noduIinucieata. b) Two ellipsoidal or spherical macronuclear
nodules and 2 or more micronuclei; this is the most common (plesiomorphic, as suggested by in-group com-
parison) type, for example, in Oxytricha granulifera, Stylonychia mytilus. c) Two macronuclear nodules with a
single micronucleus in-between; common, for example, in Tachysoma pellionellum, Oxytricha setigera. d)
Four macronuclear nodules and one to several micronuclei; not very common, for example, in Cyrtokymena
quadrinucleata, Gastrostyla steinii, Oxytricha islandica. e) About 8 macronuclear nodules and few to several
micronuclei; not very common, for example, in Rigidocortex octonucleatus, Gastrostyla muscorum, Urosoma
octonucleata.

pairs with an average of2200 base pairs (PRESCOTI 1992a, SWANTON et at 1980b). There
are about 20 000 different molecules, and each appears to contain only one transcription
unit or gene-encoding region. Each of the 20 000 molecules is present, on average, in
about 100 copies (LAUTH et at 1976). The development of the new macronucleus from
the diploid macronucleus anlage in the exconjugants of Stylonychia mytilus is shown in
Figure 3a. All hypotricbs (including euplotids) have macronuclear replication (or reor-
ganisation) bands where DNA replication is localised (Fig. 2a, b, 3b, lOe; see OUNS &
OUNS 1994). This small, clear band (disc) traverses the macronuclear nodule once before
division, a character that is shared only with the oligotrichs (for example, Halteria

Fig. 2a, b Macronucleus of Gastrostyla steinii (from WALKER & GooDE 1976. a, b, electron micrographs). a) -i
The replication band of a macronuclear nodule, bar = 111m. Arrow indicates the direction of replication band
travel. b) The third (from anterior) macronuclear nodule of an interphase specimen, bar = 211m. Closed arrow
indicates direction of replication band travel, open arrows mark fibrillar bundles within the macronuclear
nodule, arrowhead points to the narrow connecting isthmus. a = typical interphase chromatin, b I, b2 = granu-
lar zone of replication band, c = reorganising chromatin, f = fibrillar zone of replication band.
MORPHOLOGY 3
4 GENERAL SECTION

Table 1 Oxytrichid ciliates with uncommon types of nuclear apparatus. The most common type, namely two
macronuclear nodules and two or more micronuclei, is shown in Figure lb. Ma = macronuclear nodule(s), Mi
= micronucleus

Nuclear apparatus Species

I moniliform Ma (Fig. la) Conieulostomum spp. (Fig. 168a, k), Onychodromus quadricornutus
(Fig. 195b), Stylonychia noduJinucleata (Fig. 156)
2 Maand 1 Mi (Fig. lc) Ancystropodium maupasi (Fig. 198a, g), Cyrtohymena gracilis (Fig.
96f), C. sapropelica (Fig. 96h), Cyrtohymena sp. (Fig. 101h, i), His-
trieulus minimus (Fig. 178b), Oxytricha ba/ladyna (Fig. 33a, d), O.
crassistilata (Fig. 35), O. euglenivora (Fig. 37), O. geleii (Fig. 83a),
O. granulosa (Fig. 40b), O. longicirrata (Fig. 46b), O. opisthamusco-
rum (Fig. 73a, b), O. pseudofurcata (Fig. 53a), O. pseudofosiformis
(Fig. 76), O. setigera (Fig. 55d), O. sphagni (Fig. 59), Stylonychia sty-
lomuscorum (Fig. 167d), Tachysoma bicirratum (Fig. 139h), T. humi-
cola (Fig. 138b, e, 139a, e), T. ovata (I 41 p, q), T. pellionellum (Fig.
136a), Urosomaida perlhensis (Fig. 114a)
3Ma Sterkiella thompsoni (Fig. 182t: k)
4 Ma (Fig. ld) Cyrtohymena quadrinucleata (Fig. 103b), Gastrostyla steinii (Fig.
199a, b), Laurentiella strenua (Fig. 1969), Onychodromus grandis (Fig.
193a), Oxytricha is/andica (Fig. 68a, f), Parastylonychia plouneouri
(Fig. 187), Sterkiella admirabilis (Fig. 182a), S. cavicola (Fig. 181a), S.
quadrinucleatus (Fig. 186a, b), S. terricola (Fig. 184n), Urosoma
acuminata (Fig. 128a, b), Urosomaida dorsiincisura (Fig. 115a, c)
About 8 Ma (Fig. Ie) Laurentiella strenua (Fig. I 96g), Paraurostyla polynuc/eata (Fig.
218a), Parentocirrus hartualis (Fig. 218d), Rigidocortex octonuclea-
tus (Fig. 192a), Gastrostyla muscorum (Fig. 205a), Urosoma octonu-
c1eata (Fig. 129a, f), u. polynucleala (Fig. 130a, f)
10-19 Ma Gonoslomum kuehnelli (Fig. 123a, f), Onychodromus quadricornutus
(Fig. 195b), Pattersoniella vitiphila (Fig. 197a, g)

grandinella; GRAIN 1972, LYNN & CORLISS 1991) and some cyrtophorids and chonotri-
chids (RAn<.ov 1982). Some workers (for example, PETZ & FOISSNER 1992) thus assume a
close relationship ofhypotrichs and oligotrichs, which is also indicated by molecular bio-
logical data, for example, comparison of the 16S-like ribosomal RNA (BAROIN-
TOURANCHEAU et al. 1992, FLEURY et al. 1992, LYNN & SooIN 1988). In other ciliates,
DNA replication occurs simultaneously in all parts of the macronucleus. The mature mac-
ronucleus in hypotrichs, as in other ciliates, divides amitotically; neither a distinct organi-
sation of chromosomes, nor the formation of a spindle is seen (Fig. 3b; MURTI 1976,
PRESCOIT 1983, RAIKov 1969).
Oxytricbids have 1,2, or more micronuclei (Fig. 1). They are small (usually about
3-8 J.lm), diploid, possess high molecular weight DNA organised into chromosomes, di-
vide mitotically, lack nucleoli, demonstrate only limited, if any, RNA synthesis, and un-
dergo meiosis during the sexual phase (conjugation) in the life cycle. The mean ratio of
DNA content of macronucleus and micronuclei is 32 in Stylonychia mytilus (AMMERMANN
 MORPHOLOGY 5

t 4000

c3000
SE

~
c
o
u

~2000
a

1000

H ~ f---i It-------i t - - - - - - - - l l l t - - - - - - - - i
Stage 1 2 3 4 5 6

Fig. 3a Changes in the DNA content during the development of the macronuclear anlage in Stylonychiolem-
nae (from AMMERMANN 1971b and AMMERMANN et aI. 1974). The abscissa represents the age of the anlage,
while the DNA content (arbitrary units; according to STEINBROCK 1986 four thousand units are about 800 pg) is
shown on the ordinate. Stage 1 The macronucleus anlage is only slightly larger than a diploid micronucleus
when the conjugation partners separate from each other (0 h). Stage 2 Some hours later the anlage swells while
the spiralled chromosomes (2n about 280) become visible and arrange at the inner surface of the nucleus.
Stage 3 After 15 hours about one third of the chromosomes start moving to the inside of the nucleus, despiral-
ise and develop into giant chromosomes during intense DNA synthesis. The remaining condensed chromo-
somes disappear and are eliminated into the cytoplasm (chromosome elimination step). Stage 4 A first peak in
the amount of DNA is reached and the giant chromosomes attain their final size after about 35 hours. Stage 5
The giant chromosomes are fragmented, their bands are enclosed into independent vesicles. The DNA content
drops drastically, more than 90 % of the anlagen DNA are eliminated into the cytoplasm (DNA elimination
step). Stage 6 The end of the resulting DNA-poor stage, in which the anlage is filled with thousands of gran-
ules, is indicated by the formation of the first two nucleoli (N). After stretching of the macronuclear anlage,
five pairs of replication bands move successively over the nucleus and duplicate the DNA five times to an
amount corresponding to 60 C (1 C is the DNA content ofa haploid micronucleus in GI). The macronucleus is
then mature, and the exconjugants prepare for their division. N = nucleoli.

1965a, RAIKov 1969). AMMERMANN (1987b) found that different populations of Sty/ony-
chia /emnae possess similar amounts of micronuclear DNA, but that there are differences
of ±30 % between clones of the same popUlation. However, the DNA content of the mi-
cronuclei varies by about 100 % during the lifetime of a clone. In species that have only
one micronucleus, it is usually located between the two macronuclear nodules, for in-
stance, in Tachysoma pellionellum or in Oxytricha setigera (Fig. Ic). Natural arnicronu-
6 GENERAL SECTION

cleate races of oxytrichids

are reported and pedigreed
culture has been carried for
289 generations (DAWSON
1919, LIN & PRESCorr
1985, NG 1986, WOODRUFF
1921).
Further literature on
the nuclear apparatus of
oxytrichids (see also Oxy-
tricha ja/lax, O. nova, Sty-
fonychia mytilus complex,
2 S. femnae, S. pustufata, and
Gastrostyla steinil): ALoN-
SO & JARENO (1974), ALoN-
SO & PEREZ-SILVA (1963a,
1969), AMMERMANN (1964,
1965a, b, 1970, 1971a,
1973, 1979, 1987a), AM-
MERMANN & MUENZ (1982),
AMMERMANN et al. (1974,
1978, 1981, 1989), BAN-
5 CHErn et al. (1978b, 1980a,
b, 1982a), BESSLER & LIPPS
(1976), BIERBAUM et al.
Fig. 3b Amitotical macronuclear division in Stylonychia pustulata (1991), BISHOP (1943),
(after SUMMERS 1935 from RAIKOV 1982). 1 = appearance of replica- BLACKBURN (1990), BLACK-
tion bands. 2 and 3 = their movement through both nodules (arrows). BURN & KARRER (1986),
4 = fusion of nodules. 5 = maximal condensation of macronucleus,
mitosis of micronuclei. 6 = first division of macronucleus. 7 = second
BLACKBURN et al. (1983),
division of the daughter macronucleus. BOSTOCK & PRESCorr
(1972), BOSWELL (1981),
BOSWELL et al. (1982,
1983), BtITLER et al. (1984), BOTsCHLI (1876), CALVO et al. (1986a), CAPLAN (1975, 1976,
1977), CARTINHOUR & HERRICK (1984), CHADHA (1980), CHADHA et al. (1979), CHAKRA-
BORTY (1967), DASS & SAPRA (1965, 1967), DAWSON (1984), DAWSON & HERRICK (1982,
1984a, b), DAWSON et al. (1983, 1984), DILLER (1962), DINI et al. (1975), DOGIEL (1929),
DUTTA et al. (1982), EDER et al. (1993), ELSEVIER et al. (1978), ENRIQUES (1912), ESPOSITO
et al. (1976a, 1978), FANG & CECH (1993a, b, 1994, 1995), FANG et al. (1993), FEDRIANI et
al. (1976a), FLEURY (1983), FRICK (1967), GALL (1984), GAUDE (1981), GAUNITZ et al.
(1992), GIL (1976), GIL et al. (1972), GOROVSKY (1986), GORTZ (1983), GOTTSCHLING &
CECH (1984), GOTTSCHLING & ZAKIAN (1986), GRESLIN et al. (1988, 1989), GUPTA et al.
(1993), GUTIERREZ (1985), GUTIERREZ & PEREZ-SILVA (1981), HARPER & JAHN (1989),
 MORPHOLOGY 7

HAsHIMOTO (1952), HELFrENBEIN (1985), HELFfENBEIN & MOLLER (1988), HELFrENBEIN et

al. (1989), HERRICK (1992, 1994), HERRICK & WESLEY (1978), HERRICK et al. (1985,
1987a, b), HlCKE et al. (1990, 1994, 1995), HOFFMAN & PRESCOTI' (1996, 1997a, b), HOFF-
MAN et al. (1995), HORVATH (1939a, b), HUNfER et al. (1989), HUTNER (1975), IVANIC
(1939a, b), JACOBS & KLoBUTCHER (1996), JAHN (1988, 1991), JAHN et al. (1988), JAREI"lo
(1981a, 1987b, c, 1988, 1992), JARENO et al. (1969a), JERKA-DZIADOSZ & FRANKEL (1970),
JICKEU (1884), KAINE (1982), KAINE & SPEAR (1980, 1982), KANG et al. (1992), KAUL
(1989, 1991a, b), KAUL & SAPRA (1986, 1991), KAUL et al. (1990), KAy (1946), KLITZKE
(1916), KLOBUTCHER (1987), KLOBUTCHER & HERRICK (1997), KLOBUTCHER & JAHN (1991),
KLoBUTCHER & PRESCOTI' (1986), KLOBUTCHER et al. (1984, 1986), KLoETZEL (1969,
1970), KRAUT & LIPPS (1984), LAPORTE & THOMAS (1997, 1998), LAWN (1977), LAWN et
al. (1978), LEWIN (1912), LIEDER & RUTHMANN (1976), LIN & PREScOTI' (1986), LINGNER
et al. (1994, 1995), LIPPS (1975, 1977, 1980, 1985), LIPPS & BESSLER (1977), LIPPS &
EDER (1996), LIPPS & ERHARDT (1981), LIPPS & HANrKE (1975), LIPPS & MORRIS (1977),
LIPPS & STEINBROCK (1978), LIPPS et al. (1974, 1978, 1982, 1985), Lu et al. (1991), Lu-
PORINI & DINI (1975), MAERCKER & LIPPS (1993, 1994a, b), MAERCKER et al. (1997a, b),
MATSUSAKA & K1MuRA (1981), MAUPAS (1889), MEYER & LIPPS (1981), MITcHAM et al.
(1994), MURT! (1972,1973, 1976), NG (1990), NOBILl et al. (1981), NOCK (1981), OKA &
HONJO (1983), OKA & THOMAS (1987), OKA et al. (1980), OLINS et al. (1981), ORIAS & HI-
GASHINAKAGAWA (1990), PELC (1972), PIERI (1965, 1966), PLUTA et al. (1982, 1984), POL-
JANSKY & RAIKov (1976), PREER & PREER (1979), PRESCOTI' (1977, 1984, 1989, 1993,
1994), PRESCOTI' & DuBOIS (1996), PRESCOTI' & GRESLIN (1992), PRESCOTI' & MURTI
(1974), PRESCOTI' et al. (1971, 1979, 1982), PRICE (1993), PRICE & CECH (1987), PRzYBOS
(1986), RAE & SPEAR (1978), RAGHURAMAN & CECH (1989, 1990), RAGHURAMAN et al.
(1989), RAIKov (1989, 1992), RAIKov & .AMMERMANN (1974), RIBAS-APARICIO et al.
(1987), RIccI & BANCHEITI (1981), RICCI et al. (1978, 1979a, 1980a), RIEWE & LIPPS
(1977), RIGGIO et al. (1987a, b), ROTH (1958, 1960), SAPRA &.AMMERMANN (1974), SAPRA
& DASS (1970b, 1971), SAPRA et al. (1976, 1985, 1989), SCHLEGEL (1992), SCHLEGFL et al.
(1989, 1990), SCHULTZE et al. (1994), SEEGMILLER & HERRICK (1998), SEEGMILLER et al.
(1996, 1997), SESHACHAR & PADMAVATHI (1956), SHARMA et al. (1986), Sm (1975), SHIP-
PEN et aI. (1994), SMITH & FElOON (1992, 1993), SMITH et al. (1995), SMITH-SoNNEBORN
(1984), SOKOLOFF (1930), SPEAR (1980), SPEAR & LAUTH (1976), STEINBROCK (1981,
1983), STEINBROCK et al. (1981), STEINmLBER & LIPPS (1986), STOLL et al. (1991, 1993),
SWAGER et al. (1990), SWANTON et al. (1980a, 1982), TCHANG et aI. (1981), TORIOOE et al.
(1993, 1995, 1996), TORRES et al. (1979a, b, 1980b, 1982), VAN HOLDE et al. (1979),
WADA & SPEAR (1980), WALKER (1976a, b), WALKER & GOODE (1973, 1975), WALKER et
al. (1978), WANG et al. (1994), WATANABE et aI. (1986), WEFES & LIPPS (1990), WEGNER
et al. (1989), WEN et al. (1995), WESLEY (1975), WEYER (1930), WILLIAMS & HERRICK
(1991), WILLIAMSON (1994), WILLIS et al. (1993), WITHERSPOON et al. (1997), WITTE et al.
(1995), WOODRUFF (1913), YANO (1979, 1985a, b), YANO & SUHAMA (1990a, b, 1992),
YOUNG (1926), ZAHLER & PRESCOTI' (1988, 1989), ZHANG & PANG (1979), ZHANG et al.
(1982).
8 GENERAL SECTION

1-3 Contractile Vacuole and Cytopyge

Almost all oxytrichids have a single contractile vacuole on the left side, usually in about
mid-body or slightly in front of it. Many species have two conspicuous longitudinal col-
lecting canals that supply the central vacuole (Fig. 6a). Sometimes a small vesicle occurs
in the anterior canal. The excretory pore empties on the dorsal surface, usually above the
vacuole. In species of the Slylonychia mytilus complex and in Histriculus histrio, it is a
longitudinally oriented slit that, in live specimens, is recognisable only during excretion
(Fig. 99m).
The cytopyge of oxytrichids is a little-known organelle which is usually located in
the posterior portion of the cell (for example, Fig. 117b).

1-4 Cytoplasm and Colouring

The cytoplasm is often densely granulated. Usually it is filled with some micrometre-
sized, greasy shining globules and rod- and/or V-shaped yellowish crystals. These crys-
tals are more or less randomly distributed in the cytoplasm of Slylonychia (HAUSMANN
1982); in some species (for example, Urosomoida agilis) they are accumulated in the
posterior portion. The crystals consist of an organic and an inorganic component and are
enclosed in vacuoles (HAUSMANN 1982). The cytoplasm is colourless in most species,
only in few taxa is it yellowish or reddish (for example, Rubrioxytricha haematoplasma;
Table 2). Few species (Oxytricha chlorelligera, Pseudostrombidium planctonticum) are
green due to symbiotic green algae, not to be confused with ingested algae, which are en-
closed in usually distinctly recognisable food vacuoles. The size of the globular food
vacuoles depends largely on the size of the species and the size of the food.

1-5 Cortical Granules

Cortical granules (previously - for example, in BERGER & FOISSNER 1987a - often denoted
as subpellicular granules) occur in many species of the Oxytrichinae and are also com-
mon outside the oxytrichids, suggesting that possession of these structures is a plesiomor-
phy (Fig. 22c, Table 2). Conversely, no rigid oxytrichid has cortical granules, strongly
implying that these were lost in the last common ancestor of the Stylonychinae (Fig. 25a,
b). The ultrastructure of these granules is very poorly investigated. Probably, most of
them belong to the mucocyst-type. Their shape, size, colour, and arrangement are very
important species characters which cannot usually be seen after silver impregnation. Live
observation is thus absolutely necessary for reliable identification of oxytrichids.
 MORPHOLOGY 9

Table 2 Oxytrichidae with cortical granules andlor conspicuous coloured cytoplasm. Note that no Stylonychi-
nae with cortical granules are known. Taxa arranged a1phabeticallyl.2

Species3 Granules

Size Shape Colour Remarks

Allotricha anJ- 0.8-1 11m spherical colourless narrowly spaced in longitudinal

arctica (Fig. 92d) rows
Allotricha about I 11m spherical yellowish usually arranged in loose groups
mol/is (Fig. 91 b) and short rows around cirri and
dorsal cilia
Cyrtohymena aus- 0.4-1 11m spherical lemon arranged in groups, mainly around
tralis (Fig, 97g, h, m) cirri and dorsal cilia
Cyrtohymena 0.5-1.5 11m spherical yellow to mainly arranged around cirri and
citrina (Fig. 96b) orange-yellow dorsal cilia
Cyrtohymena ? ? brownish arranged in conspicuous longi-
gracilis (Fig. 96t) tudinal rows
Cyrtohymena delicate ? ? arranged in short rows
marina (Fig. 96j)
Cyrtohymena 0.5-1.5 11m spherical ruby to usually irregularly arranged,
muscorum pale red sometimes in short rows; granules
(Fig. 9480 d, g, h) also in cytoplasm
Cyrtohymena primi- 0.5-0.8 11m spherical yellow usually arranged in groups of3.-10
cirrata (Fig. 98c, d) along cirri and dorsal cilia
Gastrostyla pulchra 211m oval ? in bands, between marginal cirri
(Fig. 206k, m)
Gastrostyla stena- ? ? ? rosettes at bases of dorsal cilia and
cephala (Fig. 207) some cirri
Gonostomum 111m spherical colourless mucocysts(?), loosely arranged in
affine (Fig. 119c) indistinct longitudinal rows, some-
times absent
Gonostomum 1-1.5 11m rod-shaped colourless loosely arranged; stain red with
kuehnelti methyl green-pyronin, usually
(Fig. 123c, d) ejected under coverslip I
Gonostomum small spherical colourless mucocysts(?), after ejection
strenuum spindle-shaped, c. 1-1.5 11m long;
(Fig. 122b, c) do not impregnate with protargol
Notohymena ant- small spherical yellow to in groups around cirri and dorsal
arctica (Fig. 106c) yellow-green cilia
Notohymena austra- ? ? greenish arranged in small groups
lis (Fig. I07b, 108c) around cirri and dorsal cilia
Notohymena rubes- 1-1.4 11m spherical ruby arranged in small groups around
cens (Fig. l04e, g, i) cirri and dorsal cilia
Oxytricha aerugi- spherical russet and regularly arranged, also in the
nosa (Fig. 31 b) blackish cytoplasm
10 GENERAL SECTION

Table 2 Continued

Species Granules

Size Shape Colour Remarks

Oxytricha about 0.5 Jim spherical orange-yellow arranged around cirri and in
auripunctata disorganised rows on ventral
(Fig. 63b) side, stain with protargol
Oxytricha durham- 0.5 Jim spherical ? linear, oblique groups
iensis (Fig. 186d)
Oxytricha ? rod-shaped colourless? rather regularly arranged in
(aurei (Fig. 39g,j) longitudinal rows
Oxytricha gra- 0.5-1 Jim spherical colourless arranged in longitudinal rows
nulifera (Fig. 65b)
Oxytricha granu- 0.7 Jim spherical yellow-green in longitudinal rows
losa (Fig. 40b)
Oxytricha 2-3 x 0.5-1 Jim rod-shaped colourless distal end with inconspicuous
longigranulosa knob; stain red with methyl green-
(Fig. 71lHf) pyronin, but were not ejected;
arranged in short longitudinal rows
Oxytricha ollowi 2 x 0.5 Jim rod-shaped colourless in many loose rows
(Fig. 73d, t)
Oxytricha many dark: (refractile?) granules in
procera (Fig. 74) cytoplasm, especially in posterior
portion
Oxytricha pseudo- cytoplasm blackish
similis (Fig. 77)
Oxytricha quadri- 0.5-1 Jim spherical colourless arranged in longitudinal rows
cirrata (Fig. 78)
Oxytricha rubri- 1 Jim spherical shining red arranged around cirri and dorsal
puncta (Fig. 79b) cilia; cytoplasm colourless
Oxytricha tenella ? ? ? irregularly arranged, apparently in
(Fig.6Ic) several rows
Oxytricha trun- species indetenninata! rusty granu-
cata (Fig. 222p-t) les and small globules in cytoplasm
Paraurostyla 2 x 1 Jim rod-shaped colourless perpendicular to pellicle, very
granulifera closely spaced
(Fig.216c-e)
Paraurostyla about I Jim spherical greenish-yellow mainly grouped along cirral rows
weissei (Fig. 212i) and dorsal kineties
Parentocirrus hor- 1-2 Jim spherical not mentioned do not stain with methyl green-
lUalis (F ig. 218d, t) (colourless?) pyronin or protargol (according to
text, no special cortical granules;
according to figure legend, con-
spicuous cortical granules present)
 MORPHOLOGY 11

Table 2 Continued

Species Granules
Size Shape Colour Remarks

Parurosoma cytoplasm near cytostome and at

dubium (Fig. 146a) base of tail with grey-brownish
pigments (?) and cytoplasmic
crystals
Rubrioxytricha fer- about 111m spherical? brownish irregularly arranged in longitudinal
ruginea (Fig. 14Sd) rows; cytoplasm homogeneously
rusty brown
Rubrioxytricha 0.5-1.4 11m spherical lemon-yellow, regularly arranged in single rows;
haematoplasma sometimes cytoplasm slightly to distinctly
(Fig. 144d) with a greenish homogeneously reddish
shimmer
Taclrysoma drages- about 111m spherical not mentioned 2-4 granules arranged in irregular
coi (Fig. 141g, h) (colourless?) rows
Taclrysoma granu- about 0.5 11m spherical yellow to irregularly distributed in
liferum (Fig. 140c) orange-yellow loosely arranged groups
Tachysoma ovata 1.2-1.5 11m ellipsoidal not mentioned sparse and irregularly arranged
(Fig. 1411-0) long (colourless?) either singly or in pairs
Te"itricha stram- 3-5 x 0.8-1 11m fusiform to colourless perpendicular to pellicle,
enticola (Fig. 219c-f) rod-shaped very closely spaced
Urosoma l acumin- usually protargol-affine granules
ata (Fig. 128b, e) around cirral bases
Urosoma l cau- about 0.5 11m spherical colourless arranged around cirri and dorsal
data (Fig. 125h) cilia
Urosoma l gigan- <111m spherical colourless arranged in short rows
tea (Fig. 134f)
Urosoma l <111m spherical colourless arranged in rows
karini (Fig. 132b)
Urosoma l polynu- <111m spherical colourless mucocysts?
c1eata (Fig. 130c)
Urosomoida about 0.5 11m spherical colourless to mainly arranged around cirri and
agilis (Fig. IlOh) yellowish, dorsal cilia
rarely reddish
Urosomoida about 0.5 11m spherical yellowish usually reticularly arranged
dorsiincisura
(Fig. Il5d)
Urosomoida granu- about 111m spherical colourless irregular groups composed of
lifera (Fig. III b) up to 30 granules

I Many Urosoma species have numerous, about 2-3 11m sized, ellipsoidal, colourless structures (presumably
mitochondria) close underneath the pellicle.
2 The correct colour of the granules or the cytoplasm can only be seen with well-adjusted bright field illumina-
tion.
3 Further species: Apoamphisiella tihanyiensis (Fig. 198.ld); Hemigastrostyla enigmatica (Fig. 246c).
 12 GENERAL SECTION

Fig. 4 Two examples of tracks of cells of Sty/onychia /emnae that sit and feed (numbers), then jump back-
ward by about one body-length, leap forward and turn to the right. Cells often crawl ahead after the double
jump, before coming to a halt. The double jump (back-and-forth movement) is the most common motor activ-
ity of Sty/onychia and other hypotrichs. It does not require external stimulation (it is not "avoidance") and is
not based on "ciliary reversal", although presumably homologous to similar responses in other ciliates (from
MACHEMER 1965b and MACHEMER & DEITMER 1987).

1-6 Movement

Most oxytrichids are thigmotactic, that is, they tend to adhere to the substrate whenever
the opportunity arises (RICCI 1981b, 1990). They creep on their flattened ventral side by
means of the cirri (Fig. 148g, 15Ic). Usually the cells move hastily to and fro; some spe-
cies remain immobile for some time during feeding. Several species are recorded from

Fig. 5 Typical one-minute tracks of Stylonychia /emnae in an unstirred environment (drop size 11-14 mm, ~
0.7 mm high; moist chamber; constant cold illumination at 300 Ix; room temperature; from MACHEMER & 0Err-
MER 1987). The age of the individual cell is given at the starting point (open circle); arrows indicate direction

of locomotion. a) Development of locomotive patterns in a young cell lead to clockwise walks and back-and-
forth movements. b) The same cell proceeded to perform counter-clockwise runs and crawls. c) Regular left-
ward running and back-and-forth movement activity of a 24 h old cell; back-and-forth movements occur at the
edges of the drop. d) A 14 hour-cell speeding along fairly straight tracks. e) Fast curved running of an old cell.
f) Old cell performing rightward crawls, curvy runs, and sequences of forward and backward swimming (undu-
lating tracks); these indicate a deterioration of living conditions. The motor activity was reduced to uncoordi-
nated slow crawls 5 min later.
 MORPHOLOGY 13

(, "0'

~ .-0""
1"'10'

b
Sa

e
14 GENERAL SECTION

the pelagic zone of stagnant and running waters, only Pseudostrombidium p/anctonticum
is, however, euplanktonic. It swims under rotation about the main body axis, always mov-
ing up and down. Conversely, the little known Ancystropodium maupasi (Fig. 198d) at-
taches to the substrate by means of the transverse cirri at the end of the retractable tail-
like process during food consumption; according to FAURE-FREMIET (1909) it can even
make stepping movements.
Species of the Sty/onychia mytilus complex and "Oxytricha hi/aria" have been sub-
ject to detailed studies on their locomotive behaviour and ciliary functions (see last para-
graph for literature). According to MACHEMER (1974), the most common mode oflocomo-
tion is walking on solid substrates. It is mainly performed by the frontal-ventral-
transverse cirri moving slowly rearward in a stiff swing (power stroke) and returning rap-
idly, bent in the opposite direction (recovery stroke; SLEIGH 1968). Walking in any direc-
tion (forward, backward, straight or in arcs) at varying speeds (0.1-2.5 mm S·I; GARNHAM
1966, MACHEMER 1965b) is strikingly smooth, apparently because it is assisted (i) by the
marginal cirri and (ii) by the adoral zone of membranelles. The function of the three long
caudal cirri in species of the Stylonychia mytilus complex is probably to stabilise locomo-
tion and to assist in quick turns. Swimming in both forward and backward directions is
elicited by stimulation, but also appears to occur spontaneously. The common swimming
velocity (1 mm S·I) is less than that of walking, since the cell moves in low-pitch left-hand
helices with the ventral side pointing toward the axis of translation. During swimming, all
cirri are oriented anteriad or posteriad and perform high frequency, anticlockwise, helical
oscillations. Alternatively, the three large frontal cirri may lash at reduced frequency, but
also with a helical component. The adoral membranelles support swimming through the
continuous production of water currents. Walking and swimming are regularly inter-
rupted by a brief synchronous reversal and synchronous reorientation of the cirri,
whereby the animal is thrown backward by approximately one body length and then
pushed forward in a direction to the right of the initial one (Fig. 4, 5; MACHEMER 1965b,
1974).
For the identification of some oxytrichid taxa, it is advantageous to know if the cell is
flexible or rigid. This can be observed only on freely moving cells, that is, not squeezed
under a coverslip! In the Oxytrichinae (for example, Oxytricha, Tachysoma) and some
taxa of uncertain position (for example, Paraurostyla), the anterior portion of the cell
bends left or right during creeping (Fig. 135p). Conversely, the Stylonychinea (for exam-
ple, Histriculus, Stylonychia, Sterkiella) move like a rotating board.
Further literature on movement and motile behaviour of oxytrichid species: BAM.
FORTH (1962), BARBANERA et al. (1997), BRAMSTEDT (1935), BULLINGTON (1925), DEITMER
et al. (1983, 1984), DRYL (1965), DRYL & NOWAKOWSKA (1966), DRYL & TOTWEN-NowA-
KOWSKA (1972, 1975, 1985), ERRA et al. (1989, 1990, 1996), FAURE-FREMIET (1908), JEN.
NINGS (1899b, 1901), JENNINGS & JAMIESON (1902), LUDWIG (1929), MACHEMER (1965b, c,
1966a, 1969a, b, 1970), MACHEMER & DEITMER (1987), MACHEMER & PEYER (1977,
1982), MACHEMER & SUGINO (1986, 1989), MOGAMI & MACHEMER (1991b), MOGAMI et al.
(1992), PARDucz (1938), PEYER (1976), PEYER & MACHEMER (1978a, 1979), PINTO et al.
 MORPHOLOGY 15

(1991), POTIER (1900), RICCI (1981a, 1987, 1994), RICCI & ERRA (1996), RICCI et al.
(1989c, d, 1992a, b), Russo et al. (1988, 1989a--<:), SERAVIN (1970), SLEIGH (1984), SUG-
INO & MACHEMER (1990), SUGINO et al. (1989), TAMAR (1979), TEUNIS & MACHEMER
(1993), TOTWEN-NoWAKOWSKA & DRYL (1976), VERWORN (1889), WATANABE (1959).

1·7 Somatic Infraciliature and Ultrastructure

The somatic ciliature of the oxytrichids is represented by rows and localised groups of
cirri on the usually flattened ventral surface and few (usually 3-6) rows of more or less
widely spaced, 2-15 Jim long and usually stiff cilia ("bristles") on the vaulted dorsal side
(Fig. 6a, b).
Most oxytrichids (that is, hypotrichs showing fragmentation of a dorsal kinety) have
18 frontal-ventral-transverse cirri arranged in 6 more or less distinct groups, namely 3
frontal cirri, 1 buccal cirrus, 4 frontoventral cirri, 3 postoral ventral cirri, 2 pretransverse
ventral cirri, and 5 transverse cirri. An unequivocal designation of each of these cirri is
possible with the simple but ingenious numbering method by WALLENGREN (1900a; Fig.
6a). The exact position of the cirri is of high systematic importance (see phylogeny sec-
tion). For instance, the frontoventral cirri are arranged almost in line in Urosoma (Fig.
19a), and not V-shaped as in most other taxa (Fig. 6a) and in Gonostomum the postoral
ventral cirri are not localised behind the buccal vertex (Fig. 6a), but right of the undulat-
ing membranes (Fig. 19c).
Gastrostyla, Kerona, Laurentiel/a, Onychodromus, Paraurostyla, Pattersoniella,
and Territricha have a slightly to distinctly increased number of frontal-ventral-transverse
cirri, which is, however, no synapomorphy. Only Laurentiel/a, Onychodromus, and Pat-
tersoniella can be rather certainly assigned to the Stylonychinae, as suggested by their
rigid cortex and by molecular biological data, implying that they evolved from 18-cirri
oxytrichids, and did not branch outside this group as is usually assumed. The systematic
position of Gastrostyla, Paraurostyla, and Territricha is uncertain because evidence for
their branching are still scant. Conversely, the number of ventral and transverse cirri is
usually distinctly reduced in Urosomoida (Fig. 19d).
Frontal cirri: They are arranged at the anterior end of the cell. This group, called
"anterior frontal cirri" by some authors (for example, FOISSNER & ADAM 1983b), invaria-
bly consists of three cirri, namely Ill, 1113, and 11113 in 18-cirri oxytrichids and some
other taxa (Gastrostyla, Territricha). In some species (for example, Pattersoniella), the
number is increased secondarily (Fig. 197f). Usually, the frontal cirri are slightly to dis-
tinctly enlarged as compared with the frontoventral cirri. BORROR (1972b) designated the
leftmost frontal cirrus (cirrus Ill) as buccal and later (BORROR 1979, BORROR & WICKLOW
1983) as paroral cirrus because it originates from the same anlage as the undulating mem-
branes (Fig. 6a, 101, m; see next paragraph for confusing terminology). However, a spe-
cial name for this cirrus seems superfluous because there are also several other cirri, es-
pecially in the opisthe, which originate from the oral primordium (Fig. 23).
 16 GENERAL SECTION

Fig. 68, b Ventral and dorsal infra-

ciliature of oxytrichids. a) Oral ap-
paratus and arrangement of cirri on
the ventral side of an oxytrichid
(Sterkie/fa cavico/a) with 18 frontal-
-ventral-transverse cirri (after protar-
gol impregnation). Only the basal
bodies (and not the cilia) of the cirri
are shown. Each frontal-ventral-
transverse cirrus is designated ac-
cording to the ingenious numbering
system by WALLENGREN (19OOa),
a where the frontal-ventral-transverse
streaks are numbered I-VI from left
to right and the cirri are numbered
1-4 beginning from the rear. Cirri which originate from the same streak during morphogenesis are connected
by a broken line. The dotted lines connect cirri which form more or less typical groups (for example, transverse
cirri). Arrowhead marks cytostome at proximal end of adoral zone of membranelles (buccal vertex), arrow
marks distal end of adoral zone of membranelles. b) Dorsal infraciliature and nuclear apparatus of Sterkiella
histriomuscorum. Arrow marks distal end of adoral zone of membrane lies. Fine structure of cirri and dorsal
dikinetids, see Figures 7a-g. AZM = adoral zone of membranelies, BC = buccal cirrus (cirrus 11/2), CC = cau-
dal cirri (usually, as here, on posterior end of dorsal kineties I, 2, and 4), CV = contractile vacuole (usually not
recognisable after protargol impregnation; during diastole usually with longitudinal collecting canals), e = en-
doral (e and p are the "undulating membranes"), FC = left (cirrus VI), middle (lU3), and right (lW3) frontal cir-
rus, FVC = frontoventral cirri (cirri lII/2, IV/3, VU3, VV4; the cirri VU3 and VU4 are homologous to the migra-
tory [= frontoterminal) cirri of the urostylids), LMR = row ofleft marginal cirri, Ma = macronuclear nodule,
Mi = micronucleus. p = paroral. PTVC = pretransverse ventral cirri (cirri VI2. VU2). PVC = postoral ventral
 MORPHOLOGY 17

Buccal cirrus: This tenn is used for cirrus lI12, invariably close to the right and of-
ten near the anterior end of the paroral (Fig. 6a). There is great confusion about the tenni-
nology of cirrus 11/2. BORROR (1972b) introduced the tenn buccal cirrus for cirrus Ill, the
leftmost frontal cirrus in my terminology (Fig. 6a). According to him, the "buccal cirrus"
(cirrus Ill) may occur to the immediate right of the paroral, or be located elsewhere near
the anterior end of the ventral surface. This is obviously incorrect, because cirrus III is
invariably located in front of the undulating membranes and never right of the paroral.
Very likely, BORROR (1972b) confused cirri III and II12. Furthennore, he erroneously
labelled cirrus 11/2 as buccal cirrus, and not cirrus Ill, as intended in his schematic Figure
1. Presumably, this misleading labelling caused some taxonomists (BUlTKAMP 1977a, b,
FOISSNER 1982) to use the tenn buccal cirrus for cirrus lIl2, which was not specifically
named by either BORROR (1972b, 1979) or previous authors. Subsequently, the name buc-
cal cirrus was used in the latter sense by many taxonomists. The confusion was further in-
creased by BORROR (1979) in that he introduced the tenn paroral cirrus, a replacement
name for "buccal cirrus" (cirrus Ill) sensu BORROR (1972b). Finally, BORROR & WICKLOW
(1983) introduced the tenn malar cirrus for cirrus lI12. Thus, malar cirrus is a synonym of
my preferred tenn buccal cirrus. In conclusion, I strongly recommend using descriptive
terms concomitantly with the well-established and simple numbering system introduced
by WALLENGREN (1900a).
Frontoventral cirri: This group comprises four cirri (llIl2, IVI3, VI13, V1I4) be-
tween the anterior portion of the right marginal row and the paroral. The V- or hook-
shaped arrangement shown in Figure 6a is widespread. Conversely, in Urosoma cirrus
III12 is in front of the other three cirri, which thus fonn a longitudinal row (Fig. 19a). No
other distinct genus-specific patterns can be distinguished. However, cirrus III12 shows a
tendency to be placed more posteriorly in species having a rigid body, for example, in
Stylonychia mytilus, Steinia spp., and Histriculus histrio (Fig. 19b). Furthennore, the
frontoventral cirri are displaced postorally in Oxytricha setigera and related species. The
frontoventral cirri VII3 and V1I4, which never fonn primordia during morphogenesis, are
homologous to the migratory cirri (BORROR & WICKLOW 1983; = frontotenninal cirri of
HEMBERGER 1985) of the urostylids (WIRNSBERGER et al. 1985a).
Postoral ventral cirri: This tenn is commonly used for the cirri IVI2, V/3, and V/4,
which are behind the proximal end of the adoral zone (Fig. 6a). These three cirri fonn a
narrow group close to the buccal vertex in Oxytricha, Allotricha, Urosomoida, Urosoma,
Cyrtohymena, Notohymena, and Tachysoma (for example, Fig. 69c). By contrast, cirrus
V13 (posterior postoral ventral cirrus) is distinctly separate from the other two cirri in
some taxa with a rigid body, for example, Sterkiella and Steinia (Fig. lOa). However,
transitions exist (for example, Australocirrus oscitans; Fig. 142d). In Gonostomum, the
postoral ventral cirri are displaced right of the proximal portion of the adoral zone of

~ cirri (cirri IVI2, V/3, V/4), RMR = row of right marginal cirri, TC = transverse cirri (cirri IIII, 1lII1, lVII, VII,
VIII), I-VI = frontal-ventral-transverse cirral streaks (primordia, anlagen), 1-4 (in Fig. 6a) = cirri within a
streak, 1-6 (in Fig. 6b) = dorsal kinetics 1-6 (kinetics 5 and 6 are dorsomarginal kinetics).
18 GENERAL SECTION

membranelIes, as proved by morphogenetic data (Fig. 19c; HEMBERGER 1982, SoNG

1990).
Pretransverse ventral cirri: This term was introduced for ventral cirri V/2 and
VII2, which are arranged in front of the transverse cirri (Fig. 6a; BERGER & FOISSNER
1997). Some taxa (Urosomoida, Urosoma po/ynucleata, U simi/is) have lost one or both
of these cirri (Fig. 19d).
Transverse cirri: This group is in the posterior quarter of the cell and usually com-
prises five cirri, namely 1111, lIlli, NIl, VII, and VIII (Fig. 6a). Some taxa have more
than 5 (for example, Onychodromus quadricornutus, Territricha, Paraurostyla), some
less than 4 (Urosomoida spp.). The loss of cirrus 1111 undoubtedly occurred independ-
ently in different taxa because species with four transverse cirri are found in different
genera. Especially in the Stylonychinae the transverse cirri are often distinctly enlarged as
compared with the pretransverse ventral cirri, for example, in Histriculus histrio (Fig.
176b) and/or separated into two distinct groups, for instance in Sty/onychia mytilus,
Coniculostomum spp., andSteinia sphagnicola (Fig. 149a).
Marginal cirri: These cirri run along the left and right body margin. Most oxytri-
chids have one left and one right row (Fig. 6a). An increased number of rows obviously
evolved independently, namely in genera with a flexible (Allotricha, Parurosoma; Fig.
91c, 146a) and those with a rigid body (Conicu/ostomum, Pleurotricha; Fig. 168e, 18ge).
Usually, the marginal rows are more or less distinctly separate posteriorly, that is, not
confluent; the gap is often difficult to recognise because it is seemingly occupied by the
caudal cirri, which, however, insert on the dorsal surface (Fig. 6b). Only in Histriculus
(and possibly in Pleurotricha) are the marginal rows in fact confluent (Fig. 22d).
A cirrus or polykinetid is a tuft of cilia, few to more than a hundred in number, func-
tioning as a single unit, although with no special or additional enveloping membrane, and
tapering or fimbriate distally. It is principally a locomotion organelle and typical of hypo-
trich ciliates, although it is not exclusively found there (CORLISS & LoM 1985).
The ultrastructure of a cirral base is rather complicated (GRIM 1972, MATSUSAKA et a1.
1984, PUYTORAC et a1. 1976; Fig. 7a, b, d-g). The base is composed of few to many basal
bodies (for example, less than 20 in a marginal and more than 50 in a frontal cirrus of
Sterkiella cavicola; MATSUSAKA et a1. 1984) arranged in a more or less modified paral-
lelogram. Each basal body is linked in a complex manner by (usually) 10 electron-dense
fibres with the other basal bodies and/or the electron-dense, cup-like peripheral sheet
("rampart"; PuYTORAC et a1. 1976) comprising the network structure. Three bundles of
microtubules are attached to the periphery of the rampart to provide (among others?) me-
chanical anchorage (Fig. 7e): (i) an anterior longitudinal fibre originating from the left
anterior end, (li) a transverse rootlet from the right margin, and (iii) a subectoplasmic
rootlet from the right posterior corner. The basal bodies of a cirrus are arranged in
oblique files. Postciliary microtubules originate just behind the posterior basal bodies and
transverse microtubular ribbons are attached to the anterior basal bodies of each file.
Usually there is a kinetodesmal fibril associated with the posterior basal body of each file
(Fig. 7b). Bundles of microtubules (neodesmal microtubules) were observed in some cirri
 ~
o
zg
o==
5o
-<

AL-

Fig. 7a, b Fine structure of a left marginal cirrus of Gastrostyla steinii (from GRIM 1972; figures are schematic drawings
based on serial thin sections). a) Diagrammatic sketch of3 left marginal cirri and associated fibres. The AL-fibres point anteri-
orly, the TR-fibres to the median of the cell. b) Semi-diagrammatic, 3-dimensional view of an abbreviated base of a left mar-
ginal cirrus with fibrils and microtubules attached. For correct orientation of cirrus, see Figure 7al To avoid confusion, the mi-
crotubules of the AL-fibre which terminate in the rampart have been omitted, as have many microtubules of the SSR-fibre. AL
= anterior longitudinal fibre (consists oftmt and microtubules which terminate in the rampart), Kd'" kinetodesmal fibril, nw =
network of fibrils connecting basal bodies ofa cirrus, pcmt ... postciliary microtubular ribbon, ra'" rampart (amorphous sheet),
a SSR = small sub-ectoplasmic rootlets, tmt =transverse microtubular ribbon, TR - transverse rootlet (terminates in rampart).
\C
-
20 GENERAL SECTION

Fig. 7c Dorsal dikinetid of Oxytricha sp. (after electron microscope micro-

graphs in GRIMES & ADLER 1976 from LYNN 1981, 1991). The "inter-kineto-
somal connective" (some kind of fibrillar material) is dotted. Kd = kineto-
desmal fibril (dashed; present only in primordial bristle units), perot = post-
ciliary microtubular ribbon associated to triplet 9 of posterior basal body, tmt
= transverse microtubular ribbon associated to anterior basal body.

of Sterlde/la cavico/a originating from the proximal end of the

basal bodies (Fig. 7g). Parasomal sacs are lacking (LYNN
1981).
The ciliature of the dorsal side consists of rows of cilia
and zero (for example, Tachysoma) to several (for example,
13 in Gastrosty/a dorsicirrata) caudal cirri at the posterior
end of defined dorsal kineties (Fig. 6b). Most oxytrichids have
5 or 6 kineties. Almost all oxytrichid taxa show a peculiarity
in the dorsal pattern, namely fragmentation of one or more
dorsal kineties. Lack of fragmentation in Gonostomum, Uro-
soma, and Urosomoida evolved secondarily, as suggested by
cladistic analysis.
Dorsal kineties consist of basal body pairs. The anterior basal body bears a 2-15 Ilm
(usually 3-6 Ilm) long, stiff cilium, frequently named bristle (GRIMES & ADLER 1976).
The posterior one has a ciliary stub with 9 doublet microtubules, but lacks the central
pair. Each basal body pair, located at the bottom of a cortical pit, is surrounded by an
asymmetrical fibrillar mass, and the transverse microtubular ribbon is associated with the
anterior basal body, the postciliary microtubular ribbon with the posterior basal body
(Fig. 7c). The kinetodesmal fibril is present only in primordial bristle units (GRIMES &
ADLER 1976). The mature dorsal dikinetid of Sty/onychia mytilus differs from that of S.
pustu/ata and Oxytricha sp. in that the posterior basal body is reduced after fISsion to a
prominent fibrous structure, the parakinetosomal body (GoRTZ 1982). Possibly, this is an
autapomorphy of species of the Sty/onychia mytilus complex; however, data on closely
related taxa, like Conicu/ostomum and Steinia, are needed to evaluate this feature
reliably. Like the posterior basal body in the other species, it may serve as an anchor for
the bristle cilium. The function of the dorsal cilia is unknown. GELEI (1929, 1933) consid-
ered them to be sensory organelles. GORTZ (1982) observed that the dorsal cilia of Sty/o-
nychia mytilus may beat in non-streaming media, and that they may act individually.
However, at a current velocity of 250-300 Ilm S·I, the cilia bend with the direction of the
flow, indicating that they may in fact be sensory, responding to stimuli from water cur-
rent (GORTz 1982). Another possibility is that the dorsal ciliature is merely a rudiment
without any function.
The caudal cirri are at the posterior margin of the dorsal surface (see character 23 in
the phylogeny section). The absence of caudal cirri evolved independently in the Oxytri-
chinae (Tachysoma) and Stylonychinae (Histriculus). The relative length of the caudal
cirri was used by KAHL (1932) as characteristic for Opisthotricha and Stylonychia. How-
 MORPHOLOGY 21

Fig. 7d, e Arrangement of fibres on left and right marginal cirri (d) and cross section of a right marginal cirrus
(e) of Sty/onychia mytilus (from PUYTORAC et aI. 1976). The cross section is somewhat oblique (proximal on
the right and distal on the left side) to show the structure at different levels of the cirrus which consists of 4
oblique rows of 6-7 basal bodies. The anterior basal body of each row is attached to a transverse microtubular
ribbon (tmt), the posterior one is accompanied by a postciliary microtubular ribbon (pcmt). AL = anterior lon-
gitudinal fibre (consists oftmt and microtubules which terminate in the rampart), Kd = kinetodesmal fibril, M
= median, nw = network of fibrils connecting basal bodies of a cirrus, pcmt = postciliary microtubular ribbon,
ra = rampart (amorphous sheet), SSR = small sub-ectoplasmic rootlets (terminate in rampart), tmt = transverse
microtubular ribbon, TR = transverse rootlet (terminates in rampart).
 22 GENERAL SECTION

Fig. 7f Cross-section ofa cirral base of Slerkiella cavicola (from MATSUSAKA et al. 1984. Transmission elec-
tron microscopy micrograph. Anterior part of cirrus towards top. Bar = 0.5 11m). There are 5 files (arrowheads)
of basal bodies, each of which has a transverse ribbon at its anterior and a postciliary ribbon at its posterior.
 MORPHOLOGY 23

Fig.7g Sagittal section of a cirral base of Sterkiella cavicola (from MATSUSAKA et al. 1984. TEM-micrograph.
Bar = 0.5 11m). Arrow marks axosomal plate, double arrows denote tenninal plate, arrowheads mark wedge-
shaped bridge. Explanation of original labelling: as = axosomes, nmt = nematodesmal microtubuli, nw = net-
work structure, pemt = postciliary microtubular ribbon, ra = rampart, tmt = transverse microtubular ribbon (in-
dicating anterior part of cirrus).

ever, at the present stage of knowledge, the length is of significance for species distinc-
tion only. The caudal cirri appear to be structurally identical to the marginals (GRIMES &
ADLER 1976), indicating that marginal and dorsal rows are homonomous (BERGER et al.
1985a). The caudal cirri have about the same length as the marginal cirri in some species
and are thus difficult to discern. However, they are very long and conspicuous in some
other species, for example, Stylonychia mytilus (Fig. 148a). Unfortunately, the exact dor-
sal infraciliature and its morphogenesis, which are very important phylogenetic markers,
are known only for a small number of species.
The plasma membrane of some (all?) non-euplotid hypotrichs (Urostyla grandis,
Pseudokeronopsis rubra, Pseudourostyla cristata, Uroleptus caudatus, Paraurostyla
weissei, Oxytricha fallax, Stylonychia mytilus, Urosoma sp.) and some oligotrichs (for
example, Strombidium) is covered by an additional layer called perilemma (BARDELE
1981, GRIMES 1972, LAVAL 1971, LAVAL-PEUTO 1975). This outer coating also covers
cilia, membranelles, and cirri. The perilemma is lacking in the oligotrich Halteria and the
euplotid hypotrichs. BARDELE (1981) assumed that the perilemma in hypotrichs is a tem-
porary structure which is discarded quite often, because numerous layers of the peri-
lemma are usually seen in the buccal cavity. The cyst wall is formed between the peri-
lemma and the plasma membrane (GRIMES 1973d). Accordingly, the perilemmapresuma-
bly functions as an envelope to keep the precursers from dispersing after deposition in the
precystic cell surface. Unfortunately, nothing is known as to how the perilemma is de-
rived or replenished. LYNN & CORLISS (1991) supposed that it may be a special kind of

~ Note the single postciliary microtubules (arrows) associated with the inner basal bodies, which are linked by
many connectives and surrounded by dense material. bb = basal bodies, nw = network structure, pemt = post-
ciliary microtubular ribbon, ra = rampart, tmt = transverse microtubular ribbon.
24 GENERAL SECTION

fixation artefact of the glycocalyx, that is, the protein and glycoprotein layer of the
plasma membrane.
Further literature on the ultrastructure of the cortex: CALVO et al. (1984, 1986a),
CHAKRABORTY (1967), ESTEVE (1972), FLEURY (1988), FLEURY et al. (1993), GRIM (1970),
JERKA-DzIADOSZ (1980, 1982), JERKA-DZIADOSZ & WIERNICKA (1992), KAUL et al. (1978b),
PuYrORAC & RODRIGUES DE SANTA ROSA (1976), ROSATI et al. (1988), TORRES et al. (1986),
WILBERT & HEuER (1971).

1-8 Oral Apparatus

The adoral zone of membranelles, the most prominent part of the oral apparatus, extends
near the left body margin from the anterior end to about mid-body. Usually, it is roughly
the shape of a question mark. Right of the zone there are two undulating membranes bor-
dering a more or less wide and deep buccal cavity (Fig. 6a). The shape of the adoral zone
of membranelles and especially the shape and arrangement of the undulating membranes
are important for the evaluation of the phylogenetic relationships within the oxytrichids.
Thus, a detailed description of the different patterns evolved within this group is given in
the phylogeny section (characters 1-3; Fig. 19e-j, 20a-f, 21a-f), and the discussion here
limited to some general remarks.
The terms paroral and endoral membranes were introduced by STERK! (1878). How-
ever, he used the first to describe the rightmost cilia of the hypotrich adoral
membranelles, which often beat independently of the remainder. But for some decades
the term paroral membrane has been rather uniformly used in a wide variety of ciliates for
the right portion of the oral ciliature (EISLER 1992). I follow LYNN (1988) in that I use pa-
roral and endoral as nouns (BERGER & FOISSNER 1997). Synonyms of the hypotrich paroral
are undulating membrane (used, for example, in BORROR 1972a, FOISSNER et al. 1991,
KAHL 1932), paroral formation 2 (FERNANDEZ-LEBORANS 1985), external paroral kinety
(FERNANDEZ-LEBORANS 1985), and outer preoral membranelle (JERKA-DZIADOSZ 1981c).
Synonyms of the endoral are paroral formation 1 (FERNANDEZ-LEBORANS 1985), internal
paroral kinety (FERNANDEZ-LEBORANS 1985), and inner preoral kinety (JERKA-DZIADOSZ
1981c). In general, the paroral extends at the outer margin of the buccal cavity, that is, on
the cell surface, while the endoral is on the bottom and right border of the cavity. This
means that the membranes extend at clearly different levels; however, if the cell is viewed
from ventral, they appear to lie side by side (for example, in Sty!onychia, Fig. 19i,20d,
148i) or to intersect (for example, in Oxytricha, Fig. 1ge, 20a), depending on their shape
and arrangement.
The buccal cavity is also rather different in shape and size and is usually described
by the terms flat or deep and wide or narrow. Flat means that the buccal field is only
slightly hollowed, whereas a deep cavity extends almost to the dorsal surface, making the
buccal field conspicuously bright. In a wide buccal cavity the paroral, that is, the right
margin of the cavity, is in the midline of the cell, whereas in a narrow buccal cavity the
paroral is displaced to the left, that is, very close to the adoral zone.
 MORPHOLOGY 25

Fig. 8 Adoral membranelle of Paraurostyla weisse; (from BAKOWSKA & JERJ<A-DzIAoosz 1980). The mem-
branelles of oxytrichids, and hypotrichs in general, usually consist of 4 rows (1-4) of basal bodies. Row 3 is
distinctly shortened on left side, row 4 consists offew basal bodies only. Bar = I ~m.

Fig. 9a, b Schematic representa- ~T

tion of a cross-section through the ,, "" T
paroral (a) and the endoral (b) of Pc "
Paraurostyla we issei (from BAKo- -T ,
~
~ T
WSKA & JERKA-DzIADOSZ 1978). PC'
The paroral consists of 4 rows, the
-T ,, -T
endoraI of only one row of basal PC'
" T
bodies. Explanation of original la- ~
~T ,,
bels: C = connections between ba- PC'
sal bodies, Pc = postciliary micro-
,,
, T
tubules, T = transverse microtu-
PC--": PC'
"" T
buies, x-x' = a plane perpendicu- ~

lar to the effective stroke of mem- ,

branellar cilia Pc --- Pc
,,
~
~ T
Pc '-- PC'
, /
""~ T
The fine structure of
Pc -- PC'
the oral apparatus of the
,,
, ~
~
T
oxytrichids is studied in PC'
Pc --- :.
few species only, namely
Gastrostyla stein;; (GRIM PC--- PC' - ,, %T
1972), Histriculus similis
(CALVO et al. 1986a), Lau- PC'
, ,
-T
rentiella acuminata (TOR-
Pe--_
x-- aCfJ
'0 co
'b
'l> X a
<9"''b
00
X'
1\- - -
RES et al. 1986), Oxytricha '" OOrP a---_X' - - - Il
", 00
6'

fallax (GRIMES 1972),

a b
Paraurostyla hymenophora
(GRIMES & L'HERNAULT
1978), Paraurostyla weissei (BAKOWSKA & JERKA-DZlAOOSZ 1980, JERKA-DZlAOOSZ 1981b,
c), Sty/onychia mytilus (NOBILl & ROSATI RAFFAELLI 1971, PUYTORAC et al. 1976). A very
detailed description is given by BAKOWSKA & JERKA-DZlAOOSZ (1978) for Paraurostyla
weissei (Fig. 8, 9). The membrane lIes of the adoral zone are composed of four rows of
basal bodies. The first and second row in proximal membrane lIes are of equal length and
the longest, the third row is slightly shorter, the fourth is the shortest, consisting of only
26 GENERAL SECTION

few basal bodies (for example, 3 in Paraurostyla weissel). The paroral of P. weissei is
composed of 4-5 rows of basal bodies arranged longitudinally, whereas the endoral is
composed of a single longitudinal row of basal bodies, all of which are ciliated (Fig. 9;
BAKOWSKA & JERKA-DZIADOSZ 1978).
JERKA-DzIADOSZ (1981b) noted that the ultrastructure of the undulating membranes of
Paraurostyla weissei and Gastrosty/a steinii is more similar to members of the Urostyli-
dae, such as Pseudourosty/a cristata and Pseudokeronopsis rubra, than to Oxytricha or
Sty/onychia. This implies that Paraurosty/a and Gastrosty/a could have branched out
from the 18-cirri oxytrichids, that is, fragmentation of dorsal kinety 3 evolved before the
18 frontal-ventral-transverse cirral pattern. However, further data are needed to confirm
this hypothesis.

1-9 Silverline System

The whole silverline system of oxytrichids consists of small (1-2 Jim) polygonal meshes
(Fig. 65e). In contrast to the euplotids it has no taxonomic value in oxytrichids.

1-10 Life Cycle

All oxytrichids have a normal life cycle, that is, the theronts feed, become trophonts, di-
vide, encyst or conjugate. Usually, there are no conspicuous morphological differences
between theronts and trophonts. Conjugation is followed by a period of sexual
immaturity. A sexually immature cell, though apparently quite normal from both the mor-
phological and physiological points of view, divides vegetatively with a regular cell
cycle, but cannot conjugate. Full maturity is preceded by a short period of adolescence; in
this period the individuals can conjugate weakly with specimens of complementary mat-
ing types. In Oxytricha the period of immaturity lasts from one month to two years
(SIEGEL 1956), in Sty/onychia mytilus usually 3--6 months (AMMERMANN 1965a), and in
"Oxytricha bi/aria" about 170 binary divisions (RICCI 1981c). Once it becomes sexually
mature, Oxytricha sp. maintains its maturity for at least two years (SIEGEL 1956). Finally,
the clones become "old", and their ability to conjugate decreases. After a certain point, a
clone can only reproduce vegetatively and will eventually die (RICCI 1981c).

Fig. lOa-g Morphogenesis of celI division in Sterkiella histriomuscorum after protargol impregnation (from -
BERGER et al. 1985a a, c-g, ventral infraciliature; b, dorsal infraciliature). a, b) Inftaciliature and nuclear appa-
ratus of a non-dividing specimen, 65 11m. The cirri originating from one streak are connected by a broken line
(see Fig. 6a for detailed designation of cirri). c, d) Left posterior part of very early stages, bar 20 )!m. The oral
primordium originates immediately left of the anterior-most transverse cirri (arrows mark transverse cirrus
IIVI). e) Very early stage, 85 )!m. The oral primordium is long and narrow. Arrows mark replication bands of
macronuclear nodules (see chapter 1.2). f) Early stage, 70 11m. The buccal cirrus IJ12 (arrow) and the postoral
ventral cirrus V14 (arrowhead) are modified to primordia. g) Early stage, 65 )!m. The frontoventral cirri IIJ12
 MORPHOLOGY 27

'. ·1:~.-,
Mi
. I~'-Ma

---I

Q t
-0:' ""
00
01)
b d

00 00
/'
0
cJ
,,'
t ·7' o /
1/ ~
0 t1
C' . ~

r'l'
I?f ~
•
c1 tf c1 f~
~\ t·\ ~ If
~. t ~.\ ~
e :.: ~

cJ ~ , '.'~o
' / 0
.~
"'~ ..

/"-oJ '
.~\~... t1 ' ".:.,. ¢
00 ~

""
~
c1 ~'~Yo- (I
f .~ ~ \,~ ~*- cJ
il·

,
c1 \.~ !M~{':,
0 c? cJ
C!
C! cJ
~fi
~1~~
,t::'"
c1
c1 " c::J
~itJ
i.~~/
cJ

!!:l

I
c1 c1
~

Ii/ ~:
i:"
t1
,;:> tf

cJ 0 d
cJ
0 '"
¢
<7 oc? cT 0
C
"0
00 '"
a
'" e
cT
f g

(arrow) and IV/3 (arrowhead) also commence with modification to primordia The right anterior part of the
oral primordium differentiates to membranelles. AZM = adoral zone of membranelles, CC = caudal cirri at end
of dorsal kineties 1,2,4, Ma = macronuclear nodule, Mi = micronucleus, OP = oral primordium, I-VI = cirral
streaks I-VI, 1-6 = dorsal kineties 1-6.
28 GENERAL SECTION

CURDS (l965b, c, 1966a) and CHARDEZ (1986) described endogenous bud formation
in Histricu/us spp. (Fig. 178j, m-s, w, 179c-f, i-n, 180); however, they very probably
observed cannibalism induced by overfeeding (PANG & ZHANG 1981). Exogenous bud
formation was described by GREENAWAY (1916) in Sty/onychia sp. (Fig. 240k), but pre-
sumably the content of a food vacuole (a small ciliate) was ejected.

1-10-1 Cell Division

Oxytrichids reproduce by isotomic transverse fission, like many other ciliates (Fig_
1Oa-v; FOISSNER 1996d). The anterior filial product is the proter, the posterior the opisthe.
In early stages of division, a replication band traverses each macronuclear nodule (Fig.
1Oe; see chapter 1.2). The nodules fuse during early and middle stages of morphogenesis
(Fig. 100). The macronucleus divides amitotically just before cytokinesis, whereas the
micronuclei(eus) divide(s) mitotically (Fig. lOp, r).
The changes of the ventral and dorsal ciliature during cell division are known from a
considerable number of species (Tables 3, 4). With the aim of morphogenetic data it is
possible to homologise the cirri of some deviating taxa For instance, the pronounced
postoral ventral cirrus of Gastrostyla (Fig. 20Ia-h) can be unequivocally homologised
with the left postoral ventral cirrus (lV/2) of the 18-(frontal-ventral-transverse)cirri oxy-
trichids (Fig. 6a).
The parental ventral and dorsal somatic ciliature is usually completely renewed dur-
ing morphogenesis; in Conicu/ostomum, however, some parental cirri and dorsal kineties
are not resorbed after division and are thus part of the "new" ciliature. The parental undu-
lating membranes undergo a reorganisation whereby the tightly packed linear array is
lost, later reorganising into the new membranes and the left frontal cirrus (Fig. 101-q;
GRIMES 1972). Together with the unchanged parental adoral zone of membranelles, the
membranes form the oral apparatus of the proter. In contrast, the oral apparatus of the op-
isthe is a new formation. It develops from an oral primordium which, in many species,
originates very close to the left transverse cirrus and/or one or more postoral ventral cirri
(Table 3; for example, Fig. 1Oc, d, 94r). Transmission electron microscopic studies of
very early dividers of Oxytricha proved that the first basal body of the oral primordium is
not originally a part of a cirrus (GRIMES 1972). This suggests that stomatogenesis of oxy-
trichids is apokinetal, that is, the involved basal bodies have no apparent pre-association
with either cirri or the parental oral apparatus; the origin of the basal body anlage appears
to be de novo (CORLISS 1979, CORLISS & LOM 1985). Alignment of basal bodies to form
the membrane lies of the adoral zone first occurs in pairs, the immature membranelies
containing only two rows of basal bodies. The third and fourth row are added by produc-
tion of new basal bodies from right to left on the anterior edge of each membranelle
(GRIMES 1972). Specific characters of the oral apparatus, namely conspicuous shapes of
the undulating membranes, are only formed in the postdividers (Fig. 95p, q, s).
In the 18-cirri oxytrichids, the ventral somatic ciliature originates from 6 roughly lon-
gitudinal streaks (anlagen, primordia) numbered from I-VI according to WALLENGREN
 MORPHOLOGY 29

Cl
c:' c:>

c?

o.

"00 ()/)

{
,0
A

c?
c:J
c:J
,cP
\\\.
\ \ o
\, o co '"
k
o
o
"o r; 0
a ()"
co
m
Fig.lOh-m Cell division in Sterkiella histriomuscorum (from BERGER et a1. 1985a Ventral and dorsal infra-
ciliature, protargol impregnation). h) Early stage, 75 11m. Postoral ventral cirrus IV13 (arrowhead) starts disor-
ganising. Arrow = primordium III. i) Middle stage, 80 11m. Arrows = primordium VI; arrowhead = right mar-
ginal primordium of proter. j, k) Middle stage, 80 11m. Primordia of marginal rows and dorsal kineties 1-3
situated at about same level. I) Middle stage, 70 11m. Arrows = primordium of dorsal kinety 5 (dorsomarginal
kinety). m) Late stage, 70 11m. Segregation of new frontal, ventral, and transverse cirri complete.
 30 GENERAL SECTION

•..
•~
.-.
-. •••
\ \0
o
o

o
o 0 "
00 " .. "
p

Fig. lOn-q Morphogenesis of cell division in Sterkiella histriomuscorum after protargol impregnation (from
BERGER et al. 1985a n, p, q, ventral infraciliature; 0, dorsal infraciliature; parental structures white, new struc-
tures black; cirri originating from one streak are connected by a broken·line). n, 0) Late stage, 70 Jim. Short ar-
rows mark primordium of dorsomarginal kinety 6 of the proter (o) and the opisthe (n); arrowheads denote frag-
mentation of dorsal kinety 3; the posterior fragment becomes the dorsal kinety 4; long arrows mark new caudal
cirri at posterior end of primordia of dorsal kineties I, 2, and 4. The fused macronucleus commences with divi-
 MORPHOLOGY 31

(j • ,
"'.\(.~
\ ~f ~'I
~,.If( ~
\~~ Ii ~ "
I\ ,\ \\'\
\ \ \
If ',, I

\",
\ \ ~
\. t\ ,\~/ ;
\ \
\J?i'~
' ,,\
\ '\

\ l--, \~~

,.
\ \ "~''
" I;,;:
il ... ..

,- "0
.
•
0

r u v
Fig. lOr-v Morphogenesis of cell division in Sterkiella histriomuscorum after protargol impregnation (from
BERGER et aI. 1985a. Parental structures white, new structures black; cirri originating from one streak are con-
nected by a broken line). r) Dorsal ciliature ofa very late stage, 105 ~m. Ventral infraciliature of this stage, see
Figure lOq. S, t) Ventral and dorsal infraciliature of a proter, 50 J.Im. u, v) Ventral and dorsal infraciliature of
an opisthe, 45 J.Im.

(1900a, 1902; Fig. 6a, 10i, j, I, m). These anlagen arise from (i) parental ciliature, (ii)
new, and/or (iii) the oral primordium. Six or less parental frontal-ventral-transverse cirri
participate in primordia fonnation (for details, see Table 3). However, sometimes it is
rather difficult to ascertain exactly where a primordium comes from. The following num-
bers of cirri originate from the streaks I-VI in IS-cirri oxytrichids: 1, 3, 3, 3,4,4. The
following parental cirri are resorbed only in the latest stages of morphogenesis in most
species: frontal cirri 1f1, Ilf3, IIlf3, frontoventral cirri Vlf3 and VII4 ("migratory cirri"),
pretransverse ventral and transverse cirri (Fig. 10m, n, p, q, s, u). Several morphogenetic
peculiarities can be used to elucidate phylogenetic relationships of oxytrichids (BERGER &

~ sion. The primordia of the undulating membranes begin separating into paroral and endoral. p) Late stage,
70 J.Im. Division of macronucleus almost complete. q) Very late stage, 105 ~m.
32 GENERAL SECTION

FOISSNER 1997). For a detailed explanation and interpretation of these data, see the phy-
logeny section. A useful graphic method for summarising morphogenetic data is provided
by EIGNER (1997).
Dorsal morphogenesis proceeds rather "simply" in most hypotrichs, that is, each dor-
sal kinety usually forms an anlage in the proter and the opisthe by intrakinetal prolifera-
tion of basal bodies (FOISSNER & ADAM 1983b). Only in oxytrichids does at least one dor-
sal kinety (in most instances kinety 3) show usually simple, rarely multiple fragmentation.
Several types of dorsal morphogenetic patterns, which are very useful phylogenetic mark-
ers (BERGER & FOISSNER 1997), exist and are described and interpreted in detail in the
phylogeny section (Fig. 24). As stated above, proliferation of basal bodies begins at two
levels within parental rows (Fig. 10k). These two regions correspond to the same levels
within which the marginal cirri proliferate on the ventral surface (Fig. IOj, 1). Dorsal pro-
liferation is initiated near existing bristles, and new basal bodies insert linearly between
bristles of the same row (GRIMES & ADLER 1976). At this stage of morphogenesis, the
anterior-posterior axis of the primordial dorsal bristle unit is perpendicular to the longitu-
dinal axis of the cell. Bristles within the proliferation regions are apparently retained and
distributed among new bristles on differentiation from the reproductive streaks. The pri-
mordia then lose continuity with their original rows and move slightly laterally (for de-
tails and electron micrographs, see GRIMES & ADLER 1976). Proliferation and basal body
pairing continues until the segregation of basal bodies. At this stage, the axis of the pri-
mordial bristle units are more closely parallel with the longitudinal axis of the cell. Elec-
tron microscopy revealed that kinetodesmal fibrils are present only in nascent bristle units
and resorbed during maturation (Fig. 7c), which is obviously a very important (apomor-
phic) character for the separation of the Euhypotrichina from the Pseudohypotrichina (=
euplotids sensu lato; kinetodesmal fibre of dorsal dikinetids permanent; FLEURY 1988,
FLEURY et al. 1985, GRIMES & ADLER 1976, RUFFOLO 1976).
One (for example, in Urosoma), two (very common), or more (for example, in Ter-
ritricha) dorsal kineties develop from the right marginal row in oxytrichids (Fig. 10l-n)
and several other taxa, for example, Parakahliella (BERGER & FOISSNER 1989b). These
"dorsomarginal kineties" (MARTIN 1982) differentiate at the anterior limits of the right
marginal cirral primordia and move dorsally (Fig. 100). Usually, they are shorter than the
other rows and do not differentiate caudal cirri; however, the ultrastructure is identical
(GRIMES & ADLER 1976). According to these authors, it depends on the nutritional status
of the cell whether, in Oxytricha fallax, one or two dorsomarginal kineties are derived
from marginal cirral primordia. However, morphometric characterisation of many oxytri-
chids revealed that the number of dorsal kineties (including dorsomarginal kineties) is
rather constant within species and even higher taxa (for example, Urosoma). Cladistic
analysis suggests that dorsomarginal kineties are secondarily lost in Gonostomum (Fig.
24c). Almost all oxytrichids have caudal cirri which always originate at the posterior end
of the dorsal kineties 1, 2, and 4 (Fig. 24a, e), or on kinety 3 if fragmentation of this ki-
nety is lost secondarily (Fig. 24b, c).
 MORPHOLOGY 33

Table 3 Summary of data on morphogenesis during binary fission in oxytrichids with 6 (l-VI) frontal-ventral-
transverse cirri streaks!

Species number of sta- number of cirri origin of DM3 M Figures

[Reference]2 ges illustrated originating from streak oral in this
ventral dorsal I II III IV V VI primordium book
Allotricha 14~ Ip 3i 3 3 3 4 4 immediately 0 P 92n-y,
antarctica [32] leftofTC 93a, b
Allotricha mol/is [3] Ii 3 3 3 4 4 0 P 91e
Coniculostomum 12~ 8p 6i,5p 3 3 3-11 3-11 4-11 immediately C P 1681-z,
monilata [1] anteriad of 169a-c
leftTC
Cyrtohymena 2Ii 6i 3 3 3 4 4 somewhat 0 P 94r-z,
muscorum [2] left of PVC 958--5
Gastrostyla 4i 3i 3 3 4-7 6-10 6-9 near left TC? <Y" P 202j-p
dorsicirrata [3]
Gastrostyla 6i 2i 122 3 3 5 7 6 narrow field 0 P 203M
minima [35] between left
TC and buccal
vertex
Gastrostyla 8i 2i 1 2 3 3 9 7 left anteriad 0 P 201a-j
steinii [4] ofleftTC
Gastrostyla 3p Ip left anteriad 0 P
steinii [5] ofleftTC
Gonostomum 9i Ii 2 2 2 4 4-5 behind G P 120a-j
affine [4] AZM
Gonostomum 10i 6i 2 2-33-4 8-11 7-10 behind G P 122f-u
strenuum [6, 36] proximal part
ofAZM
Histriculus 2i 3i 1 3 3 3 4 4 near left TC O2! P 176f-j
histrio [7, 34]
Histriculus Sp.9 [8] IIp 4p immediately 0 P
left of left TC
Kerona 7i 4i 18 15 17 20 27 25 behind buccal K P 208n-y
pediculus [4, 33] vertex
Notohymena 2Ii,4p 7i 3 3 3 4 4 immediately 0 P 1041-z,
rubescens [9] leftofTC WI 105a-n
Oxytricha 3i Ii behind G P 83d-g
geleii [10] proximal part
ofAZM
Oxytricha 12i,6p 14i 1 3 3 3 4 4 somewhat 0 P 65h-z,
granulifera [11] left behind 66a-g
PVCVl3
34 GENERAL SECTION

Table 3 Continued'

Species number of sta- number of cirri origin of DM' M Figures

[Reference), ges illustrated originating from streak oral in this
ventral dorsal I II III IV V VI primordium book
Oxytricha 20p,6is 0 3 3 3 4 4 immediately P
hymenostoma [12] leftofTCWI
Oxytricha 0 4i6 0 s, t
hymenostoma [13]
Oxytricha Ii immediately P 691
lanceolata [14] leftofTCWI
Oxytricha lli Ii I 2 3 3 4 4 immediately U P 44f-n
longa7 [31] left of
PVCIV12
Oxytricha Ii immediately P 71h
longigranulosa [I5] leftofTC WI
andnearV13
Oxytricha 6i Ii I 3 3 3 4 4 immediately U P
pseudosimilis [4) leftofTC WI
Oxytricha 0 0 immediately P
rubripuncta [14] leftofTC WI
Oxytricha 4i 4i 1 3 3 3 4 4 between PVC U P 56f-m
setigera [7] andTC
Rubrioxytricha hae- 2i Ii U P I44f-h
rnatop/osrna'7 [16]
Steinia 3i 0 P 1721-n
p/atystoma [17]
Sterkiella 9i,3p 0 immediately P 181k
cavicola8 [17] leftofTC WI
Sterkiella histrio- 15i 5i I 2 3 3 4 4 immediately 0 P 10c-v
muscorum lO [I8] left ofTC 001
Sterkiella 0 0 near left TC P
terricolall [19]
Stylonychia 2i 3i 1 3 3 3 4 4 immediately 0 P 157d-h
bifaria'9 [20] leftofTC WI
(and 001)
Sty/onychia 7i Ii 1 3 3 3 4 4 immediately 0 P
mytilus [4] leftofTC WI
Stylonychia 6p 0 w
myti/us [21]
Sty/onychia 3i 0 1 3 3 3 4 4
mytilus [22]
 MORPHOLOGY 35

Table 3 Continued'

Species number of sta- number of cirri origin of DMl M Figures

[Referencej2 ges illustrated originating from streak oral in this
ventral dorsal I II ill IV V VI primordium book
Sty/onychia 5i,p 0 3 3 3 4 4 nearTCWl P
myti/us [23]
Sty/onychia myti/us IOi 0 3 3 3 4 4 immediately 0 P 153a-j
complex [24] leftofTC WI
Sty/onychia 6i 0 3 3 3 4 4 left of PVC w I62i
pustulata'l [25]
Stylonychia 9i 0 3 3 3 4 4 left of PVC 0 P 161e-n
pustulata [20]
Stylonychia Ii 0 near left TC P 165d
putrina [17]
Tachysoma 8i 0 3 3 3 4 4 near leftTC T P 135h-o
pel/ionel/urn [4]
Tachysoma 5i 0 3 3 3 4 4 immediately U P 139j-n
terricola [4] left ofTC 1111
and whole PVC
Urosoma I2i,5p 6i 3 3 3 4 4'6 left of PVC U P 127j-y
emarginata 13 [26]
Urosoma lli 3i 3 3 3 4 4 between PVC U P 134j-w
gigantea'S [271 VI3 andPTVC
VI2
Urosoma poly- Ii behindAZM P 130h
nucleata l4 [281
Urosomoida lli,2p lli 1 2 2 2 3 4 immediately U P I13c-v
agilijormis [29, left of PVC
37] (especially
V/3) and
near leftTC
Urosomoida 7i,8p 4i,2p 1 2 2 2 3 4 left behind U P 1l0k-q
agilis18 [30] PVC
Urosomoida Ii left of PVC P
minima [4]

I With few exceptions, only data based on silver impregnated material are considered. Abbreviations: AZM ==
adoral zone of membranelies, DM = dorsal morphogenesis (for types, see footnote 3), i == illustration (draw-
ing), I = from life, M = method, p = photograph, P = protargol impregnation, PTVC = pretransverse ventral
cirri, PVC = postoral ventral cirri, s = scanning electron microscopy, t = transmission electron microscopy.
TC = transverse cirrus(i), w = Chatton-Lwoff silver nitrate impregnation, - = not known or not available.
1 Species names are adapted to nomenclature of this book. References: 1 = KAMRA & SAPRA (1990), 2 = Voss
(l991b), 3 = original data, 4 = HEMBERGER (1982), 5 = WALKER & GRIM (1973). 6 = SONG (1990). 7 =
GsCHWIND (1991), 8 = NIETO et al. (1984), 9 = Voss (I991a), IO = WILBERT (1986b), II = FOISSNER & ADAM
36 GENERAL SECTION

(1983b), 12 = GRIMES (1972), 13 = GRIMES & ADLER (1976), 14 = BERGER & FOISSNER (1987a), 15 =
BERGER & FOISSNER (1989a), 16 = BLATTERER & FOISSNER (1990), 17 = GROLIERE (1969), 18 = BERGER
et a!. (1985a), 19 = BUITKAMP (1977b), 20 = WIRNSBERGER et al. (1985a), 21 = TCHANG et al. (1965),
22 = WALLENGREN (1902), 23 = WATANABE & HORA (1983), 24 = WIRNSBERGER et al. (1986), 25 = SA.
PRA & DASS (1970a), 26 = FOISSNER (1983a), 27 = BERGER & FOISSNER (l988b), 28 = FOISSNER (1984),
29 = FOISSNER & ADAM (1983a), 30 = BUITKAMP (1975), 31 = GANNER et al. (1987b), 32 = PETZ &
FOISSNER (1996), 33 = HEMBERGER & WILBERT (1982), 34 = BERGER & FOISSNER (1997), 35 = FOISSNER
(1997a), 36 =OLMo & TELLEZ (1997), 37 = WANG & SONG (1996).
3 Type of dorsal morphogenesis: C = Conicu/ostomum pattern (Fig. 24e), G = Gonostomum pattern (Fig. 24c),

K = Kerona pattern (Fig. 208u-y; mUltiple fragmentation in dorsal kineties 1-3 and two dorsomarginal kine-
ties), 0 = Oxytricha pattern (Fig. 24a), T = Tachysoma pattern (Fig. 24d), U = Urosomoida pattern (Fig.
24b).
5 TEM-micrographs also shown.
6 Many SEM- and TEM-micrographs.
7 In GANNER et aI. (1987b) designated as Urosomoida agi/ijormis.

• In GROLIERE (1969) designated as Opisthotricha monspessu/ana.

9 In NIETO et aI. (1984) designated as Histricu/us simi/is.

lOIn BERGER et aI. (1985a) designated as Histricu/us muscorum.

II In BUlTKAMP (1977b) designated as Histricu/us simi/is tricirratus.

12 In SAPRA & DASS (1970a) designated as Sty/onychia notophora.

13In FOISSNER (1983a) designated as Urosoma macrosty/a.
14In FOISSNER (1984) designated as Hemisincirra polynucleata.
151n BERGER & FOISSNER (1988b) designated as Oxytricha gigantea.
16FolSSNER (1983a) mentioned that within streaks I-VI the following number of cirri is formed: 1,2,3,3 (4),4,
5 (4).
17In BLATTERER & FOISSNER (1990) designated as Oxytricha haematop/asma.
18In BUITKAMP (1975) designated as Oxytricha agilis.
19In WIRNSBERGER et aI. (1985a) designated as Sty/onychia vorax.
lOIn Gastrostyla dorsicirrala dorsal kineties 3 and 4 are not separate in the interphase stage (arrowheads in
Fig. 202i), that is, fragmentation is incomplete.
21 No caudal cirri are formed.
22Number of cirri from the opisthe in Figure 203p.

1·10·2 Conjugation

Like other ciliates, oxytrichids have a sexual phase in their life cycle. Most have tempo-
rary conjugation. Mating type-specific soluble factors (gamones) are involved in the pre-
conjugant cell interaction (for example, ESPOSITO et aI. 1976b, RICCI 1981c). Conjugation
is preceded by a remarkable "mating game" ("Paarungsspiel"; GRELL 1951, SIEGEL 1956).
The activity of the cells remains unchanged for about two hours (waiting period) when
cultures of complementary types are mixed. Thereafter, small groups of cells collect on
the bottom of the culture dish and the cells within each group move in contiguous circles.
Soon, two such "dancing" cells meet at their anterior adoral surfaces and become rela-
tively quiescent. The mates so joined may presently separate and fmd other partners, or
become more firmly united (Fig. 11a). If conjugation proceeds, each cell is supported on
the bottom of the dish by its posterior cirri, and the pair forms an obtuse angle. This angle
 MORPHOLOGY 37

•
a b
Fig. Ua, b Stylo11)lChia lemnae (from D. AMMERMANN in STEINBROCK 1986). a) Conjugating pair. Cells are
connected in the anterior third by a cytoplasmic bridge. b) Exconjugant cell. anI = macronucleus anlage, rna =
old degenerating macronucleus. Bars 30 ~m.

is slowly reduced so that the cells fmally adhere tightly along most of their lengths and
conjugation is completed.
The cell union was studied at the ultrastructural level by RICCI et al. (l97Sa, b; see
also RICCI 198Ic). The cortical apparatus of vegetative cells of "Oxytricha hi/aria" is
composed of a singular external pellicle and the cell membrane. Cross-sections of conju-
gating pairs, 10-13 min after the onset of the visible reaction, revealed that protuberances
of only slightly dense cytoplasm are formed at the peristomial level. They are surrounded
by the pellicle and extend toward the partner's cytoplasm. The fact that they are found
only within the peristomial area suggests that these protuberances could be the expression
of the gamone-induced cortical changes. A few minutes later, the situation dramatically
changes: couples in the early rotation stage reveal that the external peIIicles of the two
partners are already fused in the anterior portion of the ventral area. The two cell bodies
are still well separated and a highly vacuolised sack from the left partner extends within
the joined pellicles to fill the gap between the two mates. Before rotation is over, lysis of
38 GENERAL SECTION

C-3-7h D-8-9h

E-l0h F_ 12h

1_15h 30min L - 16h M -16h 30mln N _16h 50mln

o -17h30mln P _ 18h 30min Q -19h 20mln R_ - 20h

II
 MORPHOLOGY 39

the cell membranes occurs and a clear continuity between the two cytoplasms is ob-
served. At the level of the fusion areas, some mitochondrial clustering occurs. Further de-
velopment of the cell union involves the differential reabsorption of parts of their adoral
zone of membranelles and several other ciliary structures in order to form the new peri-
stomial area of the pair. The rotating partner undergoes a dramatic cortical rearrange-
ment, reabsorbing the lapel of its adoral zone, the frontal cirri, and the anterior area of the
ventral surface. The still partner reabsorbs a smaller area, which includes the collar of its
adoral zone of membranelIes and the most distal portion of the ventral surface. This dif-
ferential, well co-ordinated reabsorption results in a new, mutual adoral zone of mem-
braneHes, which is practically indistinguishable from the normal adoral zone of vegeta-
tive cells. About 15 h after joining, the cytoplasmic bridge extends over approximately
the anterior third of the length and over ahnost the entire dorsoventral height. No particu-
lar limiting structure can be observed. Finally, after about 18-20 h, the cytoplasmic
bridge progressively narrows so that the cells are united by a thin, densely vesiculated cy-
toplasmic cord. This connection is eventually broken by the active uncoordinated swim-
ming of the partners, which are now referred to as exconjugants (Fig. lIb). For the nu-
clear events during conjugation, see Figure 12.
The sexual phase lasts for nearly 125 h in species of the Sty/onychia myti/us com-
plex. The cells are paired as conjugants for the first 25 h and the remainder of the period
is occupied by the development of the separated exconjugant cells.
ESPOSITO et al. (1976a) described total conjugation in Oxytricha hymenostoma. While
the pre-conjugative pattern is quite similar to that described for other oxytrichids, the cy-
toplasmic bridge uniting the partners progressively elongates up to the posterior end of
the cells, instead of being limited to the level of the first anterior third. At this stage
(about 6 h after the onset of mating), the partners are morphologically indistinguishable,
being fused in one single cell whose cytoplasm arises equally from both conjugant cells.
This trait differentiates total conjugation in O. hymenostoma from that of other hypo-
trichs with total conjugation (Paraurostyla weissei, HECKMANN 1965, JERKA-DZIADOSZ &
JANUS 1975, MOLDENHAUER 1965, Fig. 214; Pseudourostyla laevis, TAKAHASID 1973).
About 13 h from the onset of cell pairing (at 23°C), the new cell divides to produce two
daughter cells through binary, equal transverse fission. Macronuclear meiosis and karyo-
gamy were never observed. Only two of the four initial micronuclei are typically present
at the end of the process; it is supposed that some micronuclei undergo picnosis. The two
macronuclear nodules of the rotating partner at first elongate and then fuse, about 3 h af-
ter the onset of cell union. From 3-7 h later, the macronuclei of both partners shorten and
then fuse two by two, anterior with anterior and posterior with posterior. At this point, a

+- Fig. 12 Schematic drawing of the major stages of the nuclear events during conjugation in an oxytrichid cili-
ate (from RIccI & BANCHETIl 1981). I) Stages from the beginning of cell pairing to the end of meiosis (A-H).
II) The end of maturative divisions (1, L), the formation of synkaryon (M, N), the first four nuclei produced by
its division (O-Q), and the exconjugant (R). m = migratory pronuclei, s = stationary pronuclei, Sy =
synkaryon.
 40 GENERAL SECTION

Fig. 13a-d Morphogenetic processes during conjugation in a species of the Stylonychia mytilus complex
(from TUFFRAU et aI. 1981. Protargol impregnation). a, b) Early stages of conjugation. c, d) First reorganisation
step. Arrows in (c) mark anterior end of oral primordium. For details, see text

cell provided with the typical bipartite macronucleus is formed. About 12 h after the be-
ginning of the process, the macronucleus shows replication bands which lead to the nor-
mal binary fission. The term conjugation is not quite proper for this phenomenon because
there is no meiotic process. However, it is a specific mating-type dependent process,
leading to the fusion of genetic macronuclear material belonging to different complemen-
tary cells ("hologamy-like process" or ''paraconjugation''; BANCHETII et at. 1980a, b,
 MORPHOLOGY 41

~;~
,/~.~
~ ,
,~7>

/ "z:~:

A-,
f
,::\::
"

.". . . . . . . . . . . . . . "=>0,.
.

P " -~

F'r' ,"-"
~:.:':: . !~;:
.I;,~'.

Fig. 13e-i Morphogenetic processes during conjugation in a species of the Sty/onychia mytilus complex (from
TUFFRAU et aI. 1981. (}-i, protargol impregnation). e-g) First reorganisation step. Cirri and adoral membranelles
originating during this step are dotted. h) Exconjugant after first reorganisation step. Arrow marks a cortical
flexure at site of future cytostome. Arrowhead denotes macronucleus anlage (cp. Fig. II b). i) Begin of second
reorganisation step. For details, see text.
42 GENERAL SECTION

r~-,~7,~'
"~:~:~Z~"
{,,;( ------ --~
/f.~I1l;';-::-~
14/"" /,/i::~~_
___ '£

f; if~ 'y mi~;~1

~ i/ // -:--~ "":1,,
: ;$f:---::~
III • • .'/: ;~ >i~
• ') U..

-- :•-.'1, UF~/
\

c'i~\_ ~##
I,
h

~
~
if
i:!

·;1
~ I r1 I~' \»~-jo
-<)i/)fL'-';;
p

Fig. 13j-p Morphogenetic processes during conjugation in a species of the Sty/onychia mytilus complex
(from TUFFRAU et al. 1981. j-p, protargol impregnation). j-m) Second reorganisation step. Cirri and aQoral
membranelles originating from this step are hatched. n-p) Third reorganisation step. One oCthe anterior micro-
nuclei commences mitosis. The proximal portion of the adoral zone of membranelies is disorganised (arrow in
p) and replaced by the membranelles formed by the oral primordium. Final stage of this process, see Figure
151 r. Cirri and adoral membranelles originating from the third reorganisation step are black. For details, see
text.
 MORPHOLOGY 43

NOBILl et al. 1981). Autogamy can be induced in Stylonychia spp. by using the split-pair
method (JARENO et al. 1969b, 1970). FERMOR (1913) and RAo (1958) described autogamy
in encysted specimens of Stylonychia pustulata. No nuclear processes were observed in
cysts of S. mytilus (AMMERMANN 1965a).
The changes of the infraciliature during conjugation are rather complicated (Fig.
13a-p). In a species of the Stylonychia mytilus complex, a localised dedifferentiation af-
fects the conjugant's anterior infraciliature when they mate (TUFFRAU et al. 1978a, 1981).
The result is a transitory dimorphism which precedes the three reorganisation steps
through which both partners return to the vegetative state. During the first step, while a
new somatic infraciliature is restoring itself on both the ventral and dorsal side, only the
anterior portion of the adoral zone of membranelles renews (Fig. 13c-g). The second re-
organisation completes the preceding adoral zone by adding a new membranellar zone
and it gives rise to the missing paroral structure (Fig. 13i-m). The whole somatic infra-
ciliature is restored. In a third stage, the infraciliature of both sides is reorganised for the
third time. In addition, a new series of adoral membranelies is formed to replace the pre-
ceding membranelles (Fig. 13o-p). Finally, the adoral zone ofmembranelles comprises
three parts, namely the distal part which is formed in the fIrst reorganisation step, the
middle one from the second step, and the proximal part from the third.
Further literature on the sexual phase of oxytrichids (see also chapter 1·2, Nuclear
Apparatus): AnL & BERGER (1995), ALONSO & PEREz-SILVA (1965b), AMMERMA:NN
(1965a, 1967, 1982), BAITSELL (1911, 1912), BALBIANI (1861, 1882), BANCHETfI et al.
(1978a, 1982b), BULL & PEASE (1989), BOTsCHLI (1875, 1876), CETERA et al. (1978),
CHADHA et al. (1978), DILLER (1965b), DINI & LUPORINI (1974), DINI & NYBERG (1993),
DoWNS (1952, 1956, 1959), ENGELMANN (1862, 1876), ESPOSITO & NOBILl (1982), Espo-
SITO & RIccI (1975), ESPOSITO et al. (1974, 1976b), FLEURY (1983), FLEURY & FRYD-
VERSAVEL (1981), FLEURY & GUYADER (1990), FRONTCZAK-BANIEWICZ & JERKA-DZIADOSZ
(1992), GIL (1976), GRELL (1965), lIAMMERsMITIl (1976a, 1978), HEUMANN (1975),
ILOWAISKY (1916), JARENO et al. (1969a, 1970), JERKAl)zIADOsz & JANUS (1975), KAy
(1946), LICHTENBERG (1955a), Lu et al. (1991), LUPORINI & DINI (1975), MIYAKE (1974,
1981, 1982), MOLDENHAUER (1965), NOBILl et al. (1987), PIERI (1966), PREER (1969),
RIccI (1992), RIccI & CETERA (1978), RIccI et al. (1975a, b, 1977, 1978, 1979a, 1980a,
c), SAPRA & AMMERMANN (1974), SAPRA & KLOETZEL (1974, 1975), SHARMA et al. (1986),
Sm (1976), STEIN (1859b), TCHANG & PANG (1979b), TURNER (1964), WOODRUFF (1905a,
1964), Xu & Sm (1987), YANO (1985a-d), YANO & SUHAMA (1990b, 1992), Zou & NG
(1991b, c).

1·10·3 Cyst

Many oxytrichids are able to form resting cysts. Cysts with a smooth, wrinkled and a
spiniferous surface are known (Fig. 14a, b, 181d). Reproductive cysts are not described
in this group. Factors that induce encystment are, among others, deficiency of food or
desiccation (for example, CORLISS & ESSER 1974, MICHELSON 1928, PENN 1935).
44 GENERAL SECTION

G
S
CW
Ma

Fig. 14a, b Resting cysts of oxytrichids from life (a, from FOISSNER & ADAM 1983b; b, from FOISSNER 1989).
a) Oxytricha granulifera has a cyst with a smooth surface, 32 ~m across. b) Steinia sphagnicola has a spini-
ferous cyst, 53 ~m in diameter without the about 7 ~m long spines. CW = cyst wall, G = granules on cyst wall,
Ma = macronucleus, S = mucous layer.

The ultrastructure of the cyst has been studied in several oxytrichids (for example,
CALVO et al. 1983, MATSUSAKA 1976, MATSUSAKA & HONGO 1984, ROSATI et al. 1983,
1984, VERNI et al. 1984, WALKER & HOFFMANN 1985, WALKER et al. 1975, 1980; Fig.
14c). The cilia, the basal bodies, and the microtubules are resorbed during encystment;
the mature cysts are thus termed kinetosome-resorbing (GRIMES 1973b, d, WALKER &
MAUGEL 1976). In a second group ofhypotrichs (for example, Diophrys scutum), the ma-
ture cyst is non-kinetosome resorbing and the cyst wall has three layers. By contrast, the
cyst wall of the oxytrichids is composed of four layers, namely the ectocyst, the
mesocyst, the endocyst, and the granular layer. In Stylonychia mytilus, the ectocyst ap-
pears either amorphous or lamellar, while the mesocyst appears to be fibrous in nature
and to have either a "herringbone" or lamellar configuration. The endocyst is amorphous
in structure. The cytoplasm is enveloped by a layer of deeply folded, compacted membra-
nous material beneath the granular layer of the wall (WALKER et al. 1975). The cyst wall
is derived from various precursors which are synthesised new in the cytoplasm of the pre-
cystic cell (CALVO et al. 1986b, GRIMES I973d). After the precystic cell becomes
spherical, the precursor vesicles migrate toward the cell surface, where they expel their
contents between the two cell membranes (the distal perilemma and the proximal plasma
membrane) of the vegetative cell, forming the ectocyst. The perilemma presumably func-
tions as an envelope to keep the precursers from dispersing after deposition (BARDELE
1981, GRIMES 1973d). Macronuclear nodules fuse during encystment (GRIMES 1973d,
WALKER & MAUGEL 1980). Mature cysts of oxytrichids usually retain only about 10-20 %
of their vegetative volume due to strong dehydration.
 MORPHOLOGY 45

Fig. 14c Section through a resting cyst of Oxytricha sp. (TEM-micrograph from GRIMES 1973d. Glutaralde-
hyde fixation, postfIxed in OsO.). Diameter of cyst 18-20 flm, cyst wall 2-3 flm thick. Explanation of original
labels: CW = cyst wall, MAC = macronucleus, Mi = mitochondrial bands, mic = micronucleus, nu =
nucleolus, S = starch-like grains in clusters, V = autophagic vacuoles.
46 GENERAL SECTION

NAKAMURA & MArsusAKA (1992a) found in Sterkiella cavicola that (i) excystment re-
quires protein synthesis, and incubation of cysts in excystment medium for longer than
30 min, (ii) basal body formation does not need protein synthesis and does not necessar-
ily induce cysts to complete excystment, and (iii) maintenance of new basal bodies re-
quires continuation of the excystment process.
Cortical morphogenesis during excystment (including the time before and after the
release of the cell) shows clear differences from the morphogenetic processes during divi-
sion (CALVO et al. 1988). The oral and frontal-ventral-transverse primordia originate from
an extensive field of basal bodies. The primordia ofthe marginal cirri and the dorsal bris-
tles appear at an early stage of morphogenesis. The left marginal primordium originates
from the oral primordium. The whole process is described in detail for Coniculostomum
monilata by I<AMRA & SAPRA (1991; Fig. 15a-t). This species, which has a typical 18-
frontal-ventral-transverse cirral pattern and multiple rows of right marginal cirri, requires
at least 3 morphogenetic cycles to acquire the normal vegetative ciliature during excyst-
ment. In the first cycle, the frontal-ventral-transverse cirral pattern is entirely different
from that formed during division (Fig. 15j, k). The "first ciliature" comprises 21-32
frontal-ventral-transverse cirri formed by 5 primordia. These cirri are substituted by 18
frontal-ventral-transverse cirri developed from 6 primordia in the subsequent reorganisa-
tion cycle (Fig. 151-0). Furthermore, each successive cycle adds one right marginal row
and 2 dorsal kineties; previous rows are not resorbed, unlike the frontal-ventral-transverse
cirri and the left marginal cirri (Fig. 15p-t). The mature cyst has a spherical macronu-
cleus and 1-3 micronuclei. The macronucleus maintains this shape throughout the early
cortical events of excystment. It stretches linearly and acquires the characteristic multi-
nodular shape when the cell begins to elongate. The micronuclei divide, leading to the
usual set of 4-5 micronuclei per cell.
Further literature on cysts or en- or excystment of oxytrichids: BARolN-TOURANCHEAU
et al. (1997), BERGER & ForsSNER (1987a), BRAND (1923), BUSSER & JEUNIAUX (1974),
CALVO & DE MIGUEL (1995/96), CALVO et al. (1981, 1992), CHARoEZ (1986), CIEN-
KOWSKY (1855), DEITMER (1987), FERMOR (1913), FLEURY (1983), FLEURY & GUYADER
(1990), ForSSNER (1982, 1989), GIESE (1951), GRIMES (1982b), GRIMES & HAMMERsMITH
(1980), GUTIERREZ (1985), GUTIERREZ & PEREZ-SILVA (1980, 1983a, b), GUTIERREZ et al.

Fig. 15a-f Resting cyst (a) and morphogenetic processes during excystment (b-f) in Coniculostomum moni- ~
lata (from KAMRA & SAPRA 1991. a, from life; b-f, protargol impregnation). a) Mature resting cyst, size after
protargol impregnation 76-94 Jim. b) Development ofa field of basal bodies, 70 Jim. c) Differentiation ofba-
sal body field into primordia, 65 Jim. Arrowheads mark primordia of frontal-ventral-transverse ciliature. d) Be-
ginning of differentiation of adoral membrane lies, 65 Jim. e) Formation of cirri, 105 Jim. The primordium of
the undulating membranes is divided. f) Polar view showing ventral ciliature and anterior part of developing
dorsal ciliature, 90 Jim. Arrowheads mark dorsal kineties which originate from right marginal rows (dorsomar-
ginal rows). Explanation of original labels: AZM = adoral zone of membranelles, BC = buccal cirrus, F = pri-
mordial field, IUM = inner undulating membrane (endoral), LMCP = primordium for a single left marginal
row, OP = oral primordium, OUM = outer undulating membrane (paroral), RMCP = primordium for a single
row of right marginal cirri, UMP = primordium of undulating membranes, I-V = frontal-ventral-transverse
cirri primordia J-V, 1-3 = primordia of dorsal kineties.
 MORPHOLOGY 47

/
/~,

:: .
/
:. .:";.
°
0: : " . : : °0

UMf AZM
LMCP

'. c"
".
,

RMCP-\
"' ......:
: .. ~ ·::"':'::':'~OP
i-RMCP 'iZ
-:. ..•.:.:
""

c .:;:
d

Q
o
o 0
I) \)
o 0
e C) CJ
 48 GENERAL SECTION

c
h

v 0 ------
--
Q
'"
","'0 ---.....
\ \ \ \ \ I I I I
'"'" 0 0 ::::
Co
" 0 D '"'" Q ~
-:::::

~
Co

'"'" 0 .::.
Q DD
"" 0 ~ D ~
~
~
C>
0
~ 00
C> 0 D
~
~oo~
C>

0
a
°aO
D @ Q
0
~
~~
C>

0
o0 c> 0 "
c>
c>
0 '"'"
<=>
a 0 ~~ <? "
~ ()
'"'"c>
'"'" 0 0 ~$(>
§? <?
'"'"c> 0 S:S (> 0 0
0
D " g
(> <? 0 0 '"
'"'"
<?
<?(J \) Q
00
c> <?

'"'"
(/0 0 c>
0 <?
0 '00
0° '"
0 <?
00 0
00 0 <?
'"
0 '" <?
'"
GGa
a0 '"'"
<?
<? '"
'"'"
'"
co <?
"" '" k

Fig. lSg-k Morphogenetic processes during excystment in Coniculostomum monilata (from KAMRA & SAPRA
1991. g-k, protargol impregnation). g) Dorsal view of first ciliature, 115 J.lm (seen from ventral!). Dorsal pri-
mordia I and 2 yield kineties I and 2 (primordium I is sometimes fragmented initially, as in this cell, but it al-
ways forms only I kinety). Dorsal primordium 3 undergoes fragmentation and the overlapping rows of cilia
later realign to yield kineties 3 and 4. Arrows mark caudal cirri, arrowheads denote dorsomarginal rows. b, i)
Releasing ofa cell from the cyst wall in ventral (seen from dorsal) and dorsal view, 80 J.lm. j, k) Ventral view
ofexcysted cells before (j; 100 J.lm) and after (k; 145 J.lm) elongation. Arrows mark single right marginal row.
e = cyst wall, EN = endocyst, 1-3 = primordia of dorsal kineties.
 MORPHOLOGY 49

(1980a-e, 1981, 1982a, b, 1983a, b, 1984), lIAIME (1853), lIAMMERSMITII & GRIMES
(1981), HAsHIMoro (1956, 1960, 1962, 1963, 1964), HAUSMANN & FOISSNER (1986),
ILOWAISKY (1926b), IVANIC (1931a), JARENO (1975, 1977, 1979, 1980, 1984, 1985, 1987b,
c, 1988), JARENO & TUFFRAU (1978, 1979), JEFFRIES (1959, 1962), KAMRA & SAPRA
(1991), KAuSHAL & SAXENA (1978), KAy (1945b), KNAIPPE et al. (1989), MANwEll
(1928a), MArsusAKA (1974, 1977, 1979), MArsusAKA & KIMuRA (1981), MArsusAKA et
al. (1984), MAUPAS (1888), NAKAMURA (1989), NAKAMURA & MArsusAKA (1985, 1991,
1992b), NOGucm & MArsusAKA (1989), NUSSBAUM (1886, 1897), OLMO & TELLEz (1997),
RAMMELMEYER (1931), REID & JOHN (1983), REUTER (1963b), RICCI & BANCHETfI (1982),
RICCI et al. (1985), SENDO & MArsusAKA (1982), SOKOLOFF (1945), SOKOLOV (1917), SON-
NEBORN (1977), STEIN (1859b), VELASCO et al. (1992), WALKER (1976a), WEISSE (1856),
WENZEL & OMILIAN-Rosso (1982), WEYER (1930).

1·1 0·4 Reorganisation, Regeneration, Doublets

Oxytrichids produce ciliature not only during binary fission or other normal parts of the
life cycle (conjugation, excystment) but also during physiological reorganisation and
post-traumatic regeneration (BALAMUfH 1940a, NG 1990).
Physiological reorganisation (Fig. 16a-h) is defmed as morphogenesis which re-
establishes a complete set of ciliary structures in an intact morphostatic (non-dividing)
cell (GRIMES & ADLER 1978). Usually, this process is a response to an altered nutritional
status induced by unfavourable culture conditions (for example, starvation) or other more
subtle changes in the environment. Physiological reorganisation is described for Conicu-
lostomum monilata (KAMRA & SAPRA 1990), Stylonychia mytilus (DEMBOWSKA 1938,
1939, GRIMES & ADLER 1978, Zou & NG 1991a), S. pustulata (GRIMES & ADLER 1978),
and Paraurostyla weissei (WIRNSBERGER et al. 1985b). DEMBOWSKA (1938) reported that
Stylonychia mytilus can live in pure tap water without any food organisms for 14-19
days. The whole ciliature and the nuclear apparatus is reorganised several times during
this period. Reorganisation begins after 1-6 h of starvation (Zou & NG 1991a). Reorgan-
isers of S. mytilus can be distinguished under the dissecting microscope by their shorter
body and the somewhat tapered posterior end. Reorganisation takes about 4 h at 25°C
(Zou & NG 1991a).
Cortical reorganisation commences with the development of an oral primordium near
the left transverse cirrus in Sty/onychia (Fig. 16a; GRIMES & ADLER 1978, Zou & NG
1991a). It is differentiated to 16-36 (mean ± SD = 25.9 ± 4.8; n = 29) membranelles
which replace the proximal portion of the old adoral zone in S. mytilus. New mem-
branelles thus comprise about 36 % of the reconstituted adoral zone. Interestingly, the
new membranelles at the anterior end of the primordium, abutting on the old adoral zone
of membranelles, appear as discontinuous fragments. It seems that these fragments are
later joined together (Zou & NG 1991a). The oral primordium obviously does not partici-
pate in the development of the frontal-ventral-transverse cirri during reorganisation.
Their primordia originate from old cirri (frontoventral cirri III12, IV/3, buccal cirrus lI12,
50 GENERAL SECTION

o o
o 0

'"'"
.......' \ t f I,. ,. __

::3 0 o -..
-..
Q

::.

,
~
~
--
§;f
~
~
\\\\\\~\\~'"
'"'"'"g
'"'"
'"
'"'"'"
'"
'"'"'"
'"
o o'"'"
'"

Fig. lSl--q Morphogenetic processes during excystment in Conicu/oslomum monilala (from KAMRA & SAPRA
1991. l-q, protargol impregnation). I-n) Three stages in the first reorganisation cycle, 1 = 160 Jim, m =
165 Ilm, n = 150 Ilm. An oral primordium forms 30-40 membranelles that emerge with the existing adoral
zone of membranelles. 18 new frontal-ventral-transverse cirri originate from 6 frontal-ventral-transverse pri-
mordia and replace the "old" cirri. The "old" left marginal row is replaced. A new right marginal row (double
arrows) is formed while the existing one (arrow) is displaced inwards (new cirri black). 0) A cell at the end of
the first reorganisation cycle, 160 11m. Note the presence of 18 frontal-ventral-transverse cirri and 2 (arrow and
 MORPHOLOGY 51

\\ \ \ \ \ \ \ J
1/\11 III I ,," " '1.1

....... - /

~
'.
.:"!! .' '.. , ',',
\..
., ;
-:: ::!!' ,'",
\

: , \
/
! 1
i

4: 3.- 2: 1
3- 2:

:i

, 0
o
t
0
<> s t

Fig. ISr-t Morphogenesis of dorsal ciliature (seen from ventral) during excystment in Coniculostomum moni-
lata (from KAMRA & SAPRA 1991. r-t, protargol impregnation. Arrows mark caudal cirri). r) Beginning of ftrSt
reorganisation cycle in a cell with 2 dorsomarginal rows (arrowheads), 170 11m. s) Beginning of second reor-
ganisation in a cell with 4 dorsomarginal rows (arrowheads), 235 11m. t) An excysted cell after 2 reorganisation
cycles, 245 11m, The cell resembles a vegetative cell. Arrowheads mark dorsomarginal rows. 1-4 '" dorsal kine-
ties 1-4.

and in S. mytilus possibly from postoral ventral cirrus V/4; Fig. 16b, c). Five streaks are
formed which produce 17 cirri; as usual the left frontal cirrus (cirrus 111) originates from
the anterior portion of the reorganising undulating membranes (Fig. 16c, d). The primor-
dia for the left and right marginal cirri and for the dorsal kineties occur at about the same
level as the frontal-ventral-transverse primordia. Reorganisation of these structures is
identical to that during binary fission. For the changes of the nuclear apparatus during re-
organisation, see DEMBOWSKA (1938) and Zou & NG (l991a).

~ double arrow} right marginal rows, p) A stage in the second reorganisation cycle showing the addition of a
third right marginal row (triple arrow) to the two rows (arrow and double arrow) already present, 205 11m.
Some adoral membranelles are formed and added to the proximal end of the adoral zone, The old frontal-
ventral-transverse cirri and the left marginal cirri are replaced (new cirri black), q) An excysted cell after two
reorganisation cycles with 3 right marginal rows (arrows), 235 11m,
52 GENERAL SECTION

Fig. 16a-b Physiological reorganisation of the ventral (a--d) and dorsal (e--h) ciliature of Sty/onychia (from
GRIMES & ADLER 1978). Arrow in (c) marks fusion of oral primordium with old adoral zone of membranelles.
Old cirri and dorsal bristles indicated by dots, new structures indicated by open circles. Explanation of original
labels: AZM = adoral zone ofmembranelles, BC = left frontal cirrus (cirrus III), CC = caudal cirri, FVfP =
frontal-ventral-transverse primordia, LMC = left marginal row, LMCP = left marginal primordial field, OP =
oral primordium, RMC = right marginal row, RMCP = right marginal primordial field, UM = undulating
membranes, 1-6 = dorsal kineties 1-6 (kineties 5 and 6 are dorsomarginal rows, that is, they originate
near/from right marginal row).

Post-traumatic regeneration (Fig. 17a-k). Hypotrichs (including oxytrichids) have

a remarkable regeneration capability after mutilation, for instance, by a predator or ex-
perimentally induced (for example, mechanically, or chemically due to treatment with

Fig. 17a-k Post-traumatic regeneration of a transverse fragment of Sty/onychia mytilus (a-d, after GRIMES &
L'HERNAULT 1979 from NG 1990) and of longitudinal fragments of Sty/onychia (f-Ie, after GRIMES & ADLER
1978 from NG 1990) in ventral view. Open circles represent pre-existing cirri, filled circles represent regener-
ated cirri. a-d) Regeneration of a large posterior fragment ("opimer") generated by removal of the anterior part
of the cell. The oral primordium (arrowhead) arises near the left transverse cirrus; as it expands anteriorly it
connects with the posterior-most region of the pre-existing adoral zone of membranelles, where old mem-
branelles are replaced by new membranelles. The frontal-ventral-transverse primordium originates by disaggre-
gation of some pre-existing frontoventral cirri and differentiates into 5 streaks (double arrowhead); the undu
 MORPHOLOGY 53

17a
c d

o
00 0
00
00

I f
e \
• 0 0
o 0
000
oo~,
o ~~
o 0 o~
00

......
00

lating membrane primordium arises from the dedifferentiated pre-existing undulating membrane. The anterior-
right part of the oral primordium might also contribute to both of these primordia. The primordia of the mar-
ginal rows differentiate within the pre-existing marginal rows. All old ciliature is replaced, except a part of the
adoral zone of membranelles. e-k) Regeneration of right and left fragments, produced by longitudinal cuts,
generally resemble that in the large opimer. The illustrations show the origin of the oral primordium (arrow-
head) and its integration into the proximal part of the pre-existing adoral zone of membranelles. In left frag-
ments (i-k) the frontal-ventral-transverse primordia and the right marginal primordium originate in continuity
with the oral primordium because old frontoventral and right marginal cirri are absent This indicates that the
oral primordium is able to generate a complete ciliature (GRIMES & ADLER 1978).
54 GENERAL SECTION

urea). Regeneration is described in more or less detail for Gastrostyla stein;; (NIETO et al.
1981), Histriculus (CALVO et al. 1984, NIETO et al. 1984), Oxytricha (BAUER & GRANOWS-
KAlA 1934, BISHOP 1943, HAsHIMOTO 1961, HORVATH 1939b, ISHIKAWA 1912, REYNOLDS
1932), Stylonychia spp. (DEMBOWSKA 1925, DOROSZEWSKI & RAABE 1966, FRICK 1968,
GRIMES & ADLER 1978, Gu et al. 1990, GRIMES & L'HERNAVLT 1979, ISHIKAWA 1912,
PRowAZEK 1899a, 1903, SCHILD 1921, Zou & NG 1991a), and Paraurostyla weissei
(FRANKEL 1973, 1975, JERKA-DZIADOSZ 1967, 1974). The complete form of as. mytilus
cell is regenerated within 6-24 h after removal of the anterior or posterior portion of the
cytoplasm (JANISCH 1959). In addition, DEMBOWSKA (1925) found that (i) notches in vari-
ous directions and the removal of cytoplasm pieces without cirri or cilia do not induce re-
generation, (ii) amputation of only one cirrus causes complete reorganisation, and (iii)
duration of regeneration is independent of the size of the trauma. GRIMES & ADLER (1978)
concluded from the regeneration of longitudinal fragments (Fig. 17e-k) that (except for
the oral primordium) all primordial fields responsible for replacing excised ciliature de-
velop in continuity with existing primordia, that is, all ciliary primordia are derived from
segments originally continuous with the oral primordium. This is a major difference to
physiological reorganisation, where the oral primordium is not included in the formation
of other primordia.
The results on the role of the macro- and micronucleus during regeneration are con-
tradictory. REYNOLDS (1932), for instance, studied regeneration in an amicronucleate Oxy-
tricha, whereas HORVATH (1939b) found in the same taxon, that amicronucleate fragments
do not regenerate, even when a macronuclear nodule is present. In Stylonychia mytilus re-
generation takes place only when the fragment contains at least one macronuclear nodule
and one micronucleus (JANISCH 1959).
Further literature on reorganisation and regeneration: BALAMUTH (1940b), DUJARDIN
(1841), FAURE-FREMIET (1948,1967), FLEURY (1983), HEWITT (1914), LEWIN (1912), Nuss-
BAUM (1884, 1886), SUMMERS (1964), TARTAR (1941), and YOUNG (1926).
Doublets are developmental anomalies where two cells form a single individual (Fig.
18a, b). TOTWEN-NoWAKOWSKA (1964) observed doublets of Stylonychia mytilus after a
mass conjugation. Experimentally doublets were induced by ultraviolet light, laser mi-
crobeam, thermic shocks, or microsurgery (GRIMES 1976, TCHANG & PANG 1965a, b,
TOTWEN-NoWAKOWSKA 1965). Homopolar and mirror-image doublets are known in oxy-
trichids. The cells of a homopolar doublet are joined dorsally (Fig. 18a) whereas mirror-
image doublets (originally called ''jumelles'' by the Chinese workers; TCHANG et al. 1964)
are always joined side to side, with the two sets of ventral ciliature on one plane and the
two dorsal sets on the opposite plane (Fig. 18b; FRANKEL 1989). The left component has
an arrangement of cortical structures that is typical for the species (thus called "right-
handed component" by Sm & FRANKEL 1990), whereas in the right component the large-
scale arrangement is reversed (thus termed "left-handed" in Fig. 18b).
For further details on the morphology, development, and physiology of doublets, see
BANCHETTI & RICCI (1986), BANCHETTI et al. (1996), DAWSON (1920), DRYL &
NOWAKOWSKA (1966), DRYL & TOTWEN-NoWAKOWSKA (1972, 1975), DRYL et al. (1981),
 MORPHOLOGY 55

a b
Fig. 188, b Doublets of oxytrichids (a, from DAWSON 1920; b, from SHI & FRANKEL 1990. a, from life; b, pro-
largol impregnation). a) Homopolar doublet of an amicronucleate Oxytricha. b) Ventral ciliature of a mirror-
image doublet of Sty/onychia mytilus. Explanation of original labels: AZM = adoral zone of membraneUes, LH
= left-handed component (see text), LM = left marginal row, RH = right-handed component (see text), T =
transverse cirri, UM = undulating membranes.

FAURE-FREMIET (1945a, b), FENG et al. (1989, 1990), FRANKEL (1984, 1989), GRIMES
(1973c, 1976), GRIMES & lliMMERSMITII (1980), GRIMES et al. (1980, 1981), lIAMMER.-
SMITII (1976b), JARENO (1981b, 1992), JERKA-DZIADOSZ (1977, 1983, 1985, 1989), LAND-
MAN (1991), PANG et a1. (1984), RICCI et a1. (1996), Sm (1987), Sm & FRANKEL (1990),
Sm et al. (1987, 1990, 1991a, b), TCHANG & PANG (1977, 1979c, d), TOTWEN-
NOWAKOWSKA (1964, 1965, 1969, 1973, 1981), ThFFRAu & TOTWEN-NoWAKOWSKA (1988),
TUFFRAU et al. (l978b), YANO & SUHAMA (1991), ZHANG & PANG (1981), ZHANG et al.
(1982).
56 GENERAL SECTION

2 Phylogeny
Oxytrichids are usually considered to be highly evolved hypotrichs (for example, KAHL
1932, BORROR 1972a). Until recently, this was almost all we knew about the phylogeny of
this group. Only WIRNSBERGER et al. (1986) attempted investigating the branching se-
quence of seven common oxytrichid genera. Since then the number of genera has in-
creased significantly, and more data on morphology and morphogenesis have become
available (Tables 3, 4). I made a further attempt to elucidate the phylogenetic relation-
ships within the Oxytrichidae (BERGER & FOISSNER 1997, 1997a) using lIENNIG'S method
which is described in detail by Ax (1984, 1995), lIENNIG (1966, 1982), and SUDlfAUS &
REHFEw (1992). In addition, the computer programme PAUP was used (SWAFFORD 1991).
In contrast to computer analysis, the traditional method ("search for the sister group") re-
quires that the plesiomorphic and apomorphic state of a character is defined before the
analysis is performed (MEIER 1992). Out-group (for example, holostichids) and in-group
comparison was used to distinguish plesiomorphies from apomorphies (Ax 1984).
Twenty-three characters were selected, namely the morphology of the oral apparatus
(3 characters), infraciliature of ventral and dorsal side (10 characters, two of them are un-
informative within the 18 cirri oxytrichids), cortical features (2 characters), and ontoge-
netic particulars (8 characters). Thus, only those taxa could be considered in the cladistic
analysis for which detailed morphological and morphogenetic data, especially on the type
species, are available, namely A/lotricha, Coniculostomum, Cyrtohymena, Gonostomum,
Histriculus, Notohymena, Oxytricha, Steinia, Sterkie/la, Stylonychia, Tachysoma, Uro-
soma, Urosomoida. Furthermore, I omitted oxytrichids which do not have the typical 18-
frontal-ventral-transverse cirral pattern (for example, Gastrostyla, Laurentiel/a, Onycho-
dromus, Paraurostyla, Pattersoniella, Territricha). Consequently, my proposal is incom-
plete and will change more or less significantly as further data are included (Fig. 25a, b).
Characters and character states. The characters and character states used are sum-
marised in Table 5 and their distribution in the taxa is given in Table 6. For a general de-
scription of oxytrichids and terminology, see chapter I and especially Figures 6a, b.
Character 1: Shape of the adoral zone of membranelles (Fig. 6a, 19c). The adoral
zone is formed like a question mark in most oxytrichids, as in the majority ofhypotrichs,
indicating that this configuration is plesiomorphic (Fig. 6a, 20a, 2Ic). Only in Urosoma
and Gonostomum is the middle portion straight and extends along the left margin of the
cell, causing the proximal part of the zone to became abruptly bent towards the centre of
the body (Fig. 19c, 20c, 2Id). Obviously, this is a rare and thus derived configuration.
Character 2: Relative length of the adoral zone of membranelles. The ratio of the
length of the adoral zone (exactly, the distance between anterior end of the cell and the
proximal end of the adoral zone) to the length of the cell is possibly of cladistic signifi-
cance in oxytrichid hypotrichs. It is less than 40 % (usually 25-35 %) in taxa with a flexi-
ble body, and in most other hypotrichs (except euplotids), indicating that such values rep-
resent the plesiomorphic state. Exceptions are rare, for example, Cyrtohymena quadrinu-
 PHYLOGENY 57

111/2-"
"-VI!4
V1/4~
_-VI/3
V1/3~
_ -IV/3 - _ _ ..... 11112

a b c '."

Fig. 19a-j Schematic illustrations of some charac-

ters and character states used in the cladistic analysis
(from BERGER & FOISSNER 1997). a, b) Character 6,
arrangement of frontoventral cirri in Urosoma (a)
and in some taxa (for example, Hislriculus hislrio)
with rigid body (b). c) Character 7, apomorphic posi-
tion of postoral ventral cirri (arrowheads) in Gona-
stomum. d) Characters 8 and 9, reduced number of
pretransverse ventral and transverse cirri (arrow) in
Urosomoida agiliformis. e-j) Characters 1 and 3,
shape of adoral zone of membranelles and arrange-
ment ofparoral and endoral. For details, see text and Figure 6a. e) Oxylricha pattern. f) Notohymena pattern.
g) Cyrtohymena pattern. b) Gonostomum pattern. i) Sty/onychia pattern. j) Steinia pattern. AZM = adoral zone
of mernbranelles, e = endoral, p = paroral, I-VI = primordia (anlagen) I-VI; 1-4 = numbering of cirri within a
primordium.

c/eata and Gonostomum spp., whose adoral zone is 40-50 % of body length, as in all
taxa with a rigid body.
A correlation between the relative length of the zone and the mode of nutrition is nei-
ther known nor apparent. However, such a correlation possibly exists with the absolute
size of the buccal cavity; usually, species with a small cavity (for example, Oxytricha
setigera, Tachysoma pellionellum) feed on bacteria and small protists, while those with a
large buccal cavity are rapacious carnivores (for example, Sterkiella cavicoia, Cyrtohy-
mena spp.).
58 GENERAL SECTION

FC----:--

.,
,
i

,,
,,
,"I
, •,
,,
t
,p
,
I
..,,
I ,
t
:rc
.
I
PTVC----. ,
•
RMR-·. .. ..-LMR
c d
 PHYLOGENY 59

\
FVC
f\

Fig. 20e, f Characters 1 and 3, shape of adoral zone of membranelles and arrangement of paroral and endoral
in oxytrichids after protargol impregnation (from BERGER & FOISSNER 1997). Bars = 20 ~m. e) Cyrtohymena
pattern in Cyrtohymena primicirrata. The distal portion of the paroral is semicircularly curved; the straight an-
terior half of the endoral extends diagonally on the bottom of the buccal cavity, while its posterior half is
curved and at the right buccal wall, thus intersecting optically with the posterior half of the paroral. I) Steinia
pattern in Steinia sphagnico/a. The sinusoidal endoral is fragmented (arrows) and does not intersect with the
curved paroral. The specimen shown has a surplus frontal cirrus (arrowhead). Explanation of original labels:
BC = buccal cirrus, E = endoral, FC = frontal cirri, FVC = frontoventral cirri, LMR = left row of marginal
cirri, MA = macronuclear nodule, P = paroral.

~ Fig. 20a-d Characters 1 and 3, shape of adoral zone of membranelles and arrangement of paroral and endoral
in oxytrichids after protargol impregnation (from BERGER & FOISSNER 1997). Bars = 20 ~m. a) Plesiomorphic
Oxytricha pattern in Sterkiella histriomuscorum. The paroral and endoral are slightly curved and intersect opti-
cally (arrow). The adoral zone is formed like a question mark as in all derived states, except for the Gono-
stomum pattern (c). b-d) Derived states (see also Fig. 20e, I). b) Notohymena pattern in Notohymena rubes-
cens. Arrow marks hooked distal end of paroral. c) Gonostomum pattern in Gonostomum affine. The proximal
part of the adoral zone is abruptly bent towards the body centre (arrow). The paroral consists of few, widely
spaced cilia and extends beyond the anterior end of the endoral. d) Sty/onychia pattern in Sty/onychia pustu-
lata. The paroral and endoral are straight or slightly curved and arranged side by side. Explanation of original
labels: AZM = adoral zone of membranelles, BC = buccal cirrus, E = endoral, FC = frontal cirri, FVC = fron-
toventral cirri, LMR = left row of marginal cirri, MA = macronucIear nodule, P = paroral, PTVC = pretrans-
verse ventral cirri, RMR = right row of marginal cirri, TC = transverse cirri.
 0\
o

o
~
~
en

B
~

Fig.21a-c Characters 1 and 3, oral apparatus of oxytrichids in the scanning electron microscope (from BERGER & FOlsSNER 1997). Bars = 10 11m. a, b) Plesiomor-
phic Oxytricha pattern in Oxytricha granulifera and Sterkiella cavicola. The slightly curved paroral, which borders the rather flat and narrow buccal cavity, inserts
in a narrow furrow of the buccal lip (arrowheads). c) Cyrtohymena pattern in Cyrtohymena quadrlnuc1eata. Note semicircularly curved distal portion of paroral
and straight anterior part of endoral extending diagonally on the deep and wide buccal cavity. Explanation of original labels: AZM = adoral zone of membranelles,
BC =buccal cirrus, E = endoral, FC = frontal cirri, LMR = left row of marginal cirri, P = paroral.
 PHYLOGENY 61

Fig. 2111-f Characters 1 and 3, oral apparatus of oxytrichids in the scanning electron microscope (from
BERGER & FOISSNER 1997). Bars = 10 J-lm. d, e) Gonostomum pattern in Gonostomum affine. The paroral con-
sists of few, widely spaced cilia. The endoral, which lies in the very small buccal cavity, is not recognisable.
Note the peculiar shape of the adoral zone. Arrowhead marks a food particle. 1) Steinia pattern in Steinia
sphagnicola. The paroral is slightly curved, the endoral is fragmented (arrows) and extends on the bottom of
the deep and moderately wide buccal cavity. Explanation of original labels: AZM = adoral zone of mem-
brane\les, BC = buccal cirrus, FC = frontal cirri, FVC = frontoventra1 cirri, P = paroral.

Character 3: Paroral, endoral, and buccal cavity (Fig. 19e-j, 20a-f, 21a-t). The
shape of the paroml and endoml and their armngement are very important for distinguish-
ing oxytrichid genem (FOISSNER 1989). The six known patterns of chamcter 3 are de-
scribed in the following pamgraphs.
Oxytricha pattern (Fig. 1ge, 20a, 21a, b): The paroml and endoml are slightly curved
and optically intersect at about the middle. The buccal cavity is usually mther flat and
narrow. This pattern prevails and occurs, for example, in Oxytricha, Tachysoma, Uroso-
moida, Sterkiella, and Histriculus, but also in many taxa outside the oxytrichids, like
Urostyla, Holosticha, and amphisiellids (EIGNER & FOISSNER 1994, ForsSNER et at. 1991).
It is thus considered to be the plesiomorphic state. I assume that all other patterns, except
that of Steinia, which very likely evolved from the Stylonychia pattern, originated directly
62 GENERAL SECTION

from the Oxytricha state, but very likely at different times and comparatively late, as indi-
cated by the rather great homogeneity of the group. The assumption of a direct evolution
provides a more parsimonious cladogram than a series of transformations. As concerns
the time scale, HAECKEL'S ontogenetic principle can be applied. All oxytricbids and re-
lated hypotrichs have an identical origin of the undulating membranes, namely they are
formed in parallel rows right of the oral primordium. Then, when the oral apparatus in-
vaginates in late dividers, an Oxytricha-like pattern becomes recognisable for a short
time. The specific patterns described below originate only in very late dividers and early
post dividers, and are obviously produced by the particular shaping of the buccal cortex.
Gonostomum pattern (Fig. 19c,h, 20c, 21 d, e): The paroral consists of few, widely
spaced cilia and extends far beyond the anterior end of the endoral, which is of usual
structure. The buccal cavity is very small and flat. This pattern occurs in Gonostomum
and Urosoma, and very likely evolved concomitantly with the peculiar shape of their ado-
ral zone (character 1).
Notohymena pattern (Fig. 19f, 20b): The Notohymena pattern differs from the Oxy-
tricha state in that the paroral, which is often more distinctly curved, has a hooked distal
end bent anteriorly and ventrally. The buccal cavity is moderately wide and rather deep.
This pattern occurs only in Notohymena.
Cyrtohymena pattern (Fig. 199, 20e, 21c): This pattern is very prominent, but has of-
ten been confused, for example, by I<AHL (1932), with the Steinia pattern, because both
taxa have a very deep and wide buccal cavity. The cavity is bordered by a long paroral
with a semicircularly curved distal portion. The straight anterior portion of the endoral
extends diagonally on the bottom of the buccal cavity, while its posterior half is curved
and at the right buccal wall, thus optically intersecting with the posterior portion of the
paroral. Consequently, the anterior parts of the membranes form a figure like an archer's
bow. This configuration is restricted to Cyrtohymena.
Stylonychia pattern (Fig. 19i, 20d, 148i): The paroral and the endoral are straight or
very slightly curved and arranged almost in parallel. The buccal cavity is wide and flat.
This pattern also occurs in Coniculostomum. A very similar configuration is described for
Onychodromopsis flexilis sensu PETZ & FOIssNER (1996; here Allotricha antarctica). This
must be explained as parallelism, because many other significant characters assign Allo-
tricha as a sister-group of Oxytricha (Fig. 2Sa, b).
Steinia pattern (Fig. 19j, 20f, 21t): Steinia has the most complex pattern, not only in
oxytrichids but also as compared with the hypotrichs in general. The sinusoidal endoral is
fragmented in the middle part and does not intersect with the curved paroral. At the ante-
rior end of the buccal cavity, close to the left anterior margin of the endoral there is a
unique cavity of unknown function extending to the dorsal surface (Fig. 22b). In life, the
Steinia pattern resembles the Cyrtohymena configuration because of the large and bright,
anteriorly semicircularly excavated buccal field.
There is some evidence that further patterns exist, especially within Oxytricha, which
is very likely para- or polyphyletic (for example, Oxytricha siseris, Fig. 58d, g, h). Fur-
 PHYLOGENY 63

•t
., a

Fig. 22a--d Important cladistic characters in

oxytrichids (from BERGER & FOISSNER 1997). a)
Ventral ciliature in a late divider of Sty/onychia
pustulata after protargol impregnation. The
frontal-ventral-transverse cirri of oxytrichids
originate from six longitudinal anlagen (arrows)
segregating 1, 3, 3, 3, 4, and 4 cirri. A dorso-
marginal kinety (arrowhead) originates near the
anterior end of the opisthe's right marginal row
primordium. Bar = 20 11m. b) Steinia sphagni-
cola from life showing the extraordinary pit (ar-
row) in the anterior region of the buccal cavity
(character state 3-5). Bar = SO 11m. c) Character
IS, cortical granules (arrows). The example
shows Notohymena rubescens, whose granules
are about 1 11m across, spherical and reddish.
Most types of cortical granules can be seen in
live specimens only. d) Character 11, confluent
marginal rows (arrow) in Histricu/us histrio.
Bar = 10 11m. LMR = left row of marginal cirri,
OM = opisthe's primordia of marginal rows,
PM = proter's primordia of marginal rows, PTVC = pretransverse ventral cirrus, RMR = right row of marginal
cirri, TC = transverse cirri.
64 GENERAL SECTION

Table 4 Origin of the frontal-ventral-transverse primordia in several oxytrichids (from BERGER & FOISSNER
1997)

Species (Reference)6 Primordium'

II III IV V VI
Proter
Urosomoida agiliformis (1) urn III2 IIII2 IVI3 new new
Urosomoida agilir (2) urn III2 IIII2 IV/3 new new
Cyrtohymena muscorum (3) urn III2 IIII2 IVI3 new new
Notohymena rubescens (4) urn lI12 III12 IV/3 new new
Oxytricha granulifera (5) urn lII2 and IIII2 IV/3 Vl4 V/3
part ofIIo
Oxytricha longer (6) urn III2 IIII2 IVI3 V/4 VI3
Allotricha antarctica (12) urn III2 IIII3 IVI3 V/4 VI3
Sterkiella histrio- urn III2 and IIII2 IV/3 IV/3 new
muscorum (7) part of or part
110 ofVoor VI03
Sty/onychia pustu/ata (8) urn III2 IIII2 IV/3 IV/3 IVI3
Sty/onychia bifaria (8) urn I1I2 IIII2 IV/3 IVl3 IVI3
Steinia sphagnico/a (9) urn W2 III12 IVI3 V/4 V/4
Sty/onychia myti/us (10) urn OP III12 IV/3 V/4 V/4
ofo
Coniculostomum urn OP IIII2 IV/3 V/40r V/4
monilata (11) ofo new?
Opisthe
Urosomoida agiliformis (1) OP OP OP IV12 V/4 VI3
Urosomoida agilir (2) ? ? ? ? ? ?
Cyrtohymena muscorum (3) OP OP OPand OPand V/3 VI3
IVI2? V/4?
Notohymena rubescens (4) OP OP OP IV12 V/4 V/3
Oxytricha granu/ifera (5) OP OP OP IV12 V/4 VI3
Oxytricha longer (6) OP OP OP IV12 V/4 V/3
Allotricha antarctica (12) OP OP OP IV12 V/4 V/3
Sterkiella histriomuscorum (7) OP OP OP IV12 Vl44 VW
Sty/onychia pustu/ata (8) OP OP OP IV12 Vl4 V/4
Sty/onychia bifaria (8) OP OP OP IV12 V/4 Vl4
Steinia sphagnico/a (9) OP OP OP IV12 V/4 V/4
Sty/onychia myti/us (10) OP OP OP V/4 V/4 V/4
Coniculostomum OP OP OP V/4 V/40r V/4
monilata (II) new?

, Abbreviations: 0 = opisthe, OP = oral primordium, urn = undulating membranes, Io-Vlo = primordia I-VI of
opisthe, Ip-VIp = primordia I-VI of proter.
2 The origin was ascertained by BERGER et a1. (1985a) from Figure 3 and 4 in BulTKAMP (1975).

3 Possibly originates from cirrus IVI3.

4 In the type species, Sterkiella cavico/a, primordia Vo and VIo originate likely de novo (W. FOISSNER, unpubl.
data).
s Populations 3 and 4 in GANNER et a1. (l987b).
6 1= FOISSNER and ADAM (1983a), 2 = BUITKAMP (1975), 3 = Voss (1991b), 4 = Voss (1991a), 5 = FOISSNER &
ADAM (1983b), 6 = GANNER et aI. (1987b), 7 = BERGER et aI. (1985a), 8 = WIRNSBERGER et aI. (1985a), 9 =
Voss & FOISSNER (1996), 10 = WIRNSBERGER et aI. (1986), II = KAMRA & SAPRA (1990), 12 = PE1Z & FOISS·
NER (1996).
 PHYLOGENY 65

Table 5 Characters, character states, and coding used for the construction of the c1adograms shown in Figures
25a, b (from BERGER & FOISSNER 1997). Character state matrix, see Table 6. Definition ofplesiomorphic and
apomorphic states refer only to the traditionally constructed c1adogram (Fig. 25a). See text for detailed discus-
sion.

No. Character Character states

Plesiomorphic Apomorphic
Shape of adoral zone formed like a question Gonostomum pattern (coded I)
of membranelles mark (coded 0)
2 Relative length of adoral <40 %, usually 25-35 %, >40 %, usually 45-50 %, of body
zone ofmernbranelles of body length (coded 0) length (coded 1)
3 Arrangement of paroral Oxytricha pattern Gonostomum pattern (coded I)
and endoral (coded 0) Notohymena pattern (coded 2)
Cyrtohymena pattern (coded 3)
Stylonychia pattern (coded 4)
Steinia pattern (coded 5)
4 Frontal cirri'
5 Buccal cirrus'
6 Arrangement of frontoventral V-shaped (coded 0) longitudinal (coded 1)
cirri
7 Position of postoral ventral cirri behind adoral zone (coded 0) right of adoral zone (coded 1)
8 Number of pretransverse ventral 2 (coded 1) 1 (coded 1)
cirri
9 Number of transverse cirri usually 5, rarely 4 (coded 0) less than 4 (coded 1)
10 Number of marginal rows 2 (coded 0) more than 2 (coded 1)
11 Distance between posterior present (coded 0) absent (coded 1)
ends of marginal rows
12 Number of dorsal kineties 5 or 6 (coded 0) 4 (coded 1),
3 (coded 2)
>6 (coded 3)
13 Caudal cirri present (coded 0) absent (coded 1)
14 Body flexible (coded 0) rigid (coded 1)
15 Cortical granules present (coded 0) absent (coded 1)
16 Primary primordia absent (coded 0) present (coded 1)
17 Origin ofproter's primordia from frontoventral new (coded I)
V and VI cirrus IV/3 (coded 0) primordium V from cirrus V/4 and
primordium VI from cirrus V/3
(coded 2)
from cirrus V/4 (coded 3)
18 Origin ofopisthe's from postoral ventral cirri new (coded 1)
primordia V and VI V/3 and/or V/4 (coded 0)
19 Postoral ventral cirrus V/3 not involved in primordia involved in primordia formation
formation (coded 0) (coded 1)
20 Origin of opisthe's cirrus IVl2 (coded 0) cirrus V/4 (coded 1)
primordium IV
21 Origin ofproter's primordium II cirrus II12 (coded 0) oral primordium (coded I)
66 GENERAL SECTION

Table 5 Continued

No. Character Character states

Plesiomorphic Apomorphic

22 Origin of right new formation of 1 row, new formation of2 or more rows, no
marginal row(s) no parental marginal cirri parental marginal cirri retained
retained (coded 0) (coded 1)
new formation of 1 row, parental
marginal cirri retained (coded 2)
23 Morphogenesis of dorsal kineties Oxytricha pattern (coded 0) Urosomoida pattern (coded 1)
Gonostomum pattern (coded 2)
Tachysoma pattern (coded 3)
Conicu/ostomum pattern (coded 4)

1 Not used in cladistic analysis.

thennore, the differences mentioned above probably concern not only the shape of the
membranes but also their fine structure (FERNANDEZ-LEBORANS 1985, FOISSNER 1995b).
Character 4: Frontal cirri (Fig. 6a). This group invariably consists of three cirri in
18-cirri oxytrichids, VI, IV3, and IIV3. Thus, it cannot be used as phylogenetic marker.
Character 5: Buccal cirrus (Fig. 6a). While its position along the paroral has no cla-
distic significance, its morphogenetic activity is a valuable apomorphy for some genera
(see character 21 ).
Character 6: Frontoventral cirri. The V-shaped (hook-shaped) arrangement shown
in Figure 6a is widespread and thus undoubtedly the plesiomorphic state. In Urosoma cir-
rus III12 is in front of the other three cirri, which thus fonn a longitudinal row (Fig. 19a).
No other genus-specific patterns have been distinguished. However, cirrus III12 shows a
tendency to be placed more posteriorly in species having a rigid body, for example, in
StylofTJ'chia mytNus, Steinia spp., and Histriculus histrio (Fig. 19b). Furthennore, the
frontoventral cirri are postoral in Oxytricha setigera and related species.
Character 7: Postoral ventral cirri. These three cirri fonn a narrow group close to
the buccal vertex in Oxytricha, Allotricha, Urosomoida, Urosoma, Cyrtohymena, Noto-
hymena, and Tachysoma (Fig. 2Ia). By contrast, cirrus V/3 (posterior postoral ventral
cirrus) is distinctly separated from the other two cirri in some taxa with a rigid body, for
example, Sterkiella and Steinia (Fig. 6a). However, transitions exist, and thus this charac-
ter is of cladistic significance only in Gonostomum, where morphogenetic studies (SoNG
1990, HEMBERGER 1982) revealed that the postoral ventral cirri are displaced right of the
proximal portion of the adoral zone of membranelIes (Fig. 19c).
Character 8: Pretransverse ventral cirri (Fig. 6a). Two such cirri are common in
oxytrichids and also occur in holostichids. Thus, this is very likely the plesiomorphic
state. The loss of one of these cirri, for example, in Urosomoida, is considered an autapo-
morphy (Fig. 19d).
 PHYLOGENY 67

Table 6 Distribution of characters in the IS-cirri oxytrichids analysed cladistically (from BERGER & FOISSNER
1997). The holostichids are used as outgroup. Coding, see Table 5. - = not used; ? = not-applicable

Coniculostomum o14 - - 00001 o 3 0 1 1 o 3 0 0 1 124

Cyrtohymena 003 - - o0 0 0 0 o0 0 0 0 o1 0 1 0 000
Gonostomum 1 1 1- - o 1 0 1 0 o 2 000 1 ? ? 1 ? 002
Histricu/us o10 - - o 0 0 0 0 10111 o ? 0 0 ? 000
Notohymena 002 - - o 0 000 o 0 0 0 0 01010 000
Allotricha o0 0 - - o 0 001 o 0 0 0 0 02010 010
Oxytricha o0 0 - - o 0 0 0 0 o 0 0 0 0 02010 000
Steinia o 15- - o 0 0 0 0 000 1 o 3 0 0 0 000
Sterkie/la o1 0 - - o 0 0 0 0 000 1 o ? 100 000
Sty/onychia mytilus o1 4 - - o 0 0 0 0 000 1 o 3 001 100
Sty/onychia hi/aria o1 4 - - o 0 0 0 0 o 0 0 1 1 00000 000
Tachysoma o0 0 - - 00000 o 0 1 0 0 1 ? ? 1 ? 003
Urosoma I 0 1- - 1 000 0 o 1 000 1 ? ? 1 ? o0 1
Urosomoida o0 0 - - 00110 o 100 0 o 1 0 1 0 001
holostichids o0 0 - - ? 000 0 o ? ? 0 0 o ? ? 0 ? 002

Character 9: Transverse cirri (Fig. 6a). The basic pattern comprises five cirri,
namely III 1, lIllI, lVII, VII, and VIII. The loss of cirrus IIII undoubtedly occurred inde-
pendently in different taxa because species with four transverse cirri are found in differ-
ent genera, for example, in Cyrtohymena, Oxytricha, Sterkiella. Thus, only a reduction of
the number to ::;3 is used as generic autapomorphy. The transverse cirri are often dis-
tinctly enlarged as compared with the pretransverse ventral cirri, for example, in Histricu-
Ius histrio (Fig. 22d) and/or separated into two distinct groups, for example, in Stylony-
chia mytilus, Coniculostomum spp., and Steinia sphagnicola (Fig. I48h, I68e, 173c).
Characters 10, 11: Number and arrangement of marginal cirral rows (Fig. 6a).
Most oxytrichids have one left and one right row of marginal cirri, suggesting this to be
the plesiomorphic state. Consequently, an increased number is the apomorphic configura-
tion which very likely evolved, however, at least twice independently, namely in taxa
with a flexible (Allotricha, Parurosoma; Fig. 9Ia, c, I46a) and a rigid body (Coniculo-
stomum, Pleurotricha; Fig. I68e, I88e).
Usually, the marginal rows are more or less distinctly separated posteriorly, that is,
not confluent; the gap is often difficult to recognise because it is seemingly occupied by
the caudal cirri, which, however, insert on the dorsal surface (Fig. 6b). Only in Histricu-
Ius are the marginal rows in fact confluent, suggesting this as derived condition (Fig.
22d).
Character 12: Number of dorsal kineties (Fig. 6b). Most oxytrichids have five (in-
cluding one dorsomarginal row) or six (including two dorsomarginal rows) dorsal kine-
ties. Lower or higher numbers are considered to be derived states.
Character 13: Caudal cirri (Fig. 6b). Since caudal cirri are widespread in oxytri-
chids, their presence is considered as plesiomorphy (WIRNSBERGER et al. 1986). Conse-
68 GENERAL SECTION

quently, the absence of caudal cirri in, for example, Tachysoma and Histricu/us is an apo-
morphy which was, however, obtained independently as suggested by the cladogram (Fig.
25a, b).
Character 14: Flexibility of body. Suppleness is typical for many oxytrichids (for ex-
ample, Oxytricha, Tachysoma, Notohymena, Urosoma, Paraurostyla) and also for many
other hypotrichs, indicating that this is the ancestral state. The flexibility of the body is of
central importance in the cladogram because it provides a rather parsimonious and rea-
sonable tree uniting taxa with considerable morphogenetic similarities, which are, how-
ever, plesiomorphic.
Character 15: Cortical granules (Fig. 22c). They are widespread in hypotrichs and
at least one species with cortical granules is known in all flexible oxytrichid genera, sug-
gesting that cortical granules are plesiomorphic. Surprisingly, cortical granules are en-
tirely lacking in rigid oxytrichids. Thus, I consider this character as a useful phylogenetic
marker, very likely correlated in some way with the cortical fme structure. Obviously, the
granules were lost in the last common ancestor of the rigid oxytrichids.
Character 16: Primary primordia (Fig. 23b). Urosoma and Gonostomum develop
long cirral streaks during early morphogenesis (FOISSNER 1983a, SONG 1990; Fig. 120e,
122i, 1271, m). Later, these so-called primary primordia divide transversely to form sec-
ondary primordia, that is, the usual oxytrichid cirral anlagen in proter and opisthe. This
type of primordia formation has also been described for Tachysoma pel/ionellum (Fig.
1351, m), but not for T. terricola (HEMBERGER 1982), suggesting either poly- or paraphyly
of Tachysoma, or independent evolution of primary primordia in the Gonostomum-Uro-
soma group and Tachysoma, respectively. All other oxytrichids develop at least some cir-
ral streaks independently in proter and opisthe. Thus, separated cirral primordia are con-
sidered to be the plesiomorphic character state (Fig. 23a).
Character 17: Origin ofprimordia V and VI of the proter. The origin of these anla-
gen is very likely of great cladistic significance. Four states are recognisable (Table 5).
First state (Table 4 and Fig. 23a): Primordia V and VI originate from cirri located in
the anterior half of the cell, that is, from the parental frontoventral cirri III12 and/or IV/3.
This configuration occurs in Stylonychia pustulata and S. bifaria, where both anlagen
originate from the parental frontoventral cirrus IV/3. Second state (Table 4 and Fig. 23d):
In Urosomoida, Notohymena, and Cyrtohymena both primordia are formed de novo.
Third state (Table 4 and Fig. 23c): In Oxytricha granulifera, O. longa, and in Onycho-
dromopsisjlexilis sensu PETZ & FOISSNER (1996; here Allotricha antarctica) primordium
V of the proter originates from the postoral ventral cirrus V/4, and primordium VI from
the posterior postoral ventral cirrus V/3. The data on O. longa are from the popUlations 3
and 4 of Urosomoida agiliformis sensu GANNER et al. (1987b). These populations also
have a significantly higher number of pretransverse ventral and transverse cirri than the
type populations of U. agiliformis (Table 3), suggesting that they are not conspecific.
Fourth state (Table 4 and Fig. 23e, f): In Steinia, Coniculostomum, and in species of the
Stylonychia mytilus complex both primordia originate from the postoral ventral cirrus
Vl4.
 0
0
Ip Ip Ip
IIp t t IIp t
0 IIIp • o lIIp
o IVp •
• 0 o IVp •
•
VP-+
VI V IV III II
••
VIp •
.. - Io Io
Vo \ I
IVo lIP) ""0
VIo - - '
:fHH VP • Io ~
• vo.... IIafJp 5
,~'o 17 VIp IIIa C'l
0 J\ VIa.... ~
•
II I
0 o
°of 0 o
0
o o 0 o 0
00 ) 00 00
0 0
o /
\ c

Fig.23a-c Schematic illustration of characters 16-21, ontogenesis of primordia I-VI (from BERGER & FOISSNER 1997). For designation of the frontal-ventral-transverse
cirri, see Figure 6a. Cirri involved in primordia fonnation are symbolised by black dots. The direction of the arrows has no meaning. Marginal rows omitted. For details,
that is, definition of states, see characters 16-21 and Table 5 (see also Fig. 23d-t). a) Stylonychiapustulata and S. b{farla. b) Primary primordia. This apomorphic type
of anlagen fonnation occurs in Gonostomum, Urosoma, and Tachysoma pellionellum. c) Oxytricha and Allotricha. AZM = adoral zone of membrane lies, OP = oral pri-
mordium, Ia-VIo = primordia I-VI of opisthe, Ip-VIp =primordia I-VI of proter. 0'\
\0
 -.l
0

0 o~
Ip
0 IIp
Ip
t
~
IIp t

0 IIIp 0 IlIp ( 0 III, 0

o IVp • o IVp •
o IVp •

vp •
•
,
• //\
VIP,
\ / / \ I
• IU
• \U a
IVa IIp
Vp Vp ~
Va tT'l
VIP~ lIla OP IVa
lIra OP Va V~~ IIa OP s:
VIa Va lIla
O'j; l"
VIa VIa en
• .H~ )\ 0
.'~ 0
tT'l
n
....,
• 0
-
1/ Z
0
°of °of 0
o 0 o 0 o 0
00 ) 00 ) 00
0 \ 0 \ 0

Fig. 23d-f Schematic illustration of characters 16-21, ontogenesis of primordia I-VI (from BERGER & FOlsSNER 1997). For designation of the frontal-ventral-transverse
cirri, see Figure 6a. Cirri involved in primordia formation are symbolised by black dots. The direction of the arrows has no meaning. Marginal rows omitted. For details,
that is, definition of states see, characters 16-21 and Table S (see also Fig. 23a-c). d) Urosomoida, Notohymena, and Cyrtohymena. In Cyrtohymena primordium V of
the opisthe originates from cirrus V/3 (see Table S). e) Steinia. f) Sty/onychia myti/us and Conicu/ostomum. AZM = adoral zone of membranelies, OP = oral primordium,
10-VIo =primordia I-VI of opisthe, IIrVIp = primordia I-VI of proter.
 PHYLOGENY 71

It is difficult to ascertain the ancestral and, respectively, the derived states. I assume
that the first state is the plesiomorphic condition, because all primordia of the proter de-
rive from cirri located in the oral half of the cell, that is, all primordia originate independ-
ently in the proter and the opisthe (Fig. 23a). This is also true for Paraurostyla (Fig.
212w, 213a; JERKA-DZIADOSZ & FRANKEL 1969, WIRNSBERGER et al. 1985b) and Gas-
trostyla (Fig. 201c; HEMBERGER 1982), which are usually considered to be the sister groups
of the 18-cirri oxytrichids (for example, WIRNSBERGER et al. 1986). However, computer
analysis assumes another arrangement.
Character 18: Origin ofprimordia V and VI of the opisthe. These primordia usually
originate from the postoral ventral cirri. Thus, new formation in Sterkiella cavicola
(BERGER & ForssNER 1997) and, possibly, also in S. histriomuscorum (Table 4) have to be
considered as a derived condition.
Character 19: Participation of cirrus VI3 in anlagen formation (Table 4 and Fig.
23). The cladistic significance of this cirrus was first recognised by WIRNSBERGER et at.
(1986). It mayor may not be involved mprimordia formation. WIRNSBERGER et al. (1986)
considered its inactivity as apomorphy and thus obtained Stylonychia and Sterkiella as a
sister group. I assume the very opposite using the hypothesis by WIRNSBERGER (1987) that
anlagen V and VI of the oxytrichids are homologous to the two posterior-most
(rightmost) and morphogenetically inactive anlagen in holostichids. Using holostichids as
outgroup, the inactivity of cirrus V/3 (Fig. 23a, e, t) is a plesiomorphy, whereas its par-
ticipation (Fig. 23c, d) is an autapomorphy, uniting all taxa with a flexible cortex. In
these taxa cirrus V/3 is usually arranged very close to the remaining postoral ventral cirri
(for example, Fig. 21a).
Character 20: Origin of primordium IV of the opisthe (Table 4 and Fig. 23). This
primordium usually originates from postoral ventral cirrus IV12, indicating that this is the
ancestral condition. In Coniculostomum and species of the Stylonychia mytilus complex
cirrus IV/2 is not involved in primordia formation, and anlage IV of the opisthe originates
from the postoral ventral cirrus V/4 (Fig. 23t). Thus, the ontogenetic inactivity of cirrus
IVI2 is considered as apomorphy.
Character 21: Origin ofprimordium II ofthe proter (Table 4 and Fig. 23). This pri-
mordium usually originates from the parental buccal cirrus, suggesting plesiomorphy.
Only in species of the Stylonychia mytilus complex and in Coniculostomum is it gener-
ated solely by the oral primordium (Fig. 23t). However, some participation of the oral
primordium in the formation of anlage II has been described in Oxytricha granulifera and
Sterkiella histriomuscorum (Table 4).
Character 22: Origin of right marginal cirral rows. Most oxytrichids have one left
and one right marginal row, both originating by simple ''within-row formation". Two or
more right marginal rows are obtained by two different strategies, namely by new forma-
tion of all marginal rows, for example, in Allotricha (Fig. 9Ie), or, as in Coniculostomum
(Fig. 169d-h), by retention of parental right marginal rows.
Character 23: Morphogenesis of the dorsal ciliature (Fig. 24). The value of the dor-
sal morphogenesis patterns for sorting hypotrichs and evaluating their phylogenetic rela-
72 GENERAL SECTION

. ..
·n
,

•· ,, .·
·· ··
• ••
0' ,
0

0
0
•• ··••• .
..,!.
0

.
0
0
, "
,,

Fig. 24a-e Schematic illustration of char-

acter 23, ontogenesis of the dorsal ciliature
in oxytrichids (from BERGER & FOISSNER
... :, ; I 1997). Arrows mark fragmentation of dor-
• • •+
,- :.
.'. :.-:.
sal kineties, arrowheads denote dorsomar-

..
ginal kineties. New dorsal kineties nar-
o
:• .. rowly, old (parental) kineties widely
spaced. Black squares are new caudal cirri,
white squares are old caudal cirri. a) The
" plesiomorphic Oxylricha pattern is charac-
. .
, : J
terised by fragmentation of dorsal kinety 3
and formation of one or two dorsomarginal
kineties. No parental ciliature is retained
after division. b) The Urosomoida pattern
evolved from the Oxylricha pattern by the
loss of fragmentation in kinety 3 and one
of the two dorsomarginal kineties. c) The
Gonoslomum pattern evolved from the
Urosomoida pattern by the loss of the dor-
somarginal kinety. d) The Tachysoma pel-
lionellum pattern possibly evolved from the Oxytricha pattern by additional fragmentation of dorsal kinety 2.
e) The Coniculostomum pattern evolved from the Oxytricha pattern by retention of some parental kineties (as-
terisks). AZM = adoral zone of membranelies.

tionships was recognised only recently (FOISSNER & ADAM 1983b, MARTIN 1982, MARTIN
et al. 1983). All oxytrichids have at least three dorsal kineties which fonn an anlage each
in the proter and the opisthe by intrakinetal proliferation of basal bodies (Fig. 24a-e).
Furthennore, dorsomarginal kineties originate from or very close to the primordium of
the right marginal row (Fig. 24a, b, d, e). However, specialisations occur so that five pat-
 PHYLOGENY 73

terns of dorsal ciliature fonnation can be distinguished in the oxytrichids cladistically

analysed.
Oxytricha pattern (type 4 in ForSSNER & ADAM 1983b; Fig. 24a). Kinety 3 fragments
into two unequal pieces, the posterior one becoming kinety 4, and one or two dorsomar-
ginal kineties are fonned. This is the most common, and thus very likely plesiomorphic
state. It is found in many Oxytricha species and in all Allotricha, Notohymena, Cyrtohy-
mena, Sterkiella, Histriculus, Stylonychia, and Steinia species investigated so far. This
type occurs in no other hypotrich group, except for Gastrostyla, Paraurostyla, and Par-
entocirrus strongly suggesting a close relationship of these taxa with the 18-cirri oxytri-
chids.
Urosomoida pattern (type 2 in ForSSNER & ADAM 1983b; Fig. 24b). Fragmentation of
kinety 3 is lost and only one dorsomarginal kinety is formed. This pattern occurs in Uro-
somoida and Urosoma and in some Oxytricha species (for example, O. lanceolata; Fig.
69d). These taxa are in different clades, indicating that the loss of the fragmentation was
independently achieved three times.
Gonostomum pattern (type 1 in FOISSNER & ADAM 1983b; Fig. 24c). New kineties are
formed only by within-row proliferation. Within the oxytrichids this strategy is confined
to Gonostomum, and I assume that it evolved from the Urosomoida pattern by the loss of
the dorsomarginal kineties. The same pattern occurs in most other hypotrichs (for exam-
ple, holostichids), however, here it is considered as plesiomorphic state with respect to
the oxytrichids.
Tachysoma pattern (Fig. 24d). This configuration is unique to Tachysoma pe/lionel-
lum, type of Tachysoma. It differs from the ancestral state in that not only dorsal kinety 3,
but also kinety 2 fragments. One dorsomarginal kinety is fonned, so that altogether six
dorsal kineties are obtained as in the ancestral state (HEMBERGER 1982). Unfortunately,
this pattern does not match that described for T. terricola (HEMBERGER 1982, 1985),
which shows the Oxytricha pattern, suggesting that Tachysoma is not monophyletic.
However, further morphogenetic data are needed before splitting.
Coniculostomum pattern (Fig. 24e). This pattern is identical to the plesiomorphic
state, except that some parental dorsomarginal kineties are retained after division. The re-
tention of a parental dorsal kinety in Parakahliella, which very likely belongs to the
Kahliellidae (BERGER & ForSSNER 1989b, BERGER et al. 1985a, EIGNER 1995), is presuma-
bly a kind of parallelism.
Generally, two trends in dorsal ciliature formation can be observed within the Oxytri-
chidae. On the one hand, a tendency to simplification (loss of fragmentation of dorsal ki-
nety 3 and/or loss of dorsomarginal kineties) in the soft-bodied taxa, and on the other, a
tendency to complication (multiple fragmentation, retention of parental kineties) in the
taxa with a rigid body. Very likely, each of the derived states evolved independently from
the plesiomorphic Oxytricha state - except for the Gonostomum pattern, which is obvi-
ously a transformation of the Urosomoida pattern - because no parsimonious tree was
obtained if, for instance, the simple Gonostomum pattern (which also occurs in the holo-
stichids) was used at the base of the cladogram. In this case it has to be assumed that dor-
74 GENERAL SECTION

somarginal kineties and fragmentation of dorsal kinety 3 evolved independently at least

twice, which is unlikely.
Cladistic analysis: I used both the traditional method (that is mental character analy-
sis; Fig. 25a) and a computer program (Fig. 25b). As expected, the results (cladograms)
are vel)' similar. Thus, fIrst the manually-generated tree is discussed, followed by a dis-
cussion of the differences to the computer-generated tree.
The autapomorphy of the I8-cirri oxytrichids (composed of the monophyla Oxytri-
chinae and Stylonychinae) is of course the characteristic number, arrangement, and origi-
nate of the 18 frontal-ventral-transverse cirri. It is unclear whether the fragmentation of
dorsal kinety 3 evolved simultaneously or somewhat earlier, because this curious charac-
ter occurs also in Paraurostyla, Gastrostyla, and Territricha. These three taxa have a
slightly to distinctly increased number of frontal-ventral-transverse cirri indicating that
they are more ''primitive'' than the 18-cirri oxytrichids. However, molecular biological
data and cladistic analysis of Onychodromus prove that surplus cirri are not always an in-
dication of a plesiomorphic state. Both characters (cirral pattern and fragmentation of
dorsal kinety) mentioned above are slightly modifIed in Urosomoida, Urosoma, and
Gonostomum.
The cladogram based on 21 characters divides the I8-cirri oxytrichids into two major
branches, termed Oxytricha branch or Oxytrichinae and Stylonychia branch or Stylony-
chinae (Fig. 258, b). The other clades are not named because, as is widely known, pre-
liminary cladistic results suffer from the same instability as phenetic or intuitively derived
classifIcations.
The Oxytrichinae. This monophylum is defined by a distinct ontogenetic autapomor-
phy, namely the participation of cirrus V/3 in primordia formation (character 19). The
Gonostomum-Urosoma-Tachysoma lineage has primary primordia (character 16) as the
sole synapomorphy. However, in Tachysoma primary primordia occur only in the type-
species, T. pellionellum, and not in T. ferricola, the second species so far morphogeneti-
cally characterised (HEMBERGER 1982). Different strategies not only in the ventral, but
also in the dorsal morphogenesis suggest that Tachysoma is an artifIcial assemblage uni-
fIed by the loss of caudal cirri, a character which vel)' likely evolved independently in
several oxytrichids, namely Tachysoma, Histriculus, Pleurotricha, and Parastylonychia.
Gonostomum is one of the most deviating I8-cirri oxytrichid. It has a peculiar oral
apparatus (characters 1,3) and an unusual position of the postoral ventral cirri, right of
the adoral zone ofmembranelles (Fig. 19c, 20c, 2Id, e). Furthermore, the decreased num-
ber of transverse cirri and the simplifIcation of dorsal ciliature formation (character state
23-2) are valuable autapomorphies. However, Gonostomum has synapomorphies (char-
acters 1, 3, 12,23) with Urosoma, supporting the sister group relationship proposed by
ForSSNER (1983a). WIRNSBERGER et al. (1986) ignored the primary primordia (character
16) and the distinctive oral apparatus (characters 1,3), but used the decreased number of
pretransverse ventral and transverse cirri as autapomorphies. Thus, they arrived at Gono-
stomum and Urosomoida as sister groups. Kahliella (BERGER & FOISSNER 1987a) and
Wallackia (BERGER & ForSSNER 1989a) have a similar oral apparatus to Gonostomum and
 PHYLOGENY 75

Urosoma, indicating a close relationship of oxytrichids and kahliellids, as recently pro-

posed by EIGNER (1995).
The Oxytricha-Allotricha-Urosomoida-Notohymena-Cyrtohymena lineage lacks an
autapomorphy, that is, its monophyly is questionable. However, two strong (ontogenetic)
apomorphies were found for the following dichotomy, namely special types of primordia
formation (character 17), confrrmed in seven species, including the types. Unfortunately
no autapomorphy could be found for Oxytricha, indicating an almost unchanged survival
of the stem species and/or misidentified states of some characters. The important cladistic
characters of Urosomoida are the loss of some pretransverse ventral and transverse cirri
(characters 8, 9) and the loss of the fragmentation of dorsal kinety 3 (character state
23-1). These reductions also occur in the Urosoma-Gonostomum group and must thus be
explained as parallelisms. I did not find a synapomorphy for the genera Notohymena and
Cyrtohymena. However, both have a valuable autapomorphy, namely a distinctive pattern
of the undulating membranes (character 3). Possibly, Cyrtohymena differs from Notohy-
mena and Urosomoida in the origin of the anlagen II-V of the opisthe (Table 4). How-
ever, data on other Cyrtohymena species are needed to evaluate the significance of this
difference.
The Stylonychinea. This clade is characterised by 3 autapomorphies, namely the long
oral apparatus (character 2), the rigid body (character 14), and the lack of cortical gran-
ules (character 15). This distinction is strongly supported by molecular biological data
which show that Oxytricha granulifera (a member of the Oxytrichinae) is distinctly sepa-
rated from Stylonychia pustulata, Onychodromus quadricornutus, and "Oxytricha nova"
(which is a Sterkiella; FOISSNER & BERGER 1998), taxa having a rigid cortex and thus be-
longing to the Stylonychia branch.
Sterkiella is well defmed by a unique, but sophisticated combination of morphologi-
cal characters (body rigid or only slightly flexible, undulating membranes bent and inter-
secting, marginal rows separated posteriorly, caudal cirri present). However, the only
autapomorphy is the new formation of primordia V and VI of the opisthe (character 18).
Character 11, confluent marginal rows (Fig. 22d), is a valuable autapomorphy of His-
trieulus. It is the last taxon analysed cladistically within the Stylonychia branch with the
plesiomorphic Oxytricha pattern of undulating membranes. However, Pleurotricha (ex-
cluded from the cladogram, because of insufficient morphogenetic data) shows the same
membrane pattern (Fig. 18ge) and also lacks caudal cirri (character 13), indicating a sis-
ter group relationship between Histrieulus and Pleurotricha.
The Stylonychia-Conieulostomum-Steinia group is characterised by parallel undulat-
ing membranes (character state 3-4). The cladogram reveals some peculiarities within this
group: 1) Stylonychia pustulata and S. hifaria are distinctly separate from the type spe-
cies, S. mytilus, suggesting that Stylonychia is not monophyletic. This is supported by
molecular biological data (SCHLEGEL & STEINBRUCK 1986).2) Steinia is very likely the sis-
ter group of S. mytilus and Conieulostomum because they share a peculiar morphogenetic
feature, namely primordia V and VI of the proter originate from cirrus V14 (character
state 17-3). This is why I assume that the Steinia pattern of the undulating membranes
 76 GENERAL SECTION

Oxytrichidae
A
Oxytrichinae Stylonychinae
A A
.~
~ §
~
§
g .-9 ~ ~ .~ "- ff
f .to! .to! ~
.~
~ !g fi,s fi
s~ III
§ 0 .f,!
,.@
'5 §-§ § ..;j
~
""'" '5
Qj
~ .~
.t:: .~
~
<:: ~ fi ~
0
s -§ ~ ~ ? .~
J:i -St; ""~
.~
·s
S ~ G r$
,!!}
f .0:::1 <:: ,!!}
~ S ;!!
" (/) (/)Q.t'~ rJ 0)

23-2
12-2 23-41_
9-1 • 22-21~
7-1 22-11 12-31.
2-1 •• 6-1
'--r--
3-2 ~ 3-31
'---
10-11
-r--
•
'---
10-11
••
2 3-11
2-11 •••• 23-1
12-1
•
•
•
3-11
1-11. • 23-3
13-1
9-1
8-1
21-1
20-1 3-51.

17-11. 17-21. 17-31.

13-1 ~.
16-1 •• 11-1 I 3-41~

I
18-11_

15-11.
14-11. _.
19-11_ 2-11

•• eighteen FVT cirri

Fig.25a Cladogram of common 18-cirri oxytrichid genera after manual character analysis (from BERGER &
FOISSNER 1997, slightly modified). Character states, see Table 5 and section on characters and character states.
Only autapomorphies (black squares) are shown. Parallelisms indicated by asterisks. Allotricha is identical
with Onychodromopsis, and Stylonychia bifaria is identical with S. vorax in BERGER & FOISSNER (1997). Frag-
mentation of a dorsal kinety evolved very likely earlier than the 18 frontal-ventral-transverse cirri pattern, that
is, it is no autapomorphy of the I8-cirri oxytrichids, as proposed previously (BERGER & FOISSNER 1997).
 PHYLOGENY 77

Oxytrichidae
r-----------__ ~A~ ____________~
Oxytrichinae Stylonychinae
~ ____________ ~A~ ____________ ~ r -______-JA~ ________ ~

I.
23-2 1

r ~
12-2 23-41
9-1 22-21

~.
7-1 22-1 12-31
2-1 •• ·6-1 • 10-1 • 10-11
'-- ,-- '--,-- '---
100 67 100
23-3
3-1 13-1 • 21-1
1-1 12-0 • 17-0. 20-1 3-5

100 9-1 • 75
16-1 8-1

100
23-11~
12-11. 3-2 3-3
13-1 •
67 17-2 18-1 11-1 3-41~
1
100 17-31 100
15-11
14-11
19-11 2-11 •

l
Fig. 25b Cladogram of common oxytrichid genera generated by the computer (from BERGER & FOISSNER 1997,
slightly modified). The analysis was conducted with the PAUP programme (SWAFFORD 1991) using successive
weighting and holostichids as out-group. The tree is the 50 % majority rule consensus of 72 trees. Length = 39
steps, RC = 0.95, Rl = 0.97. Character coding, see Table 5; character state matrix, see Table 6.

(character state 3-5) is a transfonnation of the Stylonychia pattern (see character discus-
sion). The peculiar fragmentation of the endoral in Steinia is one of the most impressive
autapomorphies within the oxytrichids (Fig. 20t). 3) Stylonychia mytilus is very likely
more closely related to Coniculostomum (KAMRA et al. 1994) than to S. pustulata because
of the identical origin of the FVT primordia (Table 4 and Fig. 23t). This is sustained by
their almost identical interphasic FVT cirral patterns (compare Fig. 149a with 168e).
78 GENERAL SECTION

Thus, the classification of Coniculostomum in the Kahliellidae (EIGNER 1995) or

Parakahliellidae (EIGNER 1997), based mainly on the retention of some parental ciliature
after division, is very likely artificial. It is highly unlikely that the distinctive, identical 18
frontal-ventral-transverse cirral pattern and morphogenesis found in Stylonychia my/ilus
and Conicu/ostomum evolved twice independently. Conversely, it is more parsimonious
to assume that parental marginal rows are retained in different monophyla because this is
a much simpler character than the 18-cirri pattern. Autapomorphies of Coniculostomum
are the moniliform macronucleus and an increased number of marginal rows and dorsal
kineties due to retention of parental cirri (characters 10, 22) and dorsomarginal kineties
after division (characters 12, 23; I<AMRA et al. 1994).
Computer analysis using successive weighting yields 72 most parsimonious trees,
which is high, and largely due to the great number of parallelisms in combination with a
rather low number of characters. However, the SO % majority consensus tree (Fig. 2Sb) is
rather similar to that constructed manually (Fig. 2Sa), especially concerning the basic
branching into the monophyla Oxytrichinae and Stylonychinae. Furthermore, the branch-
ing sequence within the Stylonychinae is identical. Only character 17 is differently used,
namely state 17-3 is considered a further apomorphy of the Stylonychinae and state 17-0
as apomorphy of the Sty/onychia pustu/atalbijaria group.
The branching sequence in the Oxytrichinae is slightly different from the hand-
generated tree, mainly due to the different use of characters 16, 17, and 23. However, this
is not surprising because I was unable to conclusively define the plesiomorphic and apo-
morphic states of these characters. The program assumed that fragmentation of dorsal ki-
nety 3 (character 23) was reduced only once, thus forming a Gonostomum-Urosoma-
Tachysoma-Urosomoida group. However, this implies that fragmentation occurred again
in Tachysoma. Conversely, I prefer the hypothesis that fragmentation of dorsal kineties
evolved only once, assuming that loss of fragmentation occurred twice, namely in Uroso-
moida and in the Gonostomum-Urosoma group. As in the Stylonychinae, character 17 is
also differently used in the Oxytrichinae, namely new formation (state 17-1) is assumed
as plesiomorphic within the oxytrichids.
Cladistic analysis in oxytrichids, and in ciliates in general, is only at the beginning.
Thus, the proposed hypothesis will change more or less significantly as new data are
added. However, molecular biological data are very likely indispensable to find the
branching of some "atypical" oxytrichids like Paraurosty/a, Gastrostyla, Territricha.

3 Classification
Many classifications of hypotrichs, including oxytrichids, have been suggested since the
last detailed revision by KAHL (1932). However, with few exceptions (for example, BOR-
ROR 1972a), most of them were established without detailed discussion of the characters
used. Furthermore, usually no attempts were made to elucidate the phylogenetic relation-
ships within the oxytrichids. Thus, a thorough comparison with the (preliminary) phylo-
 CLASSIFICATION 79

genetic system proposed in this book (see Fig. 25a, b and Table of Contents) is impossi-
ble. However, some general statements are made in the previous chapter and at the begin-
ning of the systematic section. Most of the deviating classifications are discussed in the
taxonomy part of the genus and species sections.
Several taxa (for example, Engelmanniella, Hemisincirra, Lamtostyla, Psammomi-
tra, Terricirra) are not considered in the present book, although classified by some
authors as Oxytrichidae and/or Oxytrichidae sensu lato. For an explanation of the exclu-
sion, see the "Taxa not considered" chapter at the end of the book. KAm.'s (1932) and
some more recent classification schemes of the oxytrichids are shown in Tables 7-13. I
did not change the original presentation, for example, incorrect dating and spelling of
names; in addition, authors are not included in my reference list. For a brief note on the
classification by EIGNER (1997), see Addendum 3.

Table 7 Classification of oxytrichid ciliates according to KAm. (1932)

2. Superclass Ciliophora DoFLElN, 190 I

2. Class Euciliata METcALF.
I. Subclass Infusoria LEBERMOLLER 1760
2. Order Spirotricha BOTscHu, 1889 emend.
4. Suborder Hypotricha STEIN sensu str.
1. Family Oxytrlchidae EHRENBERG, 1838 (includes all genera except that of the fami-
lies Euplotidae and Aspidiscidae)
15. Genus Kerona EHRENBERG, 1838
19. Genus Pleurotricha STEIN, 1859
20. Genus Gastrostyla ENGELMANN, 1861
22. Genus Gonostomum STERK!, 1878
23. Genus Ancystropodium FAURE-FR., 1909
24. Genus Oxytricha (BARY, 1824) EHRENBERG, 1838, emend. WRZESNJOWSKl, 1877
(1. Subgenus Oxytricha WRZESNJOWSKl, 1877; 2. Subgenus Tachysoma STOKES,
1887; 3. Subgenus Urosoma KOWAUlWSKI, 1882; 4. Subgenus Opisthotricha
KENT, 1881; 5. Subgenus Steinia DIESING, 1866; 6. Subgenus Histrio STERK!,
1878; 7. Subgenus Stylonychia EHRENBERG, 1838)
25. Genus Onychodromopsis STOKES, 1887
26. Genus Onychodromus STEIN, 1859

Table 8 Classification of oxytrichid ciliates according to BORROR (1972a)

Order Hypotrichida STEIN, 1859

l. Family Urostylidae BOTsOlLJ, 1889 (7. Genus Kerona EHRENBERG, 1835; 9. Genus Paraurostyla n. g.)
5. Family Oxytrichidae EHRENBERG, 1838 (I. Genus Oxytricha EHRENBERG, 1830; 2. Genus Ancystropo-
dium FAURE-FREMIET, 1907; 3. Genus Gastrostyla ENGELMANN, 1862; 4. Genus Gonostomum STERK!,
1878; 5. Genus Histriculus CORUSS, 1960; 6. Genus Laurentia DRAGESCo, 1966; 7. Genus 011)'Cho-
dromus STEIN, 1859; 8. Genus Parastylonychia DRAGESco, 1963; 9. Genus Pleurotricha STEIN, 1859;
10. Genus Psammomitra n. nom.; II. Genus Stylonychia EHRENBERG, 1830; 12. Genus Tachysoma
STOKES, 1887; 13. Genus Trachelochaeta SRAMEK-HuSEI<, 1954; 14. Genus Trachelostyla KAHL,
1932)
80 GENERAL SECTION

Table 9 Classification of oxytrichid ciliates according to STILLER (1974b)

Order Hypotrichida
Family Keronidae (Kerona EHRENBERG; ...)
Family Holostichidae (... ; Gonostomum STERKI; ...)
Family Oxytrichidae (Pleurotricha STEIN; Onychodromopsis STOKES; Gastrostyla ENGELMANN; Laurentia
DRAGESCO; Ancystropodium FAuRE-FREMIET, Urosoma KOWALEWSKI; Steinia DIESING; Oxytricha
EHRENBERG emend. WRZESNlOWSKI; Tachysoma STOKES; Opisthotricha KENT; Histrieulus CORUSS; Sty-
lonychia EHRENBERG; Onychodromus STEIN)

Table 10 Classification of oxytrichid ciliates according to CORUSS (1979)

Phylum Ciliophora DoFLElN, 1901

Class m. Polyhymenophora JANKOWSKI, 1967
Subclass Spirotricha BOTscHU, 1889
Order 4. Hypotrichida STEIN, 1859
Suborder 1. Stichotrichina FAuRE-FREMIET, 1961
Family Urostylidae BOTsCHLl, 1889 (... ; Paraurostyla BORROR, 1972; ...)
Family Holostichidae FAuRE-FREMIET, 1961 (... ; Gonostomum STERKI, 1878; Laurentiella
DRAGESCO & Nmffi, 1971; Parurosoma VON GELEI, 1954)
Family Keronidae DUJARDIN, 1840 (... ; Kerona EHRENBERG, 1835; ...)
Suborder 2. Sporadotrichina FAuRE-FREMIET, 1961
Family Oxytrichidae EHRENBERG, 1838 (Actinotricha COHN, 1866; Ancystropodium FAuRE-
FREMIET, 1907; Gastrostyla ENGELMANN, 1862; Histrieulus CORLJSS, 1960; Onychodro-
mus STEIN, 1859; Opisthotricha KENT, 1882; Oxytricha BORY DE ST. VINCENT, 1825;
Parahistrieulus GROLIERE, 1976; Parastylonychia DRAGESco, 1963; Pleurotricha STEIN,
1859; Steinia DIESING, 1866; Stylonychia EHRENBERG, 1830; Tachysoma STOKES, 1887;
Urosoma KOWALEWSKI, 1882)
lncertae sedis in suborder Sporadotrichina (Allotricha STERK!, 1878; Onychodromopsis
STOKES, 1887; ... )

Table 11 Classification ofoxytrichid ciliates according to JANKOWSKI (1979)

Order Hypotrichida STEIN, 1859

Suborder 1. Stichotrichina FAuRE-FREMIET, 1961
Superfamily Urostyloidae BOTsCHLI, 1889, superfam. n.
Family 1. Urostylidae BOTsCHU, 1889 (... ; Paraurostyla; ...)
Suborder 2. Sporadotrichina FAuRE-FREMIET, 1961
Superfamily 1. Oxytrichoidea superfam. n.
Family 1. Pleurotrichidae BOTsCHLI, 1889 (Pleurotricha, Allotricha, Onychodromopsis, Not-
hopleurotricha, Balladynella, Parurosoma, Laurentia, Laurentiella, Laurosticha, Onycho-
dromus)
Family 2. Oxytrichidae EHRENBERG, 1838
Subfamily 1. Oxytrichinae EHRENBERG, 1838, s. str., subfam. n. (Oxylricha, His/rio, His-
Irieulus, Sieinia, Opislhotricha, Urosoma, Stylonychia, Gastrostyla, Tricholeplus,
Parastylonychia, Tachysoma, Aclinotricha, Trachelostyla, Gonoslomum, Prosopsenus,
Plagiotricha, Pseudostrombidium, Balladinopsis, Parahistrieulus, Nothopleurolricha)
Subfamily 2. Ancystropodiinae subfam. n. (Ancystropodium)
 PARASITISM 81

Table 12 Classification of oxytrichid ciliates according to SMALL & LYNN (1985)

Phylum Ciliophora DoFLEIN, 1901

Subphylum Postciliodesmatophora GERASSIMOVA & SERAVIN, 1976
Class Spirotrichea BOTsCHLI, 1889
Subclass Stichotrichia n. subclass
Order Stichotrichida FAuRE-FREMIET, 1961
Suborder Stichotrichina FAuRE-FREMIET, 1961
Family Amphisiellidae n. fam. (... ; Onychodromopsis STOKES, 1887; Onychodro-
mus STEIN; 1859; Paraurostyla BORROR, 1972)
Family Gonostomatidae n. fam. (Gonostomum STERKI, 1878; ...)
Family Keronidae DUJARDIN, 1840 (Kerona EHRENBERG, 1835; ...)
Suborder Sporadotrichina FAuRE-FREMlIrr, 1961
Family Oxytrichidae EHRENBERG, 1838 (Ancystropodium FAuRE-FREMIET, 1907;
Gastrostyla ENGELMANN, 1862; Histriculus CORUSS, 1960; Laurentiella
DRAGESCO & NJINE, 1971; Oxytricha BORY DE ST. VINCENT, 1825; Parastylony-
chia DRAGESCO, 1963; Pleurotricha STEIN, 1859; Stylonychia EHRENBERG, 1830;
Tachysoma STOKES, 1887)
Family Trachelostylidae n. fam. (Psammomitra BORROR, 1972; Trachelostyla KAlIL,
1932; Urosoma KOWALEWSKI, 1882; Urosomoida HEMBERGER, 1981)

Table 13 Classification ofoxytrichid ciliates according to TUFFRAU (1987)

Order Hypotrichida STEIN, 1859

Suborder 3. Stichotrichina FAuRE-FREMIET, 1961, emend.
Family Holostichidae FAuRE-FREMIET, 1961 (... ; Coniculostomum NJINE, 1978; Gonostomum STERK!,
1878; Laurentiella BORROR, 1972; Onychodromopsis STOKES, 1887; Parurosoma, VON GELEI,
1954; ... )
Family Keronidae DUJARDIN, 1840 (... ; Kerona EHRENBERG, 1835; ...)
Suborder 4. Sporadotrichina FAuRE-FREMIET, 1961, emend.
Family Oxytrichidae EHRENBERG, 1838 (Actinotricha COlIN, 1866; Ancystropodium FAuRE-FREMIET,
1907; Gastrostyla ENGELMANN, 1862; Histriculus CoRLISS, 1960; Lamtostyla BUITKAMP, 1977;
Onychodromus STEIN, 1859; Opisthotricha KENT, 1882; Oxytricha BORY DE SAINT-VINCENT,
1925; Parahistriculus GROLIERE, 1976; Parastylonychia DRAGESCO, 1963; Paraurostyla BoRROR,
1972; Pleurotricha STEIN, 1859; Steinia DIESING, 1866; Stylonychia EHRENBERG, 1830; Tachy-
soma STOKES, 1887; Urosoma KOWALESKI, 1882; Urosomoida HEMBERGER, 1981)
Incertae sedis: Allotricha STERK(, 1878; ...

4 Parasitism
Sty/onychia species are sometimes attacked or infected by suctorians (DIECKMANN 1985,
ENGELMANN 1862, 1980b, GRENFELL 1886, KENT 1882, LOPEZ OCHOTERENA 1962,
FOISSNER
LOPEZ-OCHOTERENA 1965, PENARD 1920, PEREZ REYES & LOPEZ-OcHOTERENA 1963, QUEN-
NERSTEDT 1865, STEIN 1859b). STEIN (l859b) erroneously interpreted them as embryos of
the hypotrichous ciliates. The suctorian species described as parasite species of the S.
mytilus complex are Podophrya sty/onychiae, P. grelli, P. jlXa, P. sol, and Sphaerophrya
 82 GENERAL SECTION

12

Culture medium

Fig.26a Life cycle of Podophrya greW (from DIECKMANN 1985). 1) The swanner adheres to the host cell, Sty-
lonychia lemnae, with some tentacles. 2) The oval, ciliated swanner (15-20 /lm) deciliates and induces a pel-
licular invagination in the host cell to accommode the suctor. 3) The invagination remains open to the culture
medium. The swarmer rounds up, ingests the host cytoplasm by means of its tentacles, and grows to the adult
parasite (30-50 /lm). 4) By unequal, in rare cases equal divisions, new swarmers are formed. These leave the
invagination and can infect other host cells. 4') Sometimes fission takes place, but no ciliated swanners are
formed. 5) Adult parasite with many granulated inclusions. This stage can be observed only very rarely. Later
stages of this Podophrya cell could not be observed. 6) Early conjugating stage. 7) Late conjugating stage. In-
creasing the food uptake, the conjugants store granular material and grow considerably. 8, 9) The conjugants
then enter into the culture medium, remaining connected for some time at the bottom of the culture vessel be-
fore separating. 10,11) The exconjugants commence with the formation of a stalked cyst. 12) Completed cyst.
13,14) By successive multiple fission within the cyst, up to 16 infectious swarmers are formed; these cannot
be distinguished by the light microscope from the swanners of the asexual generation.

Fig. 26b-j Stylonychia lemnae. Infection by the suctorian Podophrya grelli (from DIECKMANN 1985. b-j, ~
phase contrast). b) A swarmer of P. greW has attached near the adoral zone of membranelles (arrow). c) Two
parasites in a pellicular invagination which remains open (arrow). d, e) Slightly and heavily infected host cell.
t) Infected host cell with early conjugating stages of the parasite (arrows). g, h) Infected host cells with 1 and 2
late conjugating stages. i, j) Conjugating pairs immediately before entering the culture medium.
PARASITISM 83
84 GENERAL SECTION

Fig. 26k-m The suctorian Podophrya gre/li, a parasite of Stylonychia lemnae (from DIECKMANN 1985. k-m,
phase contrast). k) Half-finished cyst. 1) Exconjugant. The cyst is nearly complete; the surplus portion of the
cell (arrow) is constricted and degenerates. m) Completed cyst.

pusilla (names of suctorians according to MATIlIES 1988). However, only infection by the
first and the second species are reliably described (DIECKMANN 1985, FOISSNER 1980b).
Podophrya greW is host-specific and only infects S. lemnae (Fig. 26a-m). In contrast,
Podophrya stylonychiae infects both S. lemnae and S. mytilus, but never other ciliate spe-
cies like Oxytricha fallax, Paruroleptus caudatus, Vorticella sp., or Epistylis alpestris,
not even when the parasite is highly abundant (Fig. 27a-h).
The infection rate is usually <10 %. However, DIECKMANN (1985) found that the in-
fection rate was higher (up to 15 %) during May and August than in November and Feb-
ruary. FOISSNER (l980b) observed an infection rate of about 5 % in fresh samples, but up
to 90 % in old infusions. Infection causes the death of the host. An infected cell loses the
ability to divide, to conjugate, and to encyst. It is, however, able to feed and sometimes
infected specimens grow up to 400-600 Jim. The parasite multiplies rapidly by the for-
mation of swarmers, some of which infect the same host. The cytoplasm is thus often
heavily vacuolised and contains many spherical inclusions. The nuclear apparatus of an
infected cell is strongly deformed, fragmented, and often contains large nucleoli. Because
of the high number of parasites (up to 30), the host cell is usually distinctly deformed and
movement is impaired. At this stage of infection the hypotrichs are no longer able to feed
(Fig. 26f). If the parasites conjugate, the host cell is further wasted away and contains
large vacuoles which appear empty. Finally, when the conjugating pairs leave the host, it
often becomes spherical, ceases to move, and dies (Fig. 26j).
ENGELMANN (1876, P 598) reported an endomacronuclear bacterium being cylindrical
or biscuit-shaped and 1.5-3.0 Jim long (see also GORTZ 1983, P 149). REES (1878) ob-
served in Oxytricha fallax infections of the macronucleus by parasitic algae, which were
 PARASITISM 85

later thought to be fungi or flagellates (BOTSCHLI 1889; see also GORTZ 1983). Obligatory
parasitism between oxytrichids and metazoans has not yet observed.

Fig. 27a-h Podophrya stylonychiae infecting species of the Stylonychia mytilus complex (from FOISSNER
1980b. a-c, g, h, from life; d-f, dry silver impregnation). a) Infected host cell. b) Adult specimen. c) Unequal
division. Arrow marks micronucleus. d-t) Silverline system of an adult specimen, a small post divider, and a
specimen which is modified to a swarmer. Arrows mark insertion sites oftentacles. g) Cyst formation. h) Com-
pleted cyst. Scales = 45 flm (b--t) and 12.5 flm (g, h).
86 GENERAL SECTION

5 Ecology, Occurrence, and Geographic Distribution

Oxytrichid ciliates live, throughout the year, in a wide range of biotopes: freshwater,
brackish water, sea, soil, or semiterrestrial biotopes (for example, FOISSNER 1987a, 1994d,
1998, FOISSNER et al. 1995a, b, PAITERSON et al. 1989). No species is symbiotic or para-
sitic on or in other organisms. However, some taxa (Oxytricha sp., Pleurotricha sp., Sty-
lonychia sp.) have been recorded from stools of man and domestic animals (WATSON
I 946a). Kerona pediculus lives on freshwater hydras and bryozoans.
Most freshwater species are typically bottom-dwellers, creeping on detritus, stones,
macrophytes and so on or on the air-water interface (for example, RICCI et al. 1991c).
However, some species can be found in the plankton of stagnant and running waters.
Pseudostrombidium planctonticum lives almost exclusively in the pelagial of lakes and
ponds. Some species have been found to be tolerant to rather high temperatures: Oxytri-
cha sp. 52°C (DINGFELDER 1962, NOLAND & GOJDICS 1967).
Many species are confined to either aquatic or terrestrial biotops. The vel)' common
Tachysoma pellionellum is often rather abundant in lentic and lotic waters, but was never
reliably recorded from terrestrial habitats. This indicates that it does not form cysts. Thus,
the numerous (more than 50) records from soils allover the world are very likely mis-
identifications. In contrast, Oxytricha granulifera is confmed to soil. Sterkiella his-
triomuscorum is a common eurytopic species, living in aquatic and terrestrial habitats.
Oxytrichids feed on various organisms: bacteria, algae, hyphae and spores of fungi,
auto- and heterotrophic flagellates, ciliates, and rotifers. Some species are cannibalistic,
that is, they feed on active and encysted specimens of their own species (GIESE & AlDEN
1938, WENZEL 1974). WENRICH (1918) observed specimens of Sty/onychia vorax impaled
on fungal filament. This was due to trying to swallow the tip of the growing filament and
comparatively rapid growth of the filament.
Many taxa are vel)' likely distributed world-wide. BLATJ'ERER & FOISSNER (1988) did
not, however, find Cyrtohymena muscorum in their 21 soil and moss samples from Aus-
tralia. In Europe this species is rather common in terrestrial habitats (FOISSNER 1987a).
Usually, I mention all the published records I know from all over the world There is,
however, no doubt that some (many?) determinations are incorrect. Thus, records which
are not substantiated by an illustration should be used with caution for biogeographical
interpretations. Certainly, many more oxytrichid species than described in this book exist
because very little is known about freshwater ciliates outside of Europe. Furthermore, W.
FOISSNER (pers. comm.) knows many, so far undescribed terrestrial species from all over
the world.
Several oxytrichids are used as indicators of water quality (Table 14). However,
some of them are little known (for example, Oxytricha chlorelligera, O. saprobia, Pleu-
rotricha grandis, Stylonychia stylomuscorum, Tachysoma bicirratum) or euryoecious
(Tachysoma pellionellum). A detailed description of the morphology and ecology of
these hypotrichs and other species of the saprobic system is given in our "ciliate atlas"
 ECOLOGY 87

(FOISSNER et al. 1991, 1992c, 1994b, 1995a, b) and in BERGER et al. (1997) and FOISSNER
& BERGER (1996).
Oxytrichids are not very common in activated sludge plants or other reactor types as
compared to some euplotids, like Aspidisca or Eupiotes (AuGUSTIN & FOISSNER 1992,
BARKER 1942, BARRlTI 1940, CARDlNALETIl & ZrrELLI 1991, DRAKIDES 1980, Ern, 1996,
HAMBURG-EISENBERG 1933, LACKEY 1927, LUNA-PABELLO et al. 1992, 1997, MAnoNI 1996,
SALVADO 1994, SALvAD6 et at. 1995, 1997).
Species found in the marine interstitial are summarised by CAREY (1992) and PATIER-
SON et al. (1989). No species is obligatorily anaerobic (FENCHEL & FINLAY 1995).

Table 14 Saprobic classification of oxytrichid ciliates (from FolSSNEIl et at. 1991, 1995a, FOISSNI!P. & BERGER.
1996; Histriculus histrio is classified according to SLAoECEK. & SLAoEtKovA 1997)1

Species S Valency SI page

0 b a p
Gastrostyla mystacea p 3 7 4 3.7 808
Gastrostyla steinii a 2 7 3 2.9 791
Histriculus histrio a 3 7 4 2.7 647
Histriculus vorax a 10 5 3.0 659
Kerona pediculus b--o 4 5 1 2 1.7 826
Oxytricha chlorelligera a 10 5 3.0 129
Oxytricha fallax a 8 4 3.0 137
Oxytricha hymenostoma p 2 8 4 3.8 150
Oxytricha saprobia a--p 6 4 3 3.4 174
Oxytricha setigera a-b 4 6 3 2.6 175
Oxytricha similis b-a 5 5 3 2.5 183
Paraurostyla weissei a 2 7 3 2.9 844
Pleurotricha grandis b 10 5 2.0 712
Rubrioxytrichaferruginea 0 7 3 4 1.3 484
Rubrioxytricha haematoplasma b-a 6 4 3 2.4 481
Steinia platystoma b-a 6 4 3 2.4 626
Sterkiella histriomuscorum a 2 6 2 3 3.0 683
Sty/onychia mytilus complex a 9 5 2.9 504
Stylonychia pustulata b 7 2 3 2.1 565
Stylonychia putrina a 2 7 3 2.9 586
Stylonychia stylomuscorum b 10 5 2.0 595
Stylonychia vorax b 10 5 2.0 591
Tachysoma bicirratum a-p 2 4 4 2 3.2 457
Tachysoma pellionellum b-a 4 4 1 2.5 433

1 S = indication of saprobity by simple letter, 0 = oligosaprobic, b = betamesosaprobic, a = atpharnesosaprobic,

p = polysaprobic, I = indicative weight of species (ranging between 1 and 5), SI = saprobic index (ranging
from 0-4 in the Iimnosaprobic area).
88 GENERAL SECTION

6 Collecting, Culturing, Observing, and Staining of

Oxytrichid Ciliates
A detailed description of these topics for all ciliates is given in FOIssNER (1991, 1993) and
FOISSNER et al. (1991).

6·1 Collecting and Culturing

The methods for collecting and culturing hypotrichous ciliates are treated only briefly
here as detailed culturing methods are provided in the species descriptions. Furthermore,
the general procedures as described, for instance by DRAGESCO & DRAGESCO-KERNEIS
(1986), FINLAY et al. (1988), FOISSNER et at. (1991), GALTSOFF et al. (1959), LEE et al.
(1985), MAYER (1981), and PROVASOLI et al. (1958) apply also to the hypotrichs.
Oxytrichids occur in terrestrial (litters, humic and mineral soil horizons), semiterres-
trial (for example, astatic puddles, mosses), freshwater (for example, ponds, lakes,
brooks, rivers, sewage treatment plants) and brackish water biotops. Only few exclusively
marine species are described.
There are two principle techniques available for collecting protozoans from waters:
either direct sampling of natural substrates, or artificial substrate sampling. Hypotrichs
can be sampled from natural substrates by collecting algae masses, mud, debris, macro-
phytes, small stones, and leaves, and by brushing off the aufwuchs from stones, twigs,
and vegetation (see for example, BERGER et al. 1997, BLATfERER 1995, FOISSNER et al.
1991, 1992b, HEuss 1976, LIEBMANN 1962). Plankton samples (mesh size ~10 J.1m) should
be fixed with saturated, aqueous mercuric chloride (formalin destroys all [?] oxytrichids)
or studied in life. Quantitative investigation of activated sludge samples can be done ac-
cording to the method by AUGUSTIN et al. (1989). For a detailed description of foam sam-
pling, see CAIRNS & lIENEBRY (1982). Samples should be collected in 0.5-1.0 I wide-
necked bottles and transported to the laboratory in a cooler. The investigation should
done within 24 h after collecting because the ciliate biocoenosis changes very rapidly.
The most effective means for collecting and culturing hypotrichs from soils and
mosses is the non-flooded petri dish method as described by FOISSNER (1987a; see also
FOISSNER 1993). Here, 10-50 g of fresh or air-dried soil or litter sample are placed in a
petri dish (10-15 cm across) and saturated, but not flooded, with distilled water. A ciliate,
flagellate, and naked amoeba fauna, often very rich, develops within a few days. Inspec-
tion of the cultures on days 2, 4, 6, 10, 14, and 20 usually suffices. Subsequent inspec-
tions reveal only few species due to the effects of ciliatostasis (LOfTENEGGER et al. 1987).
Paraholosticha and Keronopsis species usually occur after few hours, the very common
Gonostomum affine can be found also in old cultures.
Several conditions influence the outcome of the method: (i) air-dried soil often yields
more individuals and species than fresh soil, perhaps due to reduced microbiostasis; (ii)
the sample should contain ample litter and plant debris and must be spread over the bot-
 COLLECTING, OBSERVING, STAINING 89

tom of the petri dish in an at least I cm thick layer; (iii) the soil may not be flooded. Wa-
ter should be added to the sample until 5-20 ml drains off when the petri dish is tilted and
the soil is gently pressed with a fmger. This run-off contains the protozoa and can be used
for further preparations such as silver staining.
Some of the bacteriovorous oxytrichids thrive on various media (for example, diluted
lettuce and/or hay extracts, table waters [for example, Volvic], tap water) enriched with a
little dried yolk or a crushed wheat grain to promote bacterial growth. The well-known
species of the Stylonychia mytilus complex can be cultured in Pringsheim solution with
the algae Chlorogonium as food (AMMERMANN et al. 1974). Some predatory species grow
well with small ciliates (for example, species of the Tetrahymena pyriformis complex,
Glaucoma scintillans) as food. Flourishing cultures must be fed daily to prevent encyst-
ment or cannibalism on active or encysted individuals (for example, Gastrostyla steinii;
WENZEL 1974).

6·2 Observing Living Hypotrichs

Many physical and chemical methods have been described for retarding the movement of
ciliates in order to observe structural details (for literature see FOISSNER 1991). Chemical
immobilisation (for example, nickel sulfate) or physical slowing down by increasing the
viscosity of the medium (for example, methyl cellulose) are sometimes helpful. These
procedures often change the shape of the cell or cause premortal alterations of various
cell structures. The following simple method is therefore preferable. Place about 0.5 ml
of the raw sample on a slide and pick out (collect) the desired specimens with a micro-
pipette under a compound microscope with low magnification (for example, objective
4:1, ocular 10 x). If specimens are large enough, they can be picked out from a petri dish
under a dissecting microscope. Working with micro-pipettes, the diameter of which must
be adjusted to the size of the specimens, requires some training. Transfer the collected
specimens, which are now in a very small drop offluid, onto a slide. Apply small dabs of
Vaseline (Petroleum jelly) to each of the four comers ofa coverslip (Fig. 28a). Place this
coverslip on the droplet containing the ciliates. Press on the vase lined comers with a
mounted needle until ciliates are held firmly between slide and coverslip (Fig. 28b-d). As
the pressure is increased the ciliates gradually become less mobile and more transparent.
Hence, first the location of the main cell organelles (for example, nuclear and oral appa-
ratus, contractile vacuole) and then details (for example, cortical granules, micronucleus)
can easily be observed under low (100-400 x) and high (1000 x; oil immersion objec-
tive) magnification. The colour of the cortical granules and/or the cytoplasm must be
studied with well-adjusted bright field.
The shape of the cells is of course altered by this procedure. Therefore, specimens
taken directly from the raw culture with a large-bore (opening about 1 mm) pipette must
first be investigated under low magnification (100-400 x). Many species are too fragile
to withstand handling with micro-pipette and coverslip trapping without deterioration. In-
90 GENERAL SECTION

b
e

c d

Fig. 28a-f Live observation and staining of oxytrichid ciliates (from FOISSNER 1991). a-d) Preparation of
slides for observing living ciliates. e) Staining jar for 8 and 16 (back to back) slides, respectively. f) Watch-
glass for protargol procedure according to WILBERT.

vestigation with low magnification also requires some experience, but it guarantees that
the outline of undamaged cells are recorded. Video-microscopy (including a video-
printer) is very useful at this point of investigation, especially for the registration of the
swimming behaviour.
A compound microscope equipped with Normarski differential interference contrast
optics is best for discerning the arrangement of the cirri and the length of dorsal cilia in
living hypotrichs. If not available, use bright-field. The nuclear apparatus is well-
recognisable with differential interference contrast or phase-contrast. Many species that
were not observed in life cannot be identified after silver impregnation because important
characters (for example, size, shape, colour of cortical granules, colour of cytoplasm) are
not known. Note that the correct colour can only be seen with a well-adjusted bright field
illumination.

6-3 Staining Procedures

There are many methods for staining ciliates, but only the protargol silver impregnation
yields (usually) good results in oxytrichid hypotrichs. Thus, familiarity with this method
is an absolute prerequisite for the description of oxytrichids. It is thus described in great
detail in order also to give even beginners a fair chance to obtain good slides. Simple,
 COLLECTING, OBSERVING, STAINING 91

namely molecular, fonnulae are given for the chemicals used, since usually only these are
found in the catalogues of the suppliers (for example, MERCK).
Other silver impregnation methods (dry silver nitrate method, wet silver nitrate
method, silver carbonate method), detailed literature, and some general instructions are to
be found in the review by FOISSNER (1991), in FOISSNER (1993), and, for Gennan readers,
in FOISSNER et al. (1991).
Apart from silver impregnation, some other staining techniques are useful for taxo-
nomic work with ciliates, especially the Feulgen nuclear reaction and supravital staining
with methyl green-pyronin in order to reveal the nuclear apparatus and, respectively, the
extrusomes.

6·3·1 Feulgen Nuclear Reaction

Descriptions of this method are to be found, for example, in DRAGESCO & DRAGESCO-
KERNElS (1986) and LEE et al. (1985). The Feulgen reaction reveals the nuclear apparatus
very distinctively, but, because these organelles usually stain well with protargol, it is sel-
dom used for hypotrichs.

6·3·2 Supravital Staining with Methyl Green-Pyronin

This simple method was described by FOISSNER (1979d). It is an excellent technique for
revealing the mucocysts of most ciliates. Mucocysts are stained deeply and very distinc-
tively blue or red, and can be observed in various stages of explosion because the cells
are not killed instantly. The nuclear apparatus is also stained.

Procedure (after FOISSNER 1991)

1. Pick out desired ciliates with a micro-pipette and place the small drop of fluid in the
centre of a slide.

2. Add an equally sized drop of methyl green-pyronin and mix the two drops gently by
swivelling the slide.
Remarks: If ciliates were already mounted under the coverslip, add a drop of the dye at
one edge of the coverslip and pass it through the preparation with a piece of filter paper
placed at the other end of the coverslip.

3. Place a coverslip with vaselined comers on the preparation.

Remarks: Observe immediately. Cells die in the stain within 2 min. Mucocysts stain very
quickly and many can be observed at various stages of explosion. To reveal the nuclear
apparatus, cells should be fairly strongly squashed (= flattened). The preparation is tem-
porary. After 5-10 min the cytoplasm often becomes heavily stained and obscures other
details.
92 GENERAL SECTION

Reagents

1 g methyl green-pyronin (CHROMA-GESELLSCHAFf, Schmid GmbH and Co. 0-7316 KOn-

genIN.)
add 100 ml distilled water
This solution is vel)' stable and can be used for years.

6'3·3 Protargol Methods

Protargol methods are indispensable for descriptive research ofhypotrichs. The first pro-
cedures were provided by KIRBy (1945), MOSKOWITZ (1950), DRAGESCO (1962), and TUF-
FRAU (1964, 1967) and many more modifications were subsequently proposed (see FOIss-
NER 1991 for references). Here, two variations which produce good results are described.
These procedures work well with most ciliate species, but require at least 20 specimens.
A single specimen cannot usually be handled successfully. Depending on the procedure
used, protargol can reveal many cortical and internal structures, such as basal bodies, fib-
rillary systems, nuclear apparatus. The silverlines (which have no taxonomical value in
the oxytrlchids), however, never impregnate. The shape of the cells is usually well pre-
served in permanent slides, which is an advantage for the investigation, but makes photo-
graphic documentation more difficult. However, pictures as clear as those taken from wet
silver carbonate impregnations can be obtained with the WILBERT modification if the cells
are photographed prior to embedding in the albumen glycerol.

Procedure A (after FOIssNER 1991)

The quality of the slides is usually adequate but frequently not as good as with the WIL-
BERT modification. The latter demands more material and experience; inexperienced
workers may easily lose all the material. As in all protargol methods, the procedure is
rather time-consuming and complicated. A centrifuge may be used for step 2; staining
jars (Fig. 28e) are necessary for steps 6-16.

1. Fix organisms in BOUIN'S or STIEVE'S fluid for 10-30 min.

Remarks: The fixation time has little influence on the quality of the preparation within the
limits given. Ratio fixative to sample fluid should be at least 2: 1. Pour ciliates into fixa-
tive using a wide-necked flask in order to bring organisms in contact with the fixative as
quickly as possible. Both fixatives work well but may provide different results with cer-
tain organisms. STIEVE'S fluid may be supplemented with some drops of2 % osmium te-
troxide for better fixation of vel)' fragile hypotrichs, for example, Urosoma. This in-
creases the stability of the cells but usually reduces their impregnability.

2. Concentrate by centrifugation and wash organisms 3 to 4 times in distilled water.

 COLLECTING, OBSERVING, STAINING 93

Remarks: There are now 2 choices: either to continue with step 3, or to transfer the mate-
rial through 30-50-70 % alcohol to 70 % alcohol (ethanol), where it remains stable for
several years. Transfer preserved material back through the graded alcohol series into
distilled water prior to continuing with the next step. Impregnation may be slightly modi-
fied in preserved material.

3. Clean 8 slides (or less if material is very scarce) per sample. The slides must be grease-
free (clean with alcohol and flame). Insufficiently cleaned slides may cause the albumen
to detach. Mark slides on back if several samples are prepared together. Use staining jars
with 8 sections so that you can work with 16 slides simultaneously by putting them back
to back (Fig. 28e).

4. Put 1 drop each of albumen-glycerol and concentrated organisms in the centre of a

slide. Mix drops with a mounted needle and spread over the middle third.
Remarks: Use about equally sized drops of albumen-glycerol and suspended (in distilled
water) organisms to facilitate spreading. The size of the drops should be adjusted so that
the middle third of the slide is covered after spreading. Now remove sand, grains, etc.
The thickness of the albumen layer should be equal to that of the organisms. Some
thicker and thinner slides should, however, also be prepared because the thickness of the
albumen layer greatly influences the quality of the preparation. Cells may dry out and/or
shrink if the albumen layer is too thin; if it is too thick, it may detach, or the cells may be-
come impossible to study with the oil immersion objective.

5. Allow slides to dry for at least 12 h (overnight) at room temperature.

Remarks: Slides may be allowed to dry for up 24 h but no longer if quality is to be main-
tained. Oven-dried (2 h at 60°C) slides are usually also of poorer quality.

6. Place slides in a staining jar (Fig. 28e) filled with 95 % alcohol (ethanol) for
20-30 min. Place a stainingjar with protargol solution into an oven (60°C).
Remarks: Slides should not be transferred through an alcohol series into concentrated al-
cohol as this causes the albumen layer to detach! Decrease hardening time to 20 min if al-
bumen is already rather old and/or not very sticky.

7. Rehydrate slides through 70 % alcohol and 2 distilled water steps for 5 min each.

8. Place slides in 0.2 % potassium permanganate solution. Remove first slide (or pair of
slides) after 60 s and the rest at 15 s intervals. Collect slides in a staining jar filled with
distilled water.
Remarks: Bleaching is by permanganate and oxalic acid (step 9). The procedure de-
scribed above is necessary because each species has its optimum bleaching time. The se-
quence in which slides are treated should be recorded because the immersion time in ox-
alic acid must be proportional to that in the permanganate solution. The albumen layer
94 GENERAL SECTION

containing the organisms should swell slightly in the pennanganate solution and the sur-
face should become uneven. If it remains smooth, the albumen is too sticky and this could
decrease the quality of the impregnation. If the albumen swells strongly, it is possibly too
weak (old) and liable to detach. Use fresh KMn04 solution for each series.

9. Quickly transfer slides to 2.5 % oxalic acid. Remove first slide (or pair of slides) after
160 s, the others at 20 s intervals. Collect slides in a staining jar filled with distilled
water.
Remarks: Same as for step 8! Albumen layer becomes smooth in oxalic acid.

10. Wash slides 3 times for 3 min each in distilled water.

11. Place slides in warm (60°C) protargol solution and impregnate for 10-15 min at
60°C.
Remarks: Protargol solution can be used only once.

12. Remove staining jar with the slides from the oven and allow to cool for 10 min at
room temperature.
Remarks: In the meantime organise six staining jars for developing the slides: distilled
water - distilled water - fixative (sodium thiosulfate) - distilled water -70 % alcohol-
100 % alcohol (ethanol).

13. Remove the first slide from the protargol solution and drop some developer on the
layer of albumen. Move slide gently to spread developer evenly. As soon as the albumen
turns yellowish, pour off the developer, dip slide in the first 2 distilled water steps for
about 2 s each and stop development by submerging the slide in the fixative (sodium thio-
sulfate), where it can be left for 5-10 min.
Remarks: Now control impregnation with the compound microscope. The impregnation
intensity is sufficient if the infraciliature is just recognisable. The pennanent slide will be
too dark if the infraciliature is distinct at this stage of the procedure! The intensity of the
impregnation can be controlled by the concentration of the developer and the time of de-
velopment. 5-10 s usually suffice for the diluted developer! Development time increases
with bleaching time. Therefore commence development with those slides which were in
the bleaching solutions for 60 and 120 s, respectively. The thinner the albumen layer, the
quicker the development.

14. Collect slides in the fixative (sodium thiosulfate) and transfer to distilled water for
about 5 min.
Remarks: Do not wash too long; the albumen layer is very fragile and detaches rather eas-
ily!

15. Transfer slides to 70 %- 100 %-100 % alcohol for 5 min each.

 COLLECTING, OBSERVING, STAINING 95

16. Clear by two 10 min transfers through xylene.

17. Mount in synthetic neutral mounting medium.

Remarks: Do not dry slides between 16 and 17! Mounting medium should be rather vis-
cous to avoid air-bubbles being formed when solvent evaporates during drying. If air-
bubbles develop in the mounted and hardened slide, re-immerse in xylene for some days
until the coverslip drops off. Remount using a more viscous medium and remove possible
sand grains protruding from the gelatine. Usually, some air-bubbles are found immedi-
ately after mounting; these can be pushed to the edge of the coverslip with a fmger or
mounted needle. The preparation is stable.

Reagents

a) Bourn's fluid (prepare immediately before use; components can be stored)

15 parts saturated, aqueous picric acid (CJI3N30 7; preparation: add an excess of picric
crystals to, for example, 1 litre of distilled water; shake solution several times within a
week; some undissolved crystals should remain; filter before use)
5 parts formalin (HCHO; commercial concentration, about 37 %)
1 part glacial acetic acid (= concentrated acetic acid; C:zlI402)

b) STIEVE'S fluid (slightly modified; prepare immediately before use; components can be
stored)
38 ml saturated, aqueous mercuric chloride (dissolve 60 g HgCh in 1 litre of boiling dis-
tilled water)
10 ml formalin (HCHO; commercial concentration, about 37 %)
3 ml glacial acetic acid (= concentrated acetic acid; C2IL02)

c) Albumen-glycerol (2-4 month stability)

15 ml egg albumen
15 ml concentrated (98-100 %) glycerol (C2Ha03)

Pre-treatment of the egg albumen and preparation of the albumen-glycerol: separate the
white carefully from the yolk and embryo of 3 eggs (free range eggs are preferable to
those from battery chickens, whose egg white is less stable and sticky). Shake the white
by hand (do not use a mixer!) for some minutes in a narrow-mouthed 250 ml Erlenmeyer
flask until a stiff white foam is formed. Allow the flask to stand for about 1 min. Pour the
viscous rest of the egg white in a second Erlenmeyer flask and shake again until it is stiff.
Repeat until most of the egg white is either stiff or becomes watery; usually 4-6 Erlen-
meyer flasks of foam are obtained. Leave all flasks undisturbed for about 10 min. During
this time a glycerol-like fluid percolates from the foam. This fluid and the white from the
last flask are collected and used. Add an equal volume of concentrated glycerol and a
small thymol crystal (CIOH I40) for preservation of the mixture. Mix by shaking gently and
96 GENERAL SECTION

pour mixture into a small flask. Leave undisturbed for 2 weeks. A whitish slime settles at
the bottom of the flask. A "good" albumen-glycerol drags a short thread when touched
with a needle. The albumen is too thin (not sticky enough) or too old if this thread is not
formed. Fresh albumen which is too thin may be concentrated by leaving it open for some
weeks so that water can evaporate. If the albumen is too sticky, which may cause only
one side of the organisms to impregnate well, it is diluted with distilled water or old, less
sticky albumen to the appropriate consistency. The preparation of the albumen-glycerol
must be undertaken with great care because much depends on its quality. Unfortunately,
all commercial products which have been tried detach during impregnation.

d) 0.2 % potassium permanganate solution (stable for about 1 d)

0.2 g potassium permanganate (KMn04)
add 100 ml distilled water

e) 2.5 % oxalic acid solution (stable for about 1 d)

2.5 g oxalic acid (CJI204-2H20)
add 100 ml distilled water

f) 0.4 % protargol solution (stable for about 1 d)

100 ml distilled water
add 0.4 g protargol
Remarks: Use light-brown "protargol for microscopy" (for example, MERCK'S "Albu-
mosesilber fur die Mikroskopie" or "Proteinate d'Argent", Roques, Paris, France). Some
dark-brown, cheaper products do not work! Sprinkle powder on the surface of the water
and allow to dissolve without stirring; use a wide-mouthed bottle for dissolving the pro-
targo!.

g) Developer (mix in sequence indicated; sodium sulphite must be dissolved before hy-
droquinone is added)
95 ml distilled water
5 g sodium sulphite (Na2S03)
1 g hydroquinone (CJL;02)
Remarks: This recipe yields the stock solution which is stable for some weeks and should
be used undiluted for certain ciliates (step 13). Usually, however, it must be diluted with
tap water in a ratio of 1:20 to 1:50 to avoid too rapid development and one-sided impreg-
nation of the organisms. Freshly prepared developer is usually inadequate (the albumen
turns greenish instead of brownish). The developer should thus be prepared from equal
parts of fresh and old (brown) stock solutions. Take great care with the developer as its
quality contributes highly to that of the slides. If the developer has lost its activity (which
is not always indicated by a brown colour!) the silver is not, or only insufficiently re-
duced, and the organisms stain too faintly. A fresh developer should therefore be pre-
pared for each "impregnation week", and some old developer kept. Fresh developer can
 COLLECTING, OBSERVING, STAINING 97

be artificially aged by adding some sodium carbonate (Na2C03). However, better results
are obtained with air-aged solutions, that is, by a developer which has been kept uncov-
ered for some days in a wide-mouthed bottle. It first turns yellowish, then light brown
(most effective), and later dark brown and viscous (at this stage the developer has lost
most of its activity, but is still suitable for artificial ageing of fresh developer = 1: 1 mix-
ture mentioned above).

h) Fixative for impregnation (stable for several years)

25 g sodium thiosulfate (Na2S2035H20)
add 100 ml distilled water

Procedure B (after WILBERT 1975 and FOlSSNER 1991)

This modification produces excellent results, but demands much experience. Manipulate
large cells with micropipettes in a watch-glass, whereas the centrifuge is used for steps
1-4, 7, 8 if cells are smaller than about 150 !lm. The watch-glass method is used when
there are only few specimens oflarger cells; thus an attempt is worthwhile even if only 20
cells are available. The organisms are very soft after development and fixation, and are
thus easily compressed between slide and coverslip, which greatly facilitates photo-
graphic documentation.

1. Fix organisms as described in protargol procedure A.

2. Wash and, if so desired, preserve organisms as described in protargol procedure A.

Remarks: Wash cells either in the centrifuge (small species) or in a watch-glass. To
change fluids allow cells to settle on bottom of watch-glass and remove supernatant with
a micro-pipette under the dissecting microscope; concentrate cells in the centre of the
watch-glass by gentle swirling.

3. Transfer organisms with a small amount of distilled water to an at least tenfold quantity
of sodium hypochlorite (NaCIO) and bleach for about 3 min.
Remarks: This is the critical step in this modification. If bleaching is too strong or too
weak, all is lost: cells either dissolve or do not impregnate well. Systematic investigations
showed that not the bleaching time but the amount of active chloride in the sodium hypo-
chlorite and the pre-treatment of the cells (fixation method, fresh or preserved material)
are decisive for the quality of the preparation. Different species need different concentra-
tions. Unfortunately, the concentration of active chloride in the commercial products var-
ies (10-13 %) and is dependent on the age of the fluid. It is thus impossible to provide
more than only a few guidelines: 100 ml distilled water + 0.2-0.4 ml NaCIO (if product
is fresh and cells were not stored in alcohol) or 100 ml distilled water + 0.5-1.6 ml Na-
CIO (if product is older and cells were stored in alcohol). The transparency of the cells
under the dissecting microscope may serve as a further indicator: fixed, unbleached cells
98 GENERAL SECTION

appear dark and opaque, whereas accurately bleached cells are almost colourless and
rather transparent (depends, however, also on size and thickness of the cell). Thus, in-
crease the concentration of sodium hypochlorite stepwise if cells appear too dark with the
recommended concentrations. We routinely start with 3 different hypochlorite concentra-
tions if enough material is available.

4. Wash organisms at least 3 times with distilled water and finally once in the protargol
solution.
Remarks: Wash thoroughly, especially when fluids are changed with micro-pipettes, be-
cause even the slightest traces of the sodium hypochlorite disturb impregnation.

5. Transfer to 1 % protargol solution and impregnate for 10-20 min at 60 °C.

Remarks: This and the next step should be carried out in a watch-glass even for material
which is otherwise manipulated with the centrifuge. The impregnation time depends on
the kind of material and the degree of bleaching. Check the progress of impregnation
every 3-4 min under the compound microscope by picking out a few cells with the
micro-pipette under the dissecting microscope; add these to 1 drop of developer. Dilute
developer and/or interrupt development of adding a little fixative (sodium thiosulfate) if
impregnation is strong enough.

6. Remove most of the protargol solution with a micro-pipette and add some drops of de-
veloper to the remainder containing the organisms.
Remarks: Fresh, undiluted developer is usually used (but see step 5). Control develop-
ment in compound or dissecting microscope. As soon as the infraciliature becomes faintly
visible, development must be stopped by adding a few drops of sodium thiosulfate. Judg-
ing the right moment is a question of experience; the permanent slide will be too dark if
the infraciliature is very distinct at this stage of the procedure!

7. Stabilise the impregnation by 2, approximately 5-minute transfers through sodium thio-

sulfate.
Remarks: The developer need not be removed before fIXation. For small species this and
the next step can be carried out in a centrifuge. Larger species must be manipulated with
micro-pipettes because cells become very fragile and would be damaged in a centrifuge.
Cells are very soft at this stage and can thus be easily compressed and photographed.
Transfer some of the more darkly impregnated specimens with a very small amount of the
fixative onto a clean slide using a micro-pipette and cover with a coverslip. Organisms
are usually flattened by the weight of the coverslip; excess fluid my be removed from the
edge of the coverslip with a piece of filter paper.

8. Wash very thoroughly in distilled water (3 times with the centrifuge; 7-10 times in
watch-glass with micro-pipettes). Finally remove as much of the water as possible.
 COLLECTING, OBSERVING, STAINING 99

Remarks: Even the slightest traces of the fixative destroy the impregnation within a few
days or weeks.

9. Smear a moderately thick layer of albumen-glycerol on a clean slide with a finger.

Drop impregnated, washed cells on the albumised slide with a large-bore pipette (opening
about 1 mm) and dry preparation for at least 2 h.
Remarks: The cells are too fragile to be spread with a needle. With much care it is possi-
ble to orientate cells using a mounted eyelash. Commercial albumen-glycerol can be
used.

10. Harden albumen by two 10-minute transfers through concentrated alcohol (ethanol).
Remarks: This and the next step are best carried out in staining jars. The albumen layer
turns milky and opaque.

11. Clear by two 5-minute transfers through xylene.

Remarks: The albumen layer turns transparent.

12. Mount in synthetic neutral mounting medium.

Remarks: Do not dry slides between steps 11 and 12! Mounting medium should be rather
viscous to avoid air-bubbles being formed when solvent evaporates during drying. If air-
bubbles develop in the mounted and hardened slide, re-immerse in xylene for some days
until the coverslip drops off. Remount using a more viscous medium and remove possible
sand grains protruding from the albumen. Usually, some air-bubbles are found immedi-
ately after mounting; these can be pushed to the edge of the coverslip with a fmger or
mounted needle. The preparation is stable.

Reagents

If not stated otherwise, the same reagents as in the protargol procedure A are to be used
(see above).

6-4 Preparation for Scanning Electron Microscopy

Hypotrichs, and especially oxytrichids, cannot usually be identified solely by scanning

electron microscopy because only a limited number of characters is revealed. However,
scanning electron microscopy is useful in that it allows a three-dimensional view of the
object, as well as for documenting details which are difficult to reveal with other
methods. For a detailed instruction of preparation for scanning electron microscopy, see
FOISSNER (1991, 1993), FOISSNER et al. (1991), and other textbooks.
 100 GENERAL SECTION

7 Summary of New Taxa Described in this Book and of

Nomenclatural Acts
New genera: Rigidocortex (p 717); Rubrioxytricha (p 479).

New species: Allotricha antarctica (p 268); Oxytricha durhamiensis (p 228); Oxytricha

pseudofurcata (p 173).

New combinations: Australocirrus octonucleatus FOISSNER with Rigidocortex (p 718);

Hemisincirra polynucleata ForSSNER with Urosoma (p 419); Histriculus admirabilis
ForSSNER with Sterkiella (p 677); Histrio quadrinucleatus SICK with Sterkiella (p 694);
Holosticha geleii WILBERT with Oxytricha (p 232); Holosticha (parurosoma) dubium GB-
LEI with Parurosoma (p 492); Opisthotricha emarginata STOKES with Urosoma (p 409);
Opisthotricha parahalophi/a WANG with Oxytricha (p 170); Oxytricha australis ForSSNER
& O'DONOGHUE with Notohymena (p 340); Oxytricha bi/aria STOKES with Stylonychia
(p 557); Oxytricha (Steinia) candens var. aestuarii MARGALEF L6PEZ with Cyrtohymena
(p 308); Oxytricha caudata EHRENBERG with Urosoma (p 398); Oxytricha ferruginea
STEIN with Rubrioxytricha (p 484); Oxytricha gigantea HORVATH with Urosoma (p 425);
Oxytricha haematoplasma BLATTERER & FOISSNER with Rubrioxytricha (p 481); Oxytri-
cha macrostyla WRZESNIOWSKlEGO with Urosoma (p 404); Oxytricha pel/ionella var.
chi/ensis BORGER with Tachysoma (p 451); Oxytricha salmastra DRAGESCO & DRAGESC0-
KERNEls with Urosoma (p 424); Oxytricha tricirrata BUlTKAMP with Sterkiella (p 693);
Perisincirra simi/is ForSSNER with Urosoma (p 419); Pleurotricha indica AMMERMANN,
SAPRA & SCHLEGEL with Coniculostomum (p 620); Trachelochaeta gonostomoida HEM.
BERGER with Gonostomum (p 392); Urostyla polymicronucleata MEruuMAN with Parauro-
styla (p 846).

Resurrected taxa: Al/otricha STERKI (p 261); Allotricha mollis STERKI (p 262).

It is suggested to synonymise the following taxa (including supposed synonyms):

Coniculostomum bimarginata I<AMRA, SAPRA & AMMERMANN with Coniculostomum in-
dica (p 620); Gastrostyla vorax NUSSBAUM with Gastrostyla steinii (p 791); Gastrostyla
phi/ipinensis SHIBUYA with Gastrostyla steinii (p 791); Gastrostyla parasteinii SMAll &
LYNN with Gastrostyla steinii (p 791); Gonostomum algicola GElLERT with Gonostomum
affine (p 370); Histrio hyalinus VOROsvARy with Histriculus histrio (p 650); Histrio le-
mani DRAGESCO with Gastrostyla steinii (p 791); Laurentia macrostoma DRAGESCO with
Laurentiella strenua (p 753); Opisthotricha terrestris HORVATH with Sterkiella histrio-
muscorum (p 683); Oxytricha caudata STOKES with Urosoma caudata (p 398); Oxytricha
phytophaga GELEI & SZABADOS with Stylonychia hi/aria (p 562); Oxytricha histroides
GELLERT with Sterkiella histriomuscorum (p 683); Peritromus hydrarum CHARDEZ with
Kerona pediculus (p 826); Pleurotricha variabilis REuTER with Allotricha mol/is (p 268);
 TAXONOMY AND NOMENCLATURE 101

P/eurotricha tchadensis DRAGESCO with Pleurotricha /anceo/ata (p 700); Steinia dubia

GELLERT with Cyrtohymena muscorum (p 281); Steinia simplex DRAGESCO with Cyrtohy-
mena candens (p 308); Steinia macrostoma GROLIERE with Cyrtohymena inquieta (p
314); Steinia candens var. depressa GELLERT with Cyrtohymena inquieta (p 315); Steinia
ovalis REmER with Cyrtohymena inquieta (p 315); Stylonychia virgula FROMENTEL with
Stylonychia myti/us complex (p 548); Sty/onychia leahli DRAGESCO with Sty/onychia myti-
Ius complex (p 549); Stylonychia myti/us var. pusilla STERKI with Stylonychia notophora
(p 555); Sty/onychia quadrinucleata ALEKPEROV & MUSAEV with Sterkiella admirabi/is (p
677); Trichotaxis fossicola KAHL with Paraurostyla granulifera (p 875); Urosoma mac-
rostoma GELLERT with Gonostomum affine (p 370); Urosoma cienkowski KOWALEWSKIEGO
with Urosoma caudata (p 398); Urosoma acuta DRAGESCO with Urosoma macrostyla (p
404); Urosoma ambigua DRAGESCO & DRAGESCo-l<.ERNEIS with Urosoma macrostyla (p
404).

New names: Oxytricha alfredi for Oxytricha minor KAHL (p 125); Oxytricha leah/ovata
for Oxytricha (Opisthotricha) ovata KARL (p 157); Oxytricha oxymarina for Oxytricha
(Oxytricha) marina KARL (p 233); Sterkiella terricola for Histriculus tricirratus DRA-
GESCO & DRAGSCo-KERNEIS (p 682).

Nomina nuda: Gonostomum mereschkowskii LEPSI (p 368); Leptocirra tekirghiolica Tu.

COLESCU (p 492); Parurosoma granulifera BERGER (p 262); Parurosoma tekirghiolica Tu·
COLESCU (p 492); Stylonychia harbinensis FENG, Gu & ZHANG (p 599).

Species indeterminata: See end of each genus section.

B Systematic Section

Oxytrichidea EHRENBERG, 1838

1838 Oxytrichina EHRENBERG, Infusionsthierchen, p 362 (in part) - Name-bearing type: Oxytricha BORY DE
SAINT-VINCENT in lAMOURoux, BoRY DE SAINT-VINCENT & DESLONOCHAMPS, 1824.
1841 Keronidae DuJARDIN, Zoophytes, p 410 (in part; published as Keroniens).
1852 Oxytrichina E. - PERlY, Zur Kenntniss kleinster LebensConnen, p 153 (in part).
1858 Oxytrichina - CLAPAREoE & LACHMANN, Mem. Inst nato. genev., 5: 135 (in part).
1859 Oxytrichina, EHRao. - STEIN, Infusionsthierchen I, p 140 (in part).
1875 Keronina - FROMENTEL, Microzoaires, p 159 (in part).
1882 Oxytrichidae, ERR. - KENT, Manual infusoria 11, p 759 (in part).
1889 Pleurotrichina BOTsOlLl, Protozoa, p 1746.
1926 Pleurotrichidae, BOTscHU -CALKINS, Biology oCthe protozoa, p 394.
1932 Oxytrichidae EHRENBERG, 1838 -KAHL, Tierwelt Dtl., 25: 537 (in part).
1936 Pleurotrichinae BOTscHLI, 1889 - BHATIA, Protozoa: Ciliophora, p 371.
1960 Oxytrichidae EHRENBERG, 1838 - CoRUSS, Ciliated protozoa, p 169 (in part).
1972 OxytriehidaeEtlRENBERG, 1838-BoRRoR,J.Protozool.,19: 13.
1974 Oxytrichidae - S11LlER, Fauna Hung., 115: 102.
1979 Oxytrichoidea JANKOWSKI, Trudy zool. Inst., Lenigr., 86: 77.
1979 Oxytrichidae EHRENBERG, 1838 - CORLISS, Ciliated protozoa, p 311 (in part).
1983 Oxytrichidae - CURDs, GATES & ROBERTS, British and other fieshwater ciliated protozoa, p 428.
1985 Gooostomidae CuuiERsON, Diss. Abstr. int, 46: 2589.
1985 Oxytrichidae - SMALL & LYNN, Illustrated guide to the protozoa, p 458.
1987 Oxytrichidae EHRENBERG, 1838 - TUFFRAU, Annis Sci. nat, 8: 116 (in part).
1994 Oxytrichidae EHRENBERG 1838 - TUFFRAu & FLEURY, Traite de Zoologic, 2: 141 (in part).
1997 Oxytrichidae EHRENBERG, 1838 - BERGER & FOisSNER, Arch. Protistenk., 148: 145.
1997 Oxytrichidae EHRENBERG, 1838 - EIGNER, J. Euk. Microbiol., 44: 559 (in part).
1997 Parakahliellidae EIGNER, J. Euk. Microbiol., 44: 563 (in part).

C h a r act e r i sat ion: Hypotrichous ciliates with usually 18 frontal-ventral-trans-

verse cirri clustered to six distinct groups and/or at least one fragmenting (splitting) dor-
sal kinety (primarily kinety 3; fragmentation sometimes secondarily lost). In the 18-cirri
oxytrichids the frontal-ventral-transverse cirri originate from usually six longitudinal pri-
mordia segregating 1, 3, 3, 3, 4, 4 cirri.

Rem ark s: Previously we categorised this monophylum as family (BERGER & FOISSNER
1997), as also indicated by the defined ending -idae. Here I ignore the category as such
because it is of no scientific value (for details, see Ax 1995, P 18; SUDlIAUS & REHmD
1992, P 12; WESTIfEIDE & RIEGER 1996, P VII); however, to avoid inflation, I have re-
tained the name.
The list of synonyms includes most classifications of "oxytrichids" proposed since
the establishment of this group by EHRENBERG. "In part" means that genera are included
which are not considered in the present book. KAHL's (1932) classification and some later
proposals are shown in detail in Tables 7-13. It would be to tedious to compare every
 Oxytrichidae 103

previous classification with my proposal because, with few exceptions (for example, BOR-
ROR 1972a), most were created without detailed discussion of the characters used, espe-
cially those published after KAHL (1932). However, some general statements can be
made: (i) The typical18-cirri oxytrichids (for example, Oxytricha, Tachysoma, Urosoma,
Stylonychia, Steinia, Histriculus) are almost always included in the Oxytrichidae. (ii)
There are some deviating taxa (for example, Ancystropodium, Gonostomum, Gastrostyla,
Onychodromus, Laurentiella) which are classified in various families. (iii) Kerona was
never considered an oxytrichid, except by those authors who included all non-euplotid
hypotrichs in this group (CORUSS 1960, KAHL 1932). Conversely, SMALL & LYNN (1985)
assigned oxytrichid taxa to at least four families (Oxytrichidae, Gonostomatidae, Am-
phisiellidae, Trachelostylidae; the latter three were new!) without, however, detailed ex-
planation. Furthermore, they made some curious nomenclatural mistakes. For instance,
they transferred the type of Gonostomum STERKI, 1878, G. affine, to Trachelostyla I<AHL,
1932; however, simultaneously they considered Gonostomum as valid. In general, I dis-
cuss deviating classifications in the corresponding group section.
The characterization proposed above unifies about 170 valid species, many of which
have 18 frontal-ventral-transverse cirri clustered to six distinct groups (Fig. 6a). This pat-
tern evolved, without doubt, only once, and taxa showing this pattern constitute the main
part of the Oxytrichidae of most previous classifications (see Tables 7-13). Molecular
biological and morphogenetic data and cladistic analyses suggest that some taxa with a
deviating ventral infraciliature (Laurentiel/a, Onychodromus, Pattersoniella) - and thus
previously classified as Oxytrichidae sensu lato or in other families - very likely evolved
from 18-cirri oxytrichids (BERGER & FOISSNER 1997, SCHLEGEL et al. 1991). Paraurostyla,
Gastrostyla, Ancystropodium, and Territricha are further taxa which are often considered
to be in close relationship with the oxytrichids, although they also have a slightly to dis-
tinctly increased number of frontal-ventral-transverse cirri. Unfortunately, it is unknown
if they are sister groups of the 18-cirri oxytrichids, or if they evolved from 18-cirri oxytri-
chids in that the number of cirri increased secondarily. However, all three groups (18-
cirri oxytrichids, Laurentiella ..., Paraurostyla ...) have one cwious character in
common, namely fragmentation of at least one dorsal kinety. Kinety 3 sIlows simple frag-
mentation in the basic pattern (Fig. 24a), and I argue that it is almost impossible that this
(''unimportant'') feature evolved independently several times in exactly the same way.
Thus, I consider it as the most important autapomorphy of the Oxytrichidae, having oc-
curred either earlier or concomitantly with the 18 frontal-ventral-transverse cilTal pattern.
The very unusual dorsal ciliature of the ectocommensal Kerona pediculus originates by
multiple fragmentation of 3 dorsal kineties (Fig. 208v-y). I assume that this is a modified
pattern, like that of Onychodromus quadricornutus (Fig. 195s, t), strongly suggesting an
inclusion of Kerona in the Oxytrichidae.
For a detailed overall characterization of the Oxytrichidae, see the general section of
this book.
104 SYSTEMATIC SECTION

How to recognise an oxytrichid ciliate in practice

Before using the "Key to all genera of the Oxytrichidae" (see below), you have to know
whether the hypotrich in question belongs to this group at all. If one or more of the fol-
lowing points apply, the answer is "yes":
• The species has 18 frontal-ventral-transverse cirri arranged essentially as shown in
Figure 6a. In some species the number of transverse and pretransverse ventral cirri is
slightly to distinctly decreased or these cirri are even lacking (Fig. 19d). For some
similar soil species, see also my revision on Lamtostyla (BERGER & ForssNER 1988a),
which is now classified in the Amphisiellidae (PETz & FOISSNER 1996).
• It is a non-euplotid hypotrich with a rigid body (except Psilotricha). Do not restrict
mobility or squeeze the cell when checking the flexibility!
• The dorsal infraciliature is slightly to distinctly irregular due to fragmentation (for ex-
ample, Fig. 2 19i). Unfortunately, this character needs silver impregnation or very de-
tailed live observation with Normarski differential interference contrast optics!
• The ventral infraciliature shows one of the following patterns: Figure 29f (Gono-
stomum), Figure 29j (Onychodromopsis), Figure 30b (Coniculostomum), Figure 30i
(Onychodromus), Figure 30j (Laurentiella), Figure 30k (Pattersoniel/a), Figure 301
(Gastrostyla), Figure 30m (Kerona), Figure 30n (Paraurostyla), Figure 300 (Territri-
cha), Figure 198a, e (Ancystropodium), Figure 198.1f(Apoamphisiel/a), Figure 218h
(Parentocirrus), Figure 219.1 (Pseudouroleptus).
• It lives on hydras or bryozoans. Kerona pediculus is the only known hypotrich living
on these metazoans (Fig. 30m, 208).

Key to all genera of the Oxytrichidae

Many characters (size, shape, flexibility, cortical granules, colour of cytoplasm, arrange-
ment of cirri, length of dorsal cilia, nuclear apparatus, shape of undulating membranes)
can be recognised by careful examination of live specimens with bright field illumination
and interference contrast optics. Admittedly, this requires patience and experience. Thus,
the following key, based mainly on characters recognisable after protargol impregnation,
a well-known and widely used technique for identifying ciliates (see chapter 6 in the gen-
eral section), can also be used for unstained material. Live observation is indispensable
for species identification! The following key includes all genera considered in the pre-
sent book. A similar key for all 18-cirri oxytrichid genera is given by BERGER & FOISSNER
(1997). Species which do not have the most common type of nuclear apparatus (namely 2
macronuclear nodules and 2 or more micronuclei) are listed in Table I. Eighteen-cirri
oxytrichids with distinct cortical granules and/or distinctly coloured cytoplasm belong to
the Oxytrichinae (Table 2). If you have problems assigning a species to a group, see the
somewhat heterogeneous Oxytricha. It includes species with a deviating oral apparatus
(e. g., 0. siseris, 0. longissima) or an increased number of caudal cirri (e. g., 0. laurel).
 Oxytrichidae 105

Note: For designation of characters used in the following key, see mainly Figures 6a, b,
29a-m, 30a~.

1 Ectocommensal on hydras or bryozoans; body kidney- or bean-shaped; 6 distinctly

curved frontoventral rows (Fig. 30m, 208a, 1) ........... Kerona pediculus (p 826)
Not ectocommensal; body not kidney- or bean-shaped; frontoventral rows more or
less straight (for example, Fig. 29j, 30j, n) or lacking, that is, only 2 (for example,
Fig. 6a, 29a) or more marginal (for example, Fig. 29b, 30g) rows present ....... 2
2 Dorsal processes (horns) present (Fig. 30i, 193a, 1, r, 195a). Onychodromus (p 722)
Dorsal processes lacking ............................................. 2a
2a Anterior end narrowed, that is, set off plug-like from rest of cell ("cephalised"); few
(usually 2) cirri right of posterior end of right marginal row (Fig. 246e, arrow); ma-
rine or brackish water (Fig. 82a, b, 246a-j; 207a-f) ..... Hemigastrostyla (p 933)
No such curious character combination .................................. 3
3 Two or more right, and one or more left rows of marginal cirri, and/or one or more
frontoventral rows (for example, Fig. 29b,j, 1, 30b, g,j, 1, n, 218h) ............ 4
One right and one left row of marginal cirri, and no distinct frontoventral row(s), but
several (usually 18 [for example, Fig. 6a], some species with less [for example, Fig.
2ge], two with more [Fig. 30k, 0]) frontal-ventral-transverse cirri clustered to distinct
groups ............................................................ 14
4 Typical 18 frontal-ventral-transverse cirri pattern present (for example, Fig. 29b,
30g) .............................................................. 5
Typical 18 frontal-ventral-transverse cirri lacking (for example, Fig. 30j, n, 218h). 8
5 Body tailed (Fig. 291) .......................... Parurosoma dubium (p 492)
Body not tailed ...................................................... 6
6 Macronucleus moniliform (Fig. la, 30b) .............. Coniculostomum (p 606)
Macronucleus consists of two nodules ................................... 7
7 Body very flexible; cortical granules present; caudal cirri present (sometimes diffi-
cult to distinguish from marginal cirri, which usually insert more ventrally than cau-
dal cirri (for example, Fig. 29b, 91c) ...................... Allotricha (p 261)
Body rigid; cortical granules lacking; caudal cirri absent (Fig. 30g, 189f) ......... .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. Pleurotricha (p 698)
8 (4) Body tapered posteriorly (Fig. 219.1) or with club-shaped process (Fig. 198) 8a
Body not distinctly tapered and/or without club-shaped process ............... 9
8a Body with club-shaped process (Fig. 198) ...... Ancystropodium maupasi (p 778)
Body tapered posteriorly (Fig. 219.1) ......... Pseudouroleptus caudatus (p 889)
9 Oral apparatus (undulating membranes and adoral zone of membranelles) in Gono-
stomum pattern (Fig. 19c, h, 124a-c) ......... Gonostomum gonostomoida (p 392)
Oral apparatus not in Gonostomum pattern .............................. 10
10 Body almost triangular, rigid (Fig. 30j) .............. Laurentiella strenua (p 753)
Body more or less elliptical, flexible (for example, Fig. 29j, 301, n, 218h) ...... 11
106 SYSTEMATIC SECTION

11 One more or less continuous frontoventral row, that is, 3 cirraI rows including mar-
ginal rows (for example, Fig. 301, 202h) ................... Gastrostyla (p 789)
More than 1 frontoventral row, that is, 4 or more cirral rows including marginal rows
(for example, Fig. 30n) .............................................. 12
12 Two frontoventral rows, that is, 4 (Fig. 215a-c, 218h) to 5 (Fig. 29j) cirral rows in-
cluding marginal rows ............................................... 13
Usually 3-7 frontoventral rows, that is, 5-9 (or more) cirral rows including marginal
rows (Fig. 30n, 212r) ................................. Paraurostyla (p 841)
13 Single pretransverse ventral cirri present (Fig. 29j) ..... Onychodromopsis (p 475)
Single pretransverse ventral cirri lacking (Fig. 215c, 218h) .................... 31
14 (3) Transverse cirri more than 5 in number (Fig. 30k, 0) .................... 15
Transverse cirri 5 or less in number (for example, Fig. 29a, e, f) ............. 16

Fig. 29a-g Ventral infraciliature (protargol impregnation) of representatives of genera assigned to the Oxytri- --l
chinae. For designation of structures, see Figure 6a; sources and sizes, see species descriptions. a) Oxytricha
granulifera. Arrowhead marks postoral ventral cirrus V/3 which is close behind the buccal vertex in Oxytri-
chao b) Allotricha antarctica. Arrow marks second (= inner) right marginal cirral row. c) Cyrtohymena museD-
rum. Note strongly curved paroral (arrow). d) Notohymena rubescens. Note anteriorly hooked paroral (tiny ar-
row). e) Urosomoida agilis. Note reduced number of pretransverse ventral and transverse cirri (arrowhead).l)
Gonostomum affine. Note special oral apparatus. g) Urosoma caudata. Note linear arrangement of frontoven-
tral cirri (arrowhead). Figures on p 107.

Fig. 29b-m Ventral infraciliature (protargol impregnation unless otherwise indicated) of representatives of 4
genera assigned to the Oxytrichinae. For designation of structures, see Figure 6a; sources and sizes, see species
descriptions. h) Tachysoma pellionellum. Tachysoma lacks caudal cirri. i) Australocirrus oscitans. Note huge
oral apparatus (body flexible). j) Onychodromopsis flexilis. Note increased number of cirral rows and special
arrangement of frontal-ventral-transverse cirri. Arrow marks pretransverse ventral cirri. k) Rubrioxytricha hae-
matoplasma. Rubrioxytricha has a reddish or brownish cytoplasm. I) Parurosoma dubium (mercuric chloride
fIXation). Note increased number of marginal rows. m) Pseudostrombidium planctonticum (cilia of cirri also
shown). Euplanktonic; note increased number of caudal cirri (arrowheads). Figures on p 108.

Fig.30a-f Ventral infraciliature (protargol impregnation) of representatives of genera assigned to the Stylony- 4
chinae. For designation of structures, see Figure 6a; sources and sizes, see species descriptions. a) Stylonychia
mytilus complex. Note straight and parallel arrangement of undulating membranes. b) Coniculostomum moni-
lata. Has almost the same ciliature as Stylonychia (compare black cirri of a and b), except for the parental right
marginal rows (white cirri). c) Steinia sphagnicola. Note fragmented endoral (arrowhead) and conspicuous pit
(tiny arrow). d) Histriculus histrio. Marginal rows closed posteriorly (at least in this species; arrowhead) and
lacks caudal cirri. e) Sterkiella histriomuscorum. Arrowhead marks postoral ventral cirrus V/3, which is dis-
tinctly set off in Sterkiella. l) Parastylonychia plouneouri. Marine; lacks caudal cirri. Figures on p 109.

Fig.30g-j Ventral infraciliature (protargol impregnation) of representatives of genera assigned to the Stylony- ~
chinae. For designation of structures, see Figure 6a; sources and sizes, see species descriptions. g) Pleurotricha
lanceolata. Note increased number of marginal rows (arrow). h) Rigidocortex octonuc/eatus. Has about 8
macronuclear nodules. i) Onychodromus grandis. Onychodromus has dorsal horns. j) Laurentiella strenua.
Note triangular shape. Figures on p 110.
 Oxytrichidae 107

.
?
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-9
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 108 SYSTEMA TIC SECTION

. . • ·.
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,
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Fig. 29b- m Legend on p 106.

 Oxytr'Ieh'Idae 109

.. =:::.
, • •
., • := '
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=
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=
=
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Fig. 30a- f Legend on p 106.
110 SYSTEMATIC SECTIO
N

,
., ,
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I #
.,
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I #

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~.

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.
 Oxytrichidae 111

# ......
*
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, III

*
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,
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. .. .. Fig. 30k-o Ventral infraciliature (after

protargol impregnation) of representatives
of genera assigned to the Stylonychinae
, (k) or of uncertain position within the
, Oxytrichidae (1-0). k) Pattersoniella viti-
phila. Note increased number of frontal-
, ventral-transverse cirri. I) Gastrostyla
• , steinii. Note irregular frontoventral row.
,,, Arrow marks prominent postoral ventral
.. , cirrus. m) Kerona pediculus. Note bean-

,,, shaped outline and the highly increased

number of frontoventral cirri within the 6
• frontal-ventral-transverse cm streaks
• ~

• (1-6). Dorsal side almost completely cili-

,
~

•
• .\ t
~
ated. Lives on hydras and bryozoans. 0)
,
~
~
Paraurostyla weissei complex. Note fron-
f
toventral rows (arrows). 0) Territricha
"t f stramenticola. Note slightly increased
number of postoral ventral and transverse
I
cirri.
112 SYSTEMATIC SECTION

15 Two macronuclear nodules; 3 frontal cirri (Fig. 300); common in litter of European
beech forests ................................. Territricha stramenticola (p 884)
13-19 macronuclear nodules; 5-8 frontal cirri (Fig. 30k, 1979); in soil of Southern
hemisphere ................................ Pattersoniella vitiphila (p 766)
16 (14) Undulating membranes in Steinia pattern (Fig. 19j, 20f, 30c) .. Steinia (p 624)
Undulating membranes not as above .................................... 17
17 Undulating membranes in Cyrtohymena pattern (Fig. 199, 20e) . . . . . . . . .. . ..... .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. . ............... Cyrtohymena (p 279)
Undulating membranes not as above .................................... 18
18 Undulating membranes in Notohymena pattern (Fig. 19f, 20b) Notohymena (p 324)
Undulating membranes not as above .................................... 19
19 Undulating membranes and adoral zone of membranelles in Gonostomum pattern
(Fig. 19c, h, 20c) .................................................... 20
Undulating membranes and adoral zone not as above ...................... 21
20 Adoral zone of membranelles <30 % of body length; 4 dorsal kineties (Fig. 29g,
1280; often rather slender and tailed) ....................... Urosoma (p 396)
Adoral zone of membranelles 40-50 % of body length; 3 dorsal kineties (Fig. 29f,
117g; very common in soil) ............................ Gonostomum (p 367)
21 (19) Undulating membranes in Stylonychia pattern (for example, Fig. 19i, 20d, 30a,
b, f) .............................................................. 22
Undulating membranes in Oxytricha pattern (for example, Fig. 6a, 1ge, 20a, 29a,
30e) ............................................................. 23
22 Caudal cirri present (often prominent, sometimes, however, difficult to distinguish
from marginal cirri, which usually insert more ventrally than caudal cirri; Fig. 30a,
148a, 161a) ......................................... Stylonychia(p 501)
Caudal cirri absent (Fig. 30f; marine) ......... Parastylonychia plouneouri (p 697)
23 (21) Caudal cirri present (sometimes difficult to distinguish from marginal cirri,
which usually insert more ventrally than caudal cirri; Fig. 6b) ............... 25
Caudal cirri absent .................................................. 24
24 Marginal rows distinctly separate posteriorly; body supple (for example, Fig. 29h,
135p) .............................................. Tachysoma(p431)
Marginal rows confluent posteriorly; body rigid (Fig. 30d) ..... Histriculus (p 645)
25 (23) Number of pretransverse ventral and transverse cirri usually <6 (if 6 pretrans-
verse ventral cirri and transverse cirri present, see also Oxytricha; Fig. 19d, 2ge) ...
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. Urosomoida (p 345)
Two pretransverse ventral and 5 (sometimes only 4) transverse cirri (for example,
Fig. 29a, 30e; if only 4 dorsal kineties and 1 or 2 caudal cirri present, see also
Urosomoida) ...................................................... 26
26 Caudal cirri 10-20 in number, form rather dense corona at narrowly rounded poste-
rior end of organism (caudal cirri sometimes difficult to distinguish from marginal
cirri, which usually insert more ventrally than caudal cirri, Fig. 29m; usually
planktonic) ......................... Pseudostrombidium planctonticum (p 494)
 Oxytrichinae 113

Caudal cirri 1-3 in number ........................................... 27

27 Macronuclear nodules 7-13, usually about 8 in number (Fig. 30h, 192a-l; body
rigid) ...................................... Rigidocortex octonucleatus (p 718)
Macronuclear nodules 2 (very common), 3 (only Sterkiella thompsom), or 4 (com-
mon) in number .................................................... 28
28 Body flexible and oral apparatus very prominent (Fig. 29i, 142a-e) ............. .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. Australocirrus oscitans (p 470)
No such character combination ........................................ 29
29 Cytoplasm colourless, that is, not distinctly reddish or brownish; usually 3 caudal
cirri .............................................................. 30
Cytoplasm usually distinctly, rarely only slightly homogeneously orange to reddish
or brown; only 1 or 2 caudal cirri (Fig. 29k, 144c) ......... Rubrioxytricha (p 479)
30 Adoral zone ofmembranelles ~S % of body length; postoral ventral cirri form nar-
row group close behind buccal vertex; ratio of body width to length <40 %, usually
about 30 %; body supple (for example, Fig. 29a; if only 4 transverse cirri and 4 dor-
sal kineties present, see also Urosomoida); species identification usually very
difficult ................................................... Oxytricha (p 115)
Adoral zone of membranelles ~40 of body length; postoral ventral cirri do not form
narrow group close behind buccal vertex, that is, cirrus V/3 distinctly set off from the
other two postoral cirri; ratio of body width to length ~40 %; body rigid or slightly
flexible (for example, Fig. 30e, 18Ia-g; often misidentified as Oxytricha or Styla-
nychia) ............................................... Sterkiella (p 668)
31 (13) 2 macronuclear nodules (for example, Fig. 21Sc) ..... Apoamphisiella (p 781)
Usually 8 (range 7-12) macronuclear nodules (Fig. 218h, i) ................... .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. Parentocirrus hortualis (p 879)

Oxytrichinae EHRENBERG, 1838

1838 Oxytrichina EHRENBERG, lnfusionsthierchen, p 362 (in part) - Name-bearing type: Oxytricha BORY DE
SAINT-VINCENT in LAMOUROux, BORY DE SAINT-VINCENT & DESLONGCHAMPS, 1824.
1997 Oxytrichinae JANKOWSKI, 1979 - BERGER & FOISSNER, Arch. Protistenk., 148: 145.

C h a r act e r i sat ion: O~chidae with flexible (supple) body. Cortical granules
present, secondarily lost several times. Adoral zone of membrane lies usually :9t0 % of
body length. Primarily 18 frontal-ventral-transverse cirri clustered to six distinct groups,
number of cirri of some groups (especially transverse and pre- and postoral ventral cirri)
slightly to distinctly decreased. Postoral ventral cirrus V/3 participates in primordia for-
mation. Dorsal morphogenesis primarily in Oxytricha pattern (Fig. 24a), secondarily
fragmentation ofkinety 3 and dorsomarginal kineties absent.
114 SYSTEMATIC SECTION

Rem ark s: Recently I categorised this monophylum as subfamily, as also indicated by

the defmed ending -inae (BERGER & FOISSNER 1997). Here I ignore the category as such,
but, to avoid inflation, retain the name. Furthermore, in BERGER & FOISSNER (1997) I cred-
ited this taxon to JANKOWSKI (1979, P 77) because he published this category as follows:
Subfamily 1. Oxytrichinae EHRENBERG, 1838, s. str., subfam. n. Now I prefer to credit it to
EHRENBERG (1838), as obviously already proposed by JANKOWSKI (1979) himself.
Phylogenetic analysis strongly suggests that the 18-cirri oxytrichids branch into two
roughly equal sized groups, namely the Oxytrichinae with a plesiomorphic flexible body,
and the Stylonychinae with a rigid body as an evolutionary novelty. In the cladogram pro-
posed (Fig. 25a, b), the Oxytrichinae are defined by a distinctive ontogenetic autapomor-
phy, namely the participation of the posterior-most postoral ventral cirrus (V/3) in pri-
mordia formation (for detailed explanation of this character, see character 19 in the phy-
logeny chapter of the general section). In the Oxytrichinae this cirrus is usually arranged
very close to the remaining postoral cirri, whereas it is distinctly displaced posteriorly in
the sister group, Stylonychinae. The other features mentioned in the characterisation of
the Oxytrichinae are either plesiomorphies (for example, flexible body, cortical granules
present) or conspicuous autapomorphies of members of this group, for instance, number
of pretransverse ventral and transverse cirri decreased in Urosomoida, loss of fragmenta-
tion of dorsal kinety 3 in Urosomoida and in the Urosoma-Gonostomum group, and loss
of dorsomarginal kineties in Gonostomum.
Some of the sister group relationships proposed in the cladogram are well-defined
(for example, Gonostomum-Urosoma, Oxytricha-Allotricha), others are rather weak due
to several parallelisms (Fig. 25a, b). In addition, some taxa (Australocirrus, Onychodro-
mopsis, Rubrioxytricha, Parurosoma, Pseudostrombidium), also having the typical (or
somewhat modified) 18 frontal-ventral-transverse cirral pattern and a supple body, could
not be considered in the phylogenetic analysis because detailed morphogenetic data are
lacking. Thus, the proposed cladograms are rather incomplete and will change more or
less significantly as further taxa are analysed. Furthermore, most of the important mor-
phogenetic characters used for the analysis are substantiated for the type species only. As
a consequence of this uncertainty I did not name clades within the Oxytrichinae.
Still, some genera are not dermed by a unique character (autapomorphy) but by a
specific combination of plesiomorphies, that is, their monophyly is questionable. Thus,
and because I did not name and characterise the sister groups within the Oxytrichinae, the
characterisation of a genus includes all features necessary to separate it from the other
genera of the Oxytrichinae.

Key to genera

For identification of taxa, see general key on page 104. Species with the 18 frontal-
ventral-transverse cirral pattern and cortical granules belong to the Oxytrichinae (Table
2).
 Oxytricha 115

Oxytricha BORY DE SAINT-VINCENT in LAMOUROUX, BORY DE SAINT-

VINCENT & DESLONGCHAMPS, 1824

1824 Oxitricha BORY DE SAINT-VINCENT in LAMOUROux, BORY DE SAINT-VINCENT & DESLONGOIAMPS, Ency-
clopedie methodique, p 593 (see nomenclature and taxomony) - Type (subsequent designation by
FOISSNER 1989, P 235; see nomenclature): Oxytricha granulifera FOISSNER & ADAM, 1983.
1826 Oxitricha BORY DE SAINT-VINCENT, Essai d'une classification, p 55 (see nomenclature and taxonomy).
1830 Oxytricha pellionella BORY. - EHRENBERG, Abh. preuss. Akad. Wiss., year 1830: 43 (see nomenclature).
1838 Oxytricha - EHRENBERG, Infusionsthierchen, p 363 (see nomenclature and taxonomy).
1859 Oxytricha EHRBG. - STEIN, Organismus der Infusionsthiere I, p 182 (in part).
1866 Actinotricha COHN, Z. wiss. Zool., 16: 283 (synonym) - Type (by original designation and rnonotypy):
Actinotricha saltans CoHN, 1866.
1875 Oxytricha- FROMENTEL, Etudes microzoaires, p 160 (in part; see nomenclature and taxonomy).
1882 Oxytricha, EHRENBERG - KEN"r, Manual infusoria, p 786 (in part).
1882 Opisthotricha KEN"r, Manual infusoria II, p 785 - Type (original designation): Oxytricha parallela
ENGELMANN, 1862.
1889 Oxytricha (Oxitricha p. p. BORY 1824) EHRB. 1830 emend. STERKI 1878 - BOTscHU, Protozoa, p 1749
(in part).
1932 Oxytricha (BARY, 1824) EHRENBERG, 1838, emend. WRZESNIOWSKI, 1877 - KAHL, Tierwelt Dtl., 25: 599
(in part; incorrect spelling OfBORY).
1932 Oxytricha WRZESNlOWSKI, 1877 - KAHL, Tierwelt Dtl., 25: 601 (in part).
1932 Opisthotricha KEN"r, 1881- KAHL, Tierwelt Otl., 25: 608 (in part).
1972 Oxytricha EHRENBERG, 1830 - BoRROR, J. Protozoal., 19: 13 (in part; see nomenclature and taxonomy).
1974 Oxytricha EHRENBERG emend. WRZESNlOWSKI- STILLER, Fauna Hung., 115: 128.
1989 Oxytricha - FOlsSNER, Sber. Ost Akad. Wiss., 196: 235 (fixation of type species; see nomenclature).
1997 Oxytricha BORY DE SAINT-VINCENT in LAMOUROux, BoRY DE SAINT-VINCENT & DESLONGCHAMPS, 1824-
BERGER & FOIssNER, Arch. Protistenk., 148: 146.

C h a r act e r i sat ion: Adoral zone of membranelles formed like a question mark.
Undulating membranes in Oxytricha pattern. Frontoventral cirri in V-shaped pattern.
Postoral ventral cirri in dense cluster behind buccal vertex. Two pretransverse ventral
cirri and 5, rarely 4 transverse cirri. One right and 1 left row of marginal cirri. Usually 5
or 6, rarely 4 dorsal kineties. Caudal cirri present. Primordia V and VI ofproter originate
from cirrus V/4 and V/3 respectively. Dorsal morphogenesis in Oxytricha or Urosomoida
pattern.

Nom e n c I a t u r e: The nomenclature of Oxytricha is very complicated. BORY DE

SAINT-VINCENT established this genus as "Oxitrichd'; however, the spelling Oxytricha,
which was used for the first time by EHRENBERG (1830, P 43, 54), prevailed. In spite of
this, BORY DE SAINT-VINCENT should be accepted as the author of Oxytricha (CORLISS
1979).
BORY DE SAINT-VINCENT assigned the following species to "Oxitricha": O. lepus (ba-
sionym: Trichoda lepus MOlLER, 1773, P 89; species indeterminata); O. pulex (basionym:
Trichoda pulex MOlLER, 1773, P 80; 1786, p 225, Tab. XXXI, fig. 26; species indetermi-
nata?); O. pullicina (basionym: Trichoda pulex MOLLER, 1773, P 80; 1786, p 225, Tab.
XXXI, fig. 25); O. pelionella (basionym: Trichoda pellionella MOLLER, 1773, P 80; now
116 SYSTEMATIC SECTION

Tachysoma pellionellum); O. gallina (basionym: Trichoda gallina MOLLER, 1786, P 209;

now Uroleptus gallina (MOLLER, 1786) ForSSNER, BLATfERER, BERGER & KOllMANN, 1991);
O. pullaster (basionym: Trichoda pullaster MOLLER, 1773, P 81; now Holosticha pullas-
ter (MOLLER, 1773) FOISSNER, BLATIERER, BERGER & KOllMANN, 1991); O. transfuga (ba-
sionym: Trichoda transfuga MOLLER, 1776, P 281; now Uronychia transfuga (MOLLER,
1776) STEIN, 1859c); 0. bulla (basionym: Trichoda bulla MOLLER, 1786, P 222; species
indeterminata); 0. gibbosa (basionym: Trichoda gibba MOLLER, 1786, P 179; species in-
determinata?); o. volutator (basionym: Himantopus volutator MOLLER, 1786, P 251; spe-
cies indeterminata); o.joblotii (species indeterminata); O.felis (basionym: Trichodafelis
MOLLER, 1786, P 2l3; species indeterminata); O. variabilis (species indeterminata); O.
ambigua (basionym: Trichoda ambigua MOLLER, 1786, p 200; now Spirostomum ambi-
guum (MOLLER, 1786) EHRENBERG, 1835).
EHRENBERG (1830) used the name Oxytricha in combination with the species names
O. pellionella (now type of Tachysoma), 0. piscis (now Uroleptus piscis), O. pullaster
(now Holosticha pullaster), and o. lepus (indeterminable).
Neither BORY DE SAINT-VINCENf nor EHRENBERG (1830, 1838) fIxed a type species.
This was done for the fIrst time by FROMENIEL (1875) with Oxytricha crassa CLAPAREDE
& LAClIMANN, 1858, a species not originally included in "Oxitrichd' (see above and be-
low). Unfortunately, KAHL (1932) ignored this decision, and transferred O. crassa to the
subgenus "Holosticha (Frichotaxis)". KAHL (1932) himself did not fIx a type species in
Oxytricha. Forty years later, BORROR (1972a) synonymised Steinia with Oxytricha, fIx-
ing Oxytricha platystoma EHRENBERG, 1831, as type of Oxytricha; however, this species
was subsequently designated as the type of the resurrected Steinia (FOISSNER 1989,
p 235). This caused an almost indissoluble type problem since none of the originally in-
cluded species is now considered to be an Oxytricha and many of them are indetermin-
able. Thus, ForSSNER (1989) proposed Oxytricha granulifera FOISSNER & AoAM, 1983, as
type, because this species represents what is generally considered as a ''typical'' Oxytri-
cha, especially in the sense of KAHL (1932), who provided the last authoritative charac-
terisation of this voluminous taxon. Furthermore, Oxytricha granulifera is described in
great detail both by live observations and after protargol impregnation, it is well charac-
terised morphogenetically, and the holo- and paratype slides are deposited in a museum.
Thus, I agree with FOISSNER (1989) to "fIx" O. granulifera finally as the type of Oxytri-
cha, although this is not in agreement with Article 67 (g) of the IcZN (1985). Accordingly,
a species cannot be fIxed as type species if it was not originally included; however, fol-
lowing the code strictly would mean numerous changes of names, which is hardly in the
interest the nomenclatural stability.
KAHL (1932) divided Oxytricha into 7 subgenera, namely Oxytricha (Oxytricha), o.
(Opisthotricha), o. (Stylonychia), 0. (Histrio), o. (Fachysoma), o. (Steinia), and o.
(Urosoma), so producing some primary and secondary homonyms. The most interesting
example is the primary homonymy of 0. (Opisthotricha) muscorum, O. (Sty/onychia)
muscorum, O. (Histrio) muscorum, and o. (Steinia) muscorum, which I corrected just re-
 Oxytricha 117

cently (ForSSNER et al. 1991; see also Sterkiella histriomuscorum and Cyrtohymena mus-
corum).
Some incorrect subsequent spellings: Ocytricha sp. (GEBHARDT 1933, p 154); Onytri-
cha (MAUPAS 1888, P 276); Opistotricha (GRACIA & IGUAL 1987b, P 3; SUOZUKI 1964a,
P 249, 1964c, P 168; SUOZUKI & SHIMOIZUMI 1967, P 216); Opistotricha simi/is (HoRvATH
1950, p 156); Opistrotricha simi/is (HoRvATH 1950, p 155); Opithotricha muscorum
KAHL (WANG 1977, P 145); Oxitrica sp. (GRACIA et al. 1987b); Oxitricha spec. (Voss
1978, p 42); Oxitrycha sp. (SALvAD6 & GRACIA 1993, p 893); Oxythricha (BAMFORTH
1967, P 15; HENEBRY & GORDEN 1989, p 1019); Oxyticha (BURGES 1958, P 43); Oxytri-
chia (LACKEY 1926a, b, p 37, 45; LACKEY 1927, P 519; LACKEY 1942, P 196); Oxytrichum
(GIPSON & KALINSKY 1982, p 134); Oxytrocha sp. (CAIRNS & YONGUE 1973b, P 202).

T a x 0 nom y: The characterisation of Oxytricha is according to BERGER & FOISSNER

(l997). For explanation of patterns mentioned, see chapters 1 and 2 in the general
section. Oxytricha is the most voluminous taxon within the oxytrichids, comprising about
50 valid species and 60 species indeterminata; most of the latter have been described by
DUMAS (1929, 1930). Only about a quarter of the valid species is characterised in detail
both by live observations and after protargol impregnation. All other valid taxa are de-
scribed usually only after more or less exact live observation or inadequate staining meth-
ods, and the differences to other oxytrichids are often rather vague. However, the avail-
able data leave at least a small chance of recognising them; thus, I do not classify them as
species indeterminata. More detailed investigations of these species will presumably re-
veal that some belong to other taxa, that is, the "valid" Oxytricha species mentioned in
this book very likely do not form a monophylum. However, the available data are too
scant for a more detailed classification. Furthermore, the sparse and often rather doubtful
observations made the construction of a useful key almost impossible. Identification of
Oxytricha species is thus a very difficult task. One of the most problematic species is
Oxytricha hymenostoma, which looks like a Sterkiella or Stylonychia. However, STOKES
(l887b) described the body as soft and flexible, so preventing a synonymisation with the
rather similar and very common Sterkiella histriomuscorum or a Stylonychia.
Opisthotricha KENT was established to include Oxytricha species with elongated cau-
dal cirri. However, at the present stage of knowledge this character can be used for spe-
cies distinction only. Synonymy was already proposed by BORROR (1972a).
Oxytricha and Allotricha are very likely sister groups, as indicated by the almost
identical ventral and dorsal infraciliature and - more convincingly - by distinctive mor-
phogenetic peculiarities (see Fig. 23c). However, the autapomorphy of this Oxytricha-
Allotricha group has to be substantiated by the investigation of further species.
Allotricha, a resurrected taxon described by STERKI (1878), has at least two right marginal
rows, and can thus be easily distinguished from Oxytricha.
Species are arranged according to biotopes, namely freshwater, soil, and saline wa-
ters. Some of them (for example, Oxytricha setigera) occur in two types of biotopes. Spe-
cies are arranged alphabetically within these biotope groups.
 118 SYSTEMATIC SECTION

Key to species

The identification of Oxytricha species is very difficult and in several cases very likely
impossible, especially in practical applications (for example, water pollution assessment),
when often only few specimens are available. Furthermore, the differences to other
groups are often very difficult to ascertain, especially in life; thus, see also Urosomoida if
the number of transverse cirri and pretransverse ventral cirri is decreased and only 4 dor-
sal kineties are present, Rubrioxytricha if the cytoplasm is homogeneously reddish or
brownish, Urosoma if frontoventral cirri form a line, as in Figure 19a, and only 4 dorsal
kineties present, Sterkiella histriomuscorum if the body is not very flexible, cortical gran-
ules are lacking, and the posterior postoral ventral cirrus V /3 is distinctly set off. Detailed
live observation using oil immersion is absolutely necessary for identification. Important
characters for identification are, among other things, the biotope, the size, the length of
dorsal cilia, the nuclear apparatus, and the cortical granules. Note that characters and
data mentioned in the keys and tables are usually not repeated in the text!

1 Two macronuclear nodules ............................................ 2

More than 2 macronuclear nodules ..................................... la
la 8 macronuclear nodules; cortical granules present (Fig. 73c, d, f, h) ............. .
...................................................... O.ottowi(p218)
4-6, usually 4 macronuclear nodules; cortical granules lacking (Fig. 68a, t) ....... .
· . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. O. islandica (p 207)
2 Dorsal cilia >6 ~m ................................................... 3
Dorsal cilia::;;6 ~m .................................................. 11
3 Only I micronucleus between macronuclear nodules (Fig. Ic) ............... 6
Each macronuclear nodule with I or more micronuclei (if only I micronucleus is pre-
sent, it is usually not located between the 2 macronuclear nodules; for example, Fig.
2) ............................................................... 3a
3a Freshwater or soil ................................................... 4
Marine ........................................................... 50
4 Length in life about 200 ~m; 4 distinct caudal cirri (Fig. 52a, b) ............... .
· . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. O. parallela (p 171)
Length in life about 60-120 ~m; 3 or less caudal cirri ...................... 5
5 Undulating membranes nearly straight, arranged side by side (Fig. 77) .......... .
· . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. O. pseudosimilis (p 222)
Undulating membranes distinctly bent, arranged consecutively (Fig. 58c, g) ...... .
· ..................................................... O. siseris (p 187)
6 (3) Length in life about 40-60 ~m, rarely up to 80 ~m ....................•. 7
Length in life usually more than 80 ~m or 6 dorsal kineties (Fig. 244d-g) ...... 8a
7 Buccal cirrus at posterior end of undulating membranes; 4 dorsal kineties (Fig. 55g,
h, i, p, r) .................................................. 0. setigera (p 175)
 Oxytricha 119

Buccal cirrus at anterior end of undulating membranes; 5 dorsal kineties (Fig. 33a-e;
see also the very similar O. opisthomuscorum, lead 10-) ..... O. balladyna (p 126)
8a (6) Length in life 155-280 ~m; cortical granules present (Fig. 40a-d) ........... .
· . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. O. granulosa (p 148)
Length in life about 80-150 ~m; cortical granules lacking .................. 8b
8b In life about 4.6 times as long as broad (Fig. 59) ............. 0. sphagni (p 190)
In life about 2.5-3.5 times as long as broad (Fig. 35, 37a, 73a) ............... 9
9 Outline spindle-shaped; about 2.5 times as long as broad (Fig. 37a, b) ........... .
· . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. O. euglenivora (p 136)
Outline others; about 3-3.5 times as long as broad ........................ 10
10 Transverse cirri fringed, distally distinctly bent to the left (Fig. 35) ............. .
· . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. O. crassistilata (p 135)
Transverse cirri not fringed, distally not bent (Fig. 73a, b; see also the very similar O.
balladyna, lead 7-) ............................ 0. opisthomuscorum (p 217, 925)
11 (2) Length in life more than 200 ~m .................................... 12
Length in life less than 200 ~m ........................................ 13
12 Marginal cirri become distinctly longer toward rear (Fig. 36; very uncertain species).
· . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. O. elongata (p 136)
Marginal cirri of about same length (Fig. 47) .............. O. /ongissima (p 164)
13 (11) Symbiotic algae present (Fig. 34a, b) ............... O. chlorelligera (p 129)
Symbiotic algae absent ............................................. 13a
13a Outline very slender; only 3 very prominent adoral membranelies (Fig. 90; very rare
[uncertain?], marine species) ............................ O. tricomis (p 241)
Outline less,slender than shown in Figure 90; normal adoral zone ofmembranelles 14
14 Single micronucleus between macronuclear nodules (Fig. lc) ................ 15
Two or more micronuclei (if only 1 micronucleus present, then usually not between
macronuclear nodules) ............................................... 22
15 Length in life about 150-200 ~m (Fig. 46a, b) ............ O. longicirrata (p 163)
Length in life less than 150 ~m ........................................ 16
16 Length in life about 35-70 ~m ........................................ 17
Length in life about 80-120 ~m ....................................... 19
17 4 transverse cirri (Fig. 53a, b) ......................... O. pseudofurcata (p 173)
5 transverse cirri ................................................... 18
18 Right marginal row terminates in front of right transverse cirrus; pretransverse ven-
tral cirri distinctly displaced anteriad; saline lakes (Fig. 83a-g) ... O. geleii (p 232)
Right marginal row terminates behind transverse cirri at posterior end of cell; pre-
transverse ventral cirri not distinctly displaced anteriad; soil (Fig. 76) ........... .
· . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. 0. pseudofusiformis (p 222)
19 (16) Caudal cirri conspicuously elongated (for example, Fig. 84a, b) .......... 20
Caudal cirri not distinctly elongated (for example, Fig. 54a-c) ............... 21
20 Saline waters (Fig. 84a, b) ............................. 0. halophila (p 232)
120 SYSTEMATIC SECTION

Freshwater ........................................................ 49
21 (19) All marginal cirri long and fme; transverse cirri long and prominent; dorsal cilia
about 6 /-lm long (Fig. 54a-c) .............................. O. saprobia (p 174)
Posterior-most left and right marginal cirrus slightly elongated and enlarged; trans-
verse cirri not conspicuously enlarged; dorsal cilia about 2 /-lm long (Fig. 32) ..... .
· ..................................................... O. alfredi (p 125
22 (14) Cortical granules present and/or cytoplasm rusty or reddish by granules (Note
that O. enigmatica also has cortical granules, Fig. 82a, b, 246c) .............. 23
Cortical granules absent and cytoplasm more or less colourless .............. 31
23 Cytoplasm rusty or reddish by granules ................................. 24
Cytoplasm without distinctly coloured granules (cortical granules present!) .... 25
24 Cell about 2.7-3.3 times as long as broad; transverse cirri displaced anteriad, pro-
truding only slightly beyond posterior end of cell (Fig. 31 b, c) ................. .
· ............................. O. aeruginosa (p 122; see also Rubrioxytricha)
Cell only about 1.6 times as long as broad; transverse cirri obviously arranged rather
closely to posterior end of cell (Fig. 222o-t) .................................. .
· ............................... O. truncata, a species indeterminata? (p 254)
25 (23) Body cylindrical, about 8.5 times as long as broad (Fig. 39a, g) .............. .
· . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. O.laurei (p 146)
Body not cylindrical, about 2-3.5 times as long as broad .................. 25a
25a Marine (Fig. 186d) ............................... O. durhamiensis (p 228)
Freshwater or soil .................................................. 26
26 Body about 2 times as long as broad, outline pyriform; freshwater (Fig. 61a-e) .... .
· ............................. '. . . . . . . . . . . . . . . . . . . . . .. O. tenella (p 191)
Body about 2.5-3.5 times as long as broad, outline more or less elliptical; soil (for
example, Fig. 71a~) ................................................ 27
27 Cortical granules colourless or only slightly yellowish ..................... 28
Cortical granules orange-yellow or shiny red ............................. 30
28 Cortical granules rod-shaped, about 2-3 x 0.5-1/-lm sized, arranged in short longitu-
dinal rows; 6 dorsal kineties (Fig. 71a-d, g) ........... O. longigranulosa (p 213)
Cortical granules spherical, 0.5-1/-lm across, arranged in long longitudinal rows or
in irregular patches; 5 or 4 dorsal kineties (Fig. 65a, b, g, 116b) ............ 28a
28a Cortical granules in longitudinal rows; 5 dorsal kineties (Fig. 65b) ........... 29
Cortical granules in irregular patches; 4 dorsal kineties (Fig. 111 b) .............. .
· ......................................... Urosomoida granulifera (p 352)
29 Transverse cirri constantly 5 (n = 15) in number; right marginal row begins at level
of right frontal cirrus, terminates behind transverse cirri; anterior pretransverse
ventral cirrus distinctly separated from posterior one (Fig. 65f) ................. .
· ................................................. O. granulijera (p 197)
Transverse cirri usually 4, rarely only 3 (mean = 3.9; n = 15) in number; right mar-
ginal row begins at about level of middle frontoventral cirri and terminates
slightly above level of transverse cirri; both pretransverse ventral cirri are located at
 Oxytricha 121

about same level, immediately right of left transverse cirrus (Fig. 78a) ............ .
· . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. O. quadricirrata (p 223)
30 (27) Cortical granules orange-yellow; about 23-27 adoral membranelles (Fig. 63a-e)
· ................................................ 0. auripunctata (p 194)
Cortical granules shiny red; about 36-42 adoral membranelles (Fig. 79a-e) ....... .
· . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. O. rubripuncta (p 224)
31 (22) Soil .......................................................... 32
Freshwater or saline water ........................................... 38
32 Length in life about 150-200 !lm (Fig. 64) ............. 0. bimembranata (p 196)
Length in life usually <150!lm ........................................ 33
33 4 or 5 transverse cirri; usually not very flexible (Fig. 183, 184) ................ .
· . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. Sterkiella histriomuscorum (p 683)
Usually 5, rarely 6 transverse cirri ..................................... 35
34 (43) Usually 4 transverse cirri (that is, 6 cirri inclusively pretransverse ventral cirri;
Fig. 44a-d) ............................................ O. longa (p 158)
5 transverse cirri (that is, 7 cirri inclusively pretransverse ventral cirri; Fig. 57a-d) ..
· ..................................................... O. simi/is (p 183)
35 (33) Body in life about 4-<i times as long as broad (Fig. 74) .... O. procera (p 220)
Body in life about 2.2-3.7 times as long as broad ......................... 36
36 Body in life about 2.8-3.7 times as long as broad; transverse cirri project distinctly
beyond posterior end of cell (Fig. 69a-l) ................... O. lanceolata (p 209)
Body in life usually less than 2.8 times as long as broad; transverse cirri project only
slightly beyond posterior end of cell (Fig. 72a, b, 75) ...................... 37
37 Body margins straight or slightly convex, sides usually nearly parallel, posterior end
broadly rounded (Fig. 72a-d, 116a-e) ..................................... 37a
Body margins strongly convex, outline broadly elliptical, posterior end often nar-
rowly rounded or slightly pointed (Fig. 75) ................. O. proximata (p 221)
37a 5 dorsal kineties (Fig. 72d) .............................. O. nauplia (p 215)
4 dorsal kineties (Fig. 116e) ................... Urosomoida antarctica (p 365)
38 (31) Saline water ................................................... 39
Freshwater ........................................................ 41
39 2 cirri right of transverse cirri and posterior end of right marginal row; body slightly
twisted (Fig. 82a, b, arrow; has tiny cortical granules, Fig. 246c) O. enigmatica (p 229)
No such cirri; body not twisted ........................................ 40
40 Length in life about 80-120 !lID, about 3.5 times as long as broad, outline more or
less parallel-sided (Fig. 85b) ............................. O. oxymarina (p 233)
Length in life about 65 !lID, about 2 times as long as broad, outline broad elliptical
(Fig. 80) ............................................... O. alfredkahli (p 227)
41 (38) Length in life about 130-180 !lID, outline distinctly ovoid (Fig. 38d, e, i) ..... .
· ...................................................... O. fallax (p 137)
Length in life about 70-120 !lm, outline not distinctly ovoid ................ 42
122 SYSTEMATIC SECTION

42 Caudal cirri distinctly longer than marginal cirri (for example, Fig. 44a) ....... 43
Caudal cirri not distinctly longer than marginal cirri (for example, Fig. 48) ..... 45
43 Only 2 rather long caudal cirri, which point somewhat to the right (Fig. 57a) .... 34
3 caudal cirri ...................................................... 44
44 Body in life about 2 times as long as broad (Fig. 50a, b; see also Sterkiella
histriomuscorum) ................................... O. monspessulana (p 168)
Body in life about 4.0-5.5 times as long as broad (Fig. 49a~) .... O. minor (p 166)
45 (42) 1 supernumerary cirrus immediately behind right frontal cirrus (Fig. 60) ...... .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Oxytricha sp. (p 190)
No such cirrus immediately behind right frontal cirrus ..................... 46
46 3 postoral ventral cirri, arranged rather close to proximal end of adoral zone of
membranelles (Fig.5Ia) ............................................. 47
Posterior postoral ventral cirrus distinctly displaced posteriad (Fig. 42d, arrow), or
usually more than 3 postoral ventral cirri (Fig. 62) ........................ 48
47 Transverse cirri usually more than 5 in number (Fig. 51a, b) ... O. multiseta (p 169)
Transverse cirri 5 in number (Fig. 48) ..................... O. matritensis (p 165)
48 (46) 5 dorsal kineties; ventral infraciliature rather variable (Fig. 62) .............. .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. O. variabilis (p 193)
6 dorsal kineties; ventral infraciliature not conspicuously variable (Fig. 41a-g,
42a-h) .............................................. 0. hymenostoma (p 150)
49 (20) Body in life about 2 times as long as broad, outline distinctly ovoid; transverse
cirri conspicuously displaced anteriad, do not protrude beyond posterior end of cell
(Fig.43a) ............................................... 0. kahlovata (p 157)
Body in life about 3.3 times as long as broad, outline long-elliptical; transverse cirri
protrude distinctly beyond posterior end of cell (Fig. 84c) O. parahalophila (p 170)
50 (3a) Anterior end cephalised; 180-240 ~m long (Fig. 81) ..... O. disci/era (p 227)
Anterior end not cephalised; 50-1 00 ~m long (Fig. 86a, 87a, 88a) O. saltans (p 236)

Oxytricha aeruginosa WRZESNIOWSKIEGO, 1866 (Fig. 31 a-e)

1866 Oxytricha aeruginosa WRZESNIOWSKlEGO, Wykaz Szkoty Glownej Warszawskiej, No.5: 18 (latin diag-
nosis, no illustration). I

1867 Oxytricha aeruginosa WRZESNIOWSKIEGO, Przyczynek do historyi naturalnej wymoczk6w, p 51 (Fig.

31b-e; in Polish).
1867 Oxytricha aeruginosa WRZESNIOWSKIEGO, Rocznik ces. kr61. Towarzystwa Naukowego Krakowskiego,
12: 279 (Fig. 31 b-e; in Polish).
1870 Oxytricha aeruginosa WRZESNIOWSKI, Z. wiss. Zoo\., 20: 471 (in Gennan).
1882 Oxytricha aeruginosa, WRZ. - KENT, Manual Infusoria II, p 787.
1932 Oxytricha aeruginosa WRZESNIOWSKI, 1870 - KAHL, Tierwelt Dtl., 25: 602.
1972 Oxytricha aeruginosa WRZESNIOWSKI, 1869 - BORROR, 1. Protozoo\., 19: 13.

Nom e n c I a t u rea n d t a x 0 nom y: Oxytricha aeroginosa WRZESNIOWSKI in

MAUCH (1976, p 435) is an incorrect subsequent spelling. Oxytricha aeruginosa was pub-
 Oxytricha 123

Fig. 31a-e Oxylricha aeruginosa (a, from CHORIK 1968; b--e, after WRZESNlOWSKIEOO
1867a. &-e, from life). a-c) Ventral views, a = 150 11m, b, c = 160 11m. In (b) 1 fronto-
ventral cirrus is arranged very close to the right frontal cirrus, which is very likely a
misobservation, in (c) only 3 frontoventral cirri are present. d) Lateral view, 155 11m.
e) Resting cyst, 80 11m; p 122. Fig.3lf Oxylricha sordida, a supposed synonym of O.
aeruginosa (after WRZESNIowsKlI861). Ventral view from life, about 110 11m; p 125.

lished four times as species nova! The text and the illustrations by WRZESNIOWSKIEGO
(1867a, b) are identical. Following I<AHL (1932), later always incorrectly dated with
1870, the year the Gennan version was published. The correct name in I<AHL (1932) is
Oxytricha (Oxytricha) aeruginosa because KAHL divided Oxytricha into subgenera. The
redescriptions by AGAMALIEV (1974; Fig. 221a, b) and PATSCH (1974; Fig. 220m) are in-
sufficient because no rusty granules are mentioned. A further population is descnbed by
CHORIK (1968, P 133) from small water bodies in Moldova (Fig. 3Ia); the reliability of
this Russian redescription was not checked. The figure in LoKOT (1987) is probably cop-
ied from I<AHL (1932).
Several more or less reddish Oxytricha species have been described, but none of
these has been synonymised with the poorly known O. aeruginosa. Oxytricha ru-
bripuncta is edaphic and has red granules in the cortex. Another edaphic species, O.
auripunctata, has orange-yellow cortical granules, usually only 4 transverse cirri, and is
somewhat smaller (80-130 !lm) than O. aeruginosa. The freshwater species, Rubrioxytri-
cha haematoplasma and R jerruginea, have a homogeneously reddish and rusty coloured
cytoplasm and cortical granules. Cyrtohymena muscorum also has red granules in the cy-
124 SYSTEMATIC SECTION

toplasm; however, it lives mainly in terrestrial habitats and the undulating membranes are
in Cyrtohymena pattern (Fig. 94a-<l, g, h). Detailed redescription necessary.

M 0 r p hoi 0 g y and b i 0 log y: In life about 120-165 x 45-60 !lm. Usually long,
sometimes broadly elliptical, both ends rounded. Anterior portion narrowed and slightly
bent to left. Two macronuclear nodules. Number of micronuclei unknown, but probably 2
or more. Contractile vacuole distinctly in front mid-body. Cytoplasm more or less in-
tensely coloured by russet and black granules ("Klirnchen"). WRZESNIOWSKI (1870) ob-
served that these granules are more numerous in well-feed specimens than in starved
specimens; thus, he supposed that they originate from digested food, for example
diatoms. Movement rapid.
Adoral zone of membranelles about 113 of body length, distinctly overlapping right
margin. Buccal area medium-sized, undulating membranes distinctly curved. 3-4 (proba-
bly usually 4) frontoventral cirri. Posterior postoral ventral cirrus located about in mid-
body. Five transverse cirri. Caudal cirri of about same length as marginal cirri. Cyst
spherical, approximately 80 !lm across, with a somewhat irregular, about 9 !lm thick (mu-
cous?) layer (Fig. 31e).

o c cur r e n c e and e colo g y: Rare, uncertain species; recorded from the benthos
of fresh, brackish, and marine waters; however, identifications never substantiated by de-
tailed data! Locus classicus is Warsaw, Poland, where WRZESNIOWSKIEGO (1866) discov-
ered Oxytricha aeruginosa in a spring in the garden of the University. CHORIK (1968; un-
certain!) found it in small water bodies in Moldova (Fig. 31a).
Records not substantiated by illustrations: running waters in Nordwnrttemberg,
brooks near Bonn, and Elbe Estuary, Germany (BUCK 1961, JUTRCZENKI 1982, RrnoEL-
LoRJE 1981); among detritus and in the surface film of ditches regularly contaminated
with pesticide sprays, and in unsprayed ditches in the Elbe flood plain, Germany (CASPERS
& HECKMAN 1982); Turiec River in Slovakia (TIRJAI(QvA 1993); Russia (LoKOT 1987);
Reelfoot Lake and Conestoga drainage basin, Pennsylvania, USA (BEVEL 1938, CAIRNS
1965a); occasionally among algae in ponds along Pine Lake, Michigan, USA (KOFOID
1896).
Several records from marine biotopes, for example, in the sub-littoral of the Bay of
Biscay (FERNANDEZ-LEBORANS & NOVILLO 1993), in low-salinity lagoons (maximum salin-
ity = 0.34 %) of Caspian Sea, Divichinskiy Estuary (USSR), and Baltic Sea (AGAMALIEV
1969, 1973, 1986; identifications questionable, see taxonomy; AGAMAUYEV 1974, AGA-
MALIYEV & ALlYEV 1983, PATTERSON et al. 1989, PETRAN 1977, SICH 1990).
Biomass of 106 individuals about 50 mg (NESTERENKO & KOVALCHUK 1991). MrnAI-
LOWITSCH (1989) found 0. aeruginosa (n = 17) in saline inland waters in Germany at
4.6-14.3 °C, pH 7.1-7.6, 39-156 mg I-I CO2 free, 3.5-17.6 mg I-I O2, 0.15-7.3 mg I-I
NH/-N, 0.02-0.2 mg I-I NOz--N, 1.1-13.4 mg I-I N03--N, 108-12763 mg I-I CI-, 90 to
3590 mS m- I specific conductivity. JENNINGS & MOORE (1902b) and WAGNER (1908) ob-
served spontaneous aggregations of o. aeruginosa on slides or even in small open dishes.
 Oxytricha 125

According to SLADECEK et al. (1981), WEGL (1983), and FOIssNER (1988a), Oxytricha
aeruginosa indicates strict betamesosaprobity (s = b; b = 10, I = 5, SI = 2.0). However, it
is poorly known and thus it can not be excluded that O. aeruginosa was confused with
other (common) reddish oxytrichids, like Rubrioxytricha haematoplasma. Thus, we ex-
cluded it from the list of water quality indicators (FOIssNER et al. 1991).

Sup po sed s y non y m: Oxytricha sordida WRZESNIOWSKI, 1861, AnnIs Sci. nat.,
Zoologie, 16: 332 (Fig. 311). Remarks: WRZESNIOWSKI (1861) mentioned that the cyto-
plasm is sometimes russet, indicating that it is Oxytricha aeruginosa, which he described
a few years later. The only significant difference is the position of the transverse cirri,
namely at the tip of the cell in 0. sordida (uncertain), and distinctly anteriorly displaced
in O. aeruginosa. STEIN (1862) assumed that O. sordida is identical with Tachysoma pel-
lionel/um. The name O. sordida was mentioned for the last time (?) by KEN'r (1882,
P 788), who, obviously erroneously stated that O. sordida, 0. aeruginosa, and Urosoma
macrostyla had been described by WRZESNIOWSKI (1870) in his account on the Infusoria
of Warsaw. WRZESNIOWSKI (1861) discovered O. sordida in a pond near Warsaw, Poland.

Oxytricha alfredi nom. nov. (Fig. 32)

1932 Oxytricha minor KAHL, Tierwelt Otl., 25: 602.

Nom e n cIa t u rea n d t a x 0 nom y: The correct name in the original description
is Oxytricha (Oxytricha) minor, which is a junior secondary homonym of O. minor (MAs-
KELL, 1887) KAHL, 1932 (see below). Thus, the species by KAHL (1932)
needs a new name: Oxytricha alfredi nom. nov. This species is almost
certainly not identical with Cyrtohymena gracilis (cortical granules pre-
sent, rather long caudal cirri; Fig. 961), as suggested by BORROR (l972a).
Oxytricha minor sensu MOTE (1954; Fig. 1841) is probably Sterkiella his-
triomuscorum, and the CHORIK (1968; Fig. 231h) population is insuffi-
ciently redescribed. Oxytricha minor sensu LoKOT (1987) is a redrawing
from KAHL's (1932) O. euglenivora. Poorly known species, thus detailed
redescription necessary.

M 0 r p hoI 0 g y: In life about 80 J-lm long and 2.8 times as long as

broad; according to DINGFELDER (1962; without illustration) 85-115 J-lm
long. Elliptical. Contractile vacuole slightly in front of mid-body. Adoral
zone of membranelles about 35 % of body length. Frontal cirri slightly Fig. 32 OX)!-
enlarged. Transverse cirri protrude somewhat beyond posterior end of tricha alfredi
cell. KAHL (1932) drew 2 slightly elongated cirri at posterior end of cell; (from KAHL
1932). Ventral
it is unknown if these are the posterior-most marginal cirri, or caudal view from life,
cirri. Dorsal cilia 2-4 11m long. 80 I'm.
126 SYSTEMATIC SECTION

o c cur r e n c e and e colo g y: Not vel)' common, in polluted lotic and lentic wa-
ter bodies. KAHL (1932) did not mention the type location. He found "Oxytricha minor"
in the sapropel, probably somewhere in Germany. Records from freshwater habitats not
substantiated by illustrations: Bulgarian running water (RUSSEV et al. 1976); side arm of
Danube River, rivers, and water reservoir in Czechoslovakia (MATIS 1961, 1977, MATIS
& TIRJAKovA 1994a); Turiec River in Slovakia (TIRJAKovA 1993); at pH 6.4-7.2 in rain
puddles near Erlangen, Germany, during spring (DINGFELDER 1962); ~O ind cm-2 in
mesosaprobic running waters in Northern Italy (MAnoNI 1980, 1983, 1984, MAnoNI &
GHEm 1977); slightly and heavily polluted waters in Barcelona, Spain (MARGALEF LOPEZ
1945); Latvian river (LIEPA 1973); peiagial of water reservoirs of Kiev and Kremenchug
and cooling plant in Moldova (CHORIK & VIKOL 1973, NEBRAT 1975, 1980); Utah, USA
(SHAWHAN et al. 1947); Conestoga drainage basin in Pennsylvania, USA (CAIRNS 1965a);
south-eastem Louisiana, USA (BAMFORTH 1963). Records from terrestrial habitats also
not substantiated by illustrations (BAMFORTH 1967, 1968, 1969, 1973, 1976, 1980a, 1984,
BAMFORTH & BENNETT 1985, LoUSIER & BAMFORTH 1990, MARTIN & SHARP 1983).
Feeds on bacteria and algae. Biomass of 106 individuals about 17 mg. MATIS (1975)
found 0. alfredi in road pools and in a drainage canal near Bratislava, Slovakia, at
I-31°C, pH 5.8-6.9, and 0.7-12.5 mg}-I O2• PATRICK et al. (1967) recorded it once from
the Savannah River, USA, at 5-<10 mg I-I Cl-, 5-10 mg I-I CO2, >5-7 mg 1-1 0 2, 0.1-{).4
mg I-I Fe2+, 3-<10 mg I-I Ca2+, 0.03-{).05 mg I-I NH3-N, <0.007 mg I-I NOi-N, 0.07-{).2
mg I-I N03--N, 0.05-{).1 mg I-I POlo, 10-<30 mg I-I SOlo, 23-28 °C, 0.5 to l.0 mg I-I
BODs. REINNARTH (1979) found O. alfredi in the mud-water interface ofa eutrophic pond
in Germany at +1 to -2 cm depth, pH 7.l-7.3, -113 to +210 mV redox potential,
0.07-2.5 mg I-I NH/-N, and 1.5-5.0 mg I-I N03--N. MATIS & STRAKovA-STRIEilKOvA
(1991) found it in a brook and in two thermal lakes in Czechoslovakia at 17-42 °C and
pH 5.0-7.5. MAUCH (1976) mentioned one reference where O. alfredi was found in
sewage-polluted water.

Oxytricha balladyna SONG & WILBERT, 1989 (Fig. 33a-f, Table 15)

1932 Balladyna similis KAHL, Tierwelt Dtl., 25: 592 (Fig. 331).
1972 Holosticha similis (KAlli., 1932) - BORROR, J. Protozool., 19: 11.
1989 Oxytricha balladyna (KAHL, 1932) - SONG & WILBERT, Lauterbomia, Heft 3: 166 (Fig. 33~).
1991 Oxytricha balladyna SONG & WILBERT, 1989 - FOiSSNER, BLATTERER, BERGER & KOHMANN, Infonna-
tionsberichte Bayer. Landesamtes fUr Wasserwirtschaft, 1191: 299.

Fig. 33a-£ Oxytricha balladyna (~, from SONG & WILBERT 1989; f, from KAlli. 1932. a, c, f, from life; b, d, ~
e, protargol impregnation). a, 1) Ventral views, a = 40 11m, f= 60 11m. b) Ventral infraciliature, 44 11m. Arrow
marks anterior postoral ventral cirrus. c) Lateral view. d, e) Dorsal infraciliature, d = 40 11m, e = 45 11m. Ac-
cording to SONG & WILBERT (1989), dorsal kineties 4 and 5 are only a single kinety which is interrupted in mid-
body. Be = buccal cirrus, ee = caudal cirri, Fve = posterior frontoventral cirrus, Mi = micronucleus, RMR =
right marginal row, 1-5 = dorsal kineties 1-5.
 Oxytricha 127

: 0.
f

1 : 3
: 2 :

,
1\ 5
/1
,:' \\
I I
\ I
\ I
I
, , I
\ J
\'
I'

/
\ : :.j
c CC~~e
128 SYSTEMATIC SECTION

Nom e n cIa t u rea n d t a x 0 nom y: BORROR (1 972a) transferred it to Holosticha

WRZESNIOWSKI, 1877. Since he classified H. similis STOKES, 1886, in Keronopsis, H%sti-
cha similis (KARL, 1932) was not a junior secondary homonym in his revision. Sm.LER
(l974a, p 132) proposed, under the headline "Balladinella gen. n.", that Balladynafusi-
formis KAlIL, 1932 and B. similis had to be separated from the other Balladyna species
because the transverse cirri are longitudinally arranged and the peristomial lip is anteri-
orly curved. According to CoRUSS (1979), the second original spelling in STILLER (1974a,
P 130), namely "Balladynella n. n.", is the correct one. However, since the establishment
of the genus is not accompanied by the fixation of a type species, Balladynel/a (or Ba/-
ladinel/a) is invalid (leZN 1985, Article 13 b) and both names must be considered as no-
mina nuda. Since no redescription of Ba/ladyna fusiformis is available I do not establish
a new genus. To avoid a secondary homonymy with Oxytricha simi/is ENGELMANN, 1862,
SONG & WILBERT (1989) introduced the replacement name "ba/ladynd'. Thus they them-
selves - and not KAlIL (1932) - are the authors of the present species (if it is classified in
Oxytricha). Possibly a synonym of O. opisthomuscorum (p 217,925).
The outline, the long caudal cirri, and the size of the population of SONG & WILBERT
(1989) agree well with the original description. In contrast, the number of frontoventral
cirri and the arrangement of the transverse cirri differs significantly. Thus, I am not sure
if these two popUlations are conspecific. However, it cannot be excluded that the devia-
tions are due to the small size, which makes live observations very difficult Oxytricha
balladyna sensu SONG & WILBERT (1989) is easily confused with the very similar O.
setigera (cp. Fig. 33a with Fig. 47a, 48a). Thus, an identification should always be con-
firmed by very careful live observations (lOOOx, differential interference contrast) or on
protargol-impregnated material; they differ significantly in the position of the buccal cir-
rus and the number of dorsal kineties. Further investigation recommended.

M 0 r p hoi 0 g y and b i 0 log y: About 60-80 JIm long in life according to KAlIL
(1932); 40-60 x 20-25 JIm according to SONG & WILBERT (1989). Long oval, posterior
end rounded; according to SONG & WILBERT (1989) narrowly rounded or slightly tapered.
The remaining description is based on the data by SONG & WILBERT (1989). For the KAlIL
(1932) population, see Figure 33f. Invariably (n = 24) only 1 micronucleus (not shown in
Fig. 33f) between the 2 macronuclear nodules, reminiscent of Balladynafusiformis. Con-
tractile vacuole about in mid-body, during diastole with inconspicuous collecting canals.
Cytoplasm bright and colourless, cortical granules absent. Rapid, somewhat jumping
movement.
Adoral zone of membranelles about 36 % of body length. Undulating membranes
short, intersecting, and only inconspicuously curved. Anterior portion of peristomial lip
conspicuously curved according to original description (Fig. 33f). Invariably 18 (n = 14)
frontal, ventral, and transverse cirri. All frontoventral cirri arranged right of adoral zone
of membrane lIes, posterior one at about level of buccal vertex (Fig. 33b). Frontal and
transverse cirri slightly enlarged. Transverse cirri arranged in an almost longitudinally
orientated hook (however, see KAHL's illustration), protrude distinctly beyond posterior
 Oxytricha 129

end of cell. Marginal rows begin slightly behind level of buccal vertex, distinctly sepa-
rate posteriorly. Dorsal cilia spreading, in life about 10 /lm long. According to SONG &
WILBERT (1989) invariably 4 (n = 20) dorsal kineties, the fourth being interrupted in mid-
body. In my opinion, however, 5 kineties are present, the fourth being the posterior por-
tion ofkinety 3, and the fifth is very likely a dorsomarginal kinety (compare with O. te-
nella). Invariably 3 (n = 14) prominent, in life about 20 /lm long caudal cirri at end ofki-
neties 1,2, and (probably) 4 (Fig. 33b, d, e).

o c cur r e n c e and e colo g y: Infrequent, probably confmed to freshwater. I<AHL

(1932) did not mention the type location; he found it in the sapropel, probably some-
where (near Hamburg?) in Germany. SONG & WILBERT (1989) isolated Oxytricha balla-
dyna from the Poppelsdorfer Weiher, a eutrophic pond near Bonn, Germany. Records not
substantiated by illustrations: Czechoslovakia (MAns & TIRJAKovA 1994a, TIRJAKovA
1992b, 1993); Latvian rivers (LIEPA 1973, 1978); sporadically in an alphamesosaprobic
and a polysaprobic area of a mountain brook in Yugoslavia (pRIMC 1984); freshwater
habitats in south-eastern Louisiana, USA (BAMFORrn 1963). Biomass of 106 individuals
about 6 mg (population of SoNG & WILBERT 1989).

Oxytricha chlorelligera KAHL, 1932 (Fig. 34a, b)

1932 Oxytricha chlorelligera KAlIL, Tierwelt 00., 25: 601 (Fig. 34a).
1968 Oxytricha chlorelligera KAlIL, 1932 - CHORIK, Free-living ciliates, p 133 (Fig. 34b).
1991 Oxytricha chlorelligera KAlIL, 1932 - FOISSNER, BLATfERER, BERGER & KOllMANN, Infonnationsberichte
Bayer. Landesamtes filrWasserwirtschaft, 1/91: 277.

Nom e n cIa t u rea n d t a x 0 nom y: The cor-

rect name in the original description is Oxytricha
(Oxytricha) chlorelligera because I<AHL (1932) di-
vided Oxytricha into several subgenera. Incorrect sub-
sequent spelling: Oxytricha chlorigella (HoRvArn
1939a, p 545; HORvArn 1942, p 4). The HORvArn
(1939b; Fig. 22li) population is insufficient rede-
scribed in that he did not mention the symbiotic algae,
the most important character of this species. The illus-
tration in DRAGESCO & DRAGESCO-KERNEIS (1986,
P 473, Planche 140H) is a redrawing from I<AHL
(1932). Detailed redescription necessary.

Fig. 34a, b Oxytricha chlorelligera (a, from KAlIL 1932; b, from

CHORIK 1968). Ventral views from life, a = lI5 ~m, b = 130 ~m.
The globules are symbiotic green algae.
130 SYSTEMATIC SECTION

M 0 r p hoi 0 g y and b i 0 log y: In life about 115 Jlm long and 3-3.5 times as long
as broad. Anterior end slightly bent to the left. Macronuclear nodules only slightly sepa-
rate; micronuclei not observed, according to KARL (1932) probably only one present.
Contractile vacuole about in mid-body. Rapid movement. Adoral zone of membranelles
113 of body length. 18 frontal-ventral-transverse cirri. Frontal cirri distinctly enlarged.
Five transverse cirri, only the right two protrude slightly beyond posterior end of cell.
Probably 3 inconspicuous caudal cirri. Dorsal cilia 2-4 Jlm long.

o c cur r e n c e and e col 0 g y: Very probably confined to freshwater. KARL

(1932) did not mention the type location; he found Oxytricha chlorelligera among
Utricularia, very likely somewhere (near Hamburg?) in Germany. CHORIK (1968) ob-
served it in small water bodies in Moldova.
Records not substantiated by illustrations: pond in Czechoslovakia (HAsSDENTEUFEL-
ovA-MoRAvcovA 1955); bank of Danube River in Czechoslovakia (TIRJAKovA 1992a);
pond in botanical garden of Kiel University, and small, stagnant water bodies in the Kot-
ton forest near Bonn, Germany (Ax & Ax 1960, BrcK 1957, 1958); Lake Balaton, Kala-
nos brook, and Danube River, Hungary (GELLERT & TAMAs 1959b, MATIS & TIRJAKovA
1995, VOROSVAAY 1950); water reservoirs in the Ukraine (KRAvCHENKO 1969); low-
salinity lagoons of Caspian Sea (AGAMALIEV 1973, 1986, AGAMALlYEV & ALlYEV 1983);
reservoirs and Araks River in Azerbaijan (ALEKPEROV 1988, 1990, ALIEV 1982a); Lake
Dzhandar, Azerbaijan? (ALIEV 1988); cooling plant in Moldova (CHORIK & VIKOL 1973);
oligotrophic lakes and rivers in Latvia (LIEPA 1973, 1990, LlYEPA 1984a, VEYLANDE & Lr-
YEPA 1985); Russia (LOKOT 1987; with illustration, however, probably a redrawing); stag-
nant water bodies of the Tibetan Plateau (WANG 1977); China (NING et al. 1993, YANG
1989); pond, pools, and lentic areas along stream margins in the USA (LACKEY 1938a);
ponds in Cameroon (DRAGESCO & DRAGESCO-KERNEIS 1986, NJINE 1977, NJINE & DEVAUX
1977).
Feeds on bacteria, flagellates, and diatoms (VOROS vARy 1950). Biomass of 106 indi-
viduals about 35 mg (ForssNER et al. 1991). BICK (1957) found O. chlorelligera in a de-
ciduous forest pond only during winter, when the pond was not shaded and therefore
enough light was available for assimilation of the symbiotic algae; he provided the fol-
lowing autecological data: pH 6.4-7.0,0.2-2 mg 1.10 2,0-0.7 mg I-I N03--N, 0-2.3 mg 1-1
NH/-N, 0-1.1 mg 1-1 HzS. HEMPSTEAD & JAHN (1940) recorded it from small, shallow
pools of Silver Lake Bog (Iowa, USA) at 14.2 mg 1-1 HzS! PATRICK et al. (1967) found
Oxytricha chlorelligera in the Savannah River, USA, at 20-23 °C, 1-3 mg I-I Cl-, 7 to
9 mg 1-1 O2, 1-<5 mg 1-1 BOBs, 3-<10 mg 1-1 Ca2+, 0.07-0.2 mg 1-1 N03--N, 0.001 to
<0.007 mg 1-1 N02--N, 0.017-0.033 mg 1-1 PO/-P, 1->10 mg I-I SOl-. Tolerates a maxi-
mum salinity of 0.2 % (Ax & Ax 1960).
According to SLADECEK et al. (1981), WEGL (1983), FOISSNER (1988a), and FOISSNER et
al. (1991), Oxytricha chlorelligera is confmed to alphamesosaprobic areas: a; a = 10, I =
5, SI = 3.0. However, this has to be verified by further autecological and saprobiological
data on reliably determined populations.
 Oxytricha 131

Table 15 Morphometric data of Oxytricha auripunctata (aur, from BLATIERER & FOISSNER 1988), O. balla-
dyna (bal, from SONG & WILBERT 1989), Oxytricha sp., 0. enigmatica, O.jaurei (dsp, eni, fau, from ORAGESCO
& ORAGESCO-KERNEIS 1986),0. geleii (gel, from WILBERT 1986b), O. granulifera (gra, from FOISSNER & ADAM
1983b), O. islandica (isl, from BERGER & FOISSNER 1989a), 0. lanceolata (lal, from BERGER & FOISSNER 1987a;
1a2, from BERGER & FOISSNER 1989; 1a3, from FOISSNER 1996b), O. longa (Igl, from GaEl & SzAaADOS 1950,
sublimate fixation; 192, IgJ, population 3, respectively 4 from GANNER et al. 1987b), O. longigranulosa (101,
from BERGER & FOISSNER 1989a; 102, from BLATIERER & FOISSNER 1988), O. naup/ia (nau, from BERGER &
FOISSNER 1987a), O. ottowi (ott, from FOISSNER 1996c), O. pseudofusiformis (psf, from DRAGESCO & DRAGESCO-
KERNElS 1986), O. pseudosimilis (pss, from HEMBERGER 1985), O. quadrici"ata (qua, from BLATTERER &
FOISSNER 1988), O. rubripuncta (rul, from BERGER & FOISSNER 1987a; ru2, from SHIN 1994), O. saltans (sal,
from SONG et aI. 1991),0. setigera (sel = population from the Glockner area, se2 = population from the Tull-
nerfeld, both from FOISSNER 1982; se3, from GsCHWIND 1991), O. siseris (sis, from FOISSNER 1982), O. tenella
(ten, from SoNG & WILBERT 1989). All data are based on protargol-impregnated material unless otherwise indi-
cated. All measurements in micrometres. ? = sample size not indicated; if only 1 value is known it is listed in
the mean column, if 2 values are available they are listed as Min and Max. CV = coefficient of variation (in
%), Max = maximum value, mean = arithmetic mean, Min = minimum value, n = sample size, SO = standard
deviation

Character Species mean SO CV Min Max n

Body, length aur 96.0 9.8 10.2 83 1I6 14
bal 40.4 2.9 7.2 35 44 14
dsp 95.0 84 105 6
eni 109.0 75 128 II
fau 143.0 92 172 16
gel 51.1 5.7 ILl 42 63 12
gra 80.6 8.9 11.1 70 103 25
isl 93.8 12.1 13.0 72 1I5 10
lal 87.9 7.3 8.3 75 100 12
1a2 94.5 12.2 12.9 70 108 10
1a3 93.5 7.1 7.6 87 1I0 II
Igl 140.0 I
192 58.9 7.1 12.0 47 75 25
IgJ 60.6 9.6 15.8 39 76 25
101 87.6 8.5 9.6 72 98 13
102 75.7 6.3 8.3 65 83 10
nau 63.3 8.1 12.7 56 80 10
ott 92.2 8.4 9.1 78 103 14
psf 41.0 35 46 6
pss 95 120 ?
qua 64.9 6.7 10.3 57 79 15
rul 140.1 11.8 8.5 1I2 160 15
ru2 134.9 19.9 14.7 100 170 12
sal 76.1 6.1 8.1 64 86 16
sel 37.2 3.9 10.5 30 43 10
se2 47.4 4.5 9.4 40 54 10
se3 36.9 2.9 8.0 30 42 21
sis 76.2 6.2 8.2 64 86 II
ten 50.9 4.7 9.1 46 60 9
Body, width aur 24.2 2.5 10.5 21 29 14
bal 19.8 2.3 11.7 17 24 14
dsp 33.0 23 36 6
132 SYSTEMATIC SECTION

Table 15 Continued

Character Species mean SD CV Min Max n

Body, width eni 41.0 32 56 11
fau 42.5 34 52 14
gel 23.1 2.8 12.2 17 26 12
gra 33.6 4.6 13.8 28 43 25
isl 30.7 3.6 11.8 25 35 10
lal 30.8 3.5 11.4 27 39 12
1a2 34.2 3.4 10.0 29 42 10
laJ 34.9 4.8 13.8 28 46 11
Igl 44.0 1
192 19.4 5.0 25.9 13 30 25
IgJ 16.4 2.4 14.6 10 21 25
101 31.9 3.1 9.8 27 38 13
102 27.5 2.8 10.3 21 32 10
nau 26.1 2.6 9.8 21 29 10
ott 32.8 5.4 16.3 23 40 14
psf 17.0 16 19 6
pss 40 50 ?
qua 24.5 3.6 14.7 19 32 15
rul 45.1 4.6 10.1 34 52 15
ru2 47.5 8.7 18.3 33 62 12
sal 35.1 9.8 28.1 27 45 16
sel 13.6 1.4 10.5 12 16 10
se2 14.8 1.5 9.9 13 17 10
se3 15.1 1.1 7.4 14 17 21
sis 21.4 2.8 13.1 18 27 11
ten 26.3 3.9 14.9 21 31 9
Adoral zone of membranelles, length aur 26.7 1.8 6.6 22 29 14
bal 14.6 0.9 6.3 13 17 8
gel 16.8 1.4 8.1 15 19 12
gra 26.5 1.9 7.2 24 32 25
isl 29.1 2.7 9.2 22 31 10
lal 27.5 1.1 3.9 25 29 12
1a2 30.1 3.5 11.6 24 34 10
laJ 29.4 1.4 4.6 27 32 11
101 28.1 1.3 4.7 25 30 13
102 26.7 2.2 8.1 23 30 10
nau 21.1 1.5 6.9 18 22 10
ott 31.3 3.6 11.4 24 35 14
qua 24.3 1.3 5.5 22 27 15
rul 43.7 3.5 8.1 35 49 15
ru2 40.9 3.1 7.6 33 44 12
sal 18.4 2.1 11.2 14 21 16
sel ll.8 0.6 5.1 11 13 10
se2 13.0 1.0 7.7 11 14 10
se3 12.3 1.1 8.8 11 14 20
sis 26.4 2.1 8.1 24 31 11
ten 18.8 1.6 8.4 16 21 8
 Oxytricha 133

Table 15 Continued

Character Species mean SD CV Min Max n

Adoral membranelles, number aur 24.4 1.2 4.9 23 27 13
bal 16.2 0.6 3.8 15 19 7
dsp 38.0 30 43 6
eni 49.0 45 53 II
fau 46.0 30 SO 18
gel 19.9 0.9 4.5 18 21 13
gra 31.2 1.3 4.1 29 32 25
isl 26.7 2.3 8.5 21 29 to
lal 26.5 1.5 5.5 23 28 12
1a2 27.6 0.8 3.1 26 29 to
1a3 27.9 1.4 5.2 26 30 II
Igl 21 22 'I
192 19.9 l.l 5.8 18 22 25
193 16.7 0.9 5.5 IS 18 24
101 26.5 l.l 4.0 24 28 13
102 22.0 1.3 6.1 20 24 10
nau 24.5 1.7 6.7 22 27 10
ott 25.5 2.7 10.4 21 29 14
psf 15.0 12 18 6
pss 21.0 'I
qua 19.7 0.6 3.0 19 21 IS
rul 39.2 1.9 4.9 36 42 IS
ru2 28.1 1.4 4.9 26 30 12
sal 15.1 1.7 11.2 13 18 14
sel 14.7 1.3 9.1 13 18 10
se2 14.9 0.8 5.6 14 16 10
se3 15.2 0.5 3.4 14 16 18
sis 25.2 2.5 9.8 22 30 11
ten 26.0 1.8 7.0 23 28 7
Right marginal row, number of cirri aur 18.4 1.7 9.2 15 21 14
bal 6.3 0.9 14.9 6 8 12
dsp 31.0 26 41 6
eni 27.0 20 32 11
fau 37.0 24 45 14
gel 21.9 2.1 9.6 18 25 15
gra 32.1 2.5 7.7 29 41 25
isl 25.6 2.2 8.5 21 28 10
lal 28.4 2.0 7.1 25 32 12
1a2 28.5 1.9 6.7 26 32 10
1a3 28.6 2.2 7.6 25 33 11
Igl 20 22 'I
192 20.4 1.8 8.9 16 23 25
193 16.4 1.8 10.7 14 20 25
101 25.9 1.6 6.1 23 28 13
102 19.1 1.5 8.0 17 22 10
nau 21.5 1.8 8.6 19 24 10
ott 24.6 1.8 7.4 22 27 14
psf 9.0 7 II 6
134 SYSTEMA TIC SECTION

Table 15 Continued

Character Species mean SD CV Min Max n

Right marginal row, number of cirri pss 15.0 14 17 ?
qua 15.1 1.0 6.9 14 17 15
rul 28.6 2.4 8.5 24 32 15
ru2 22.8 1.6 7.1 20 26 10
sal 5.6 0.8 14.6 5 7 16
sel 4.3 0.6 14.9 3 5 10
se2 4.6 0.8 17.4 3 6 10
se3 7.7 0.4 5.8 7 8 27
sis 11.5 1.2 10.0 10 14 11
ten 25.6 1.5 5.9 23 27 7
Left marginal row, number of cirri aur 17.7 2.1 12.0 15 22 14
bal 8.3 0.8 9.9 7 9 12
dsp 23.0 17 31 6
eni 24.0 20 27 9
fau 41.0 31 47 13
gel 20.5 1.8 8.8 18 24 15
gra l 34.0 2.6 7.6 30 43 25
isl 26.2 2.9 11.2 22 32 10
lal 31.4 2.6 8.4 27 35 12
1a2 31.7 3.6 11.3 25 36 10
1a3 30.0 2.6 8.8 25 34 11
Igl 19 21 ?
192 19.7 1.8 9.1 17 23 25
193 15.8 1.7 10.6 13 20 25
101 22.9 1.4 6.2 21 25 13
102 19.1 1.7 8.7 17 22 10
nau 20.8 1.7 8.1 18 23 10
ott 22.5 2.2 10.0 19 26 14
psf 8.0 7 9 6
pss 15.0 14 15 ?
qua 15.9 1.6 9.8 13 18 15
rul 29.4 1.7 5.7 26 32 15
ru2 22.9 1.8 7.8 20 26 10
sal 9.2 0.7 7.1 8 10 16
sel 6.5 0.7 10.3 6 8 10
se2 6.4 0.5 7.6 6 7 10
se3 7.5 0.6 7.8 6 8 25
sis 14.2 1.2 8.4 12 17 11
ten 22.3 1.9 8.5 19 25 7
Transverse cirri, number (pretrans- Igl 6.0 1
verse ventral cirri included) 192 6.0 0.4 6.8 5 7 25
193 5.7 0.6 11.0 4 6 25

I Caudal cirri included.

 Oxytricha 135

Oxytricha crassistilata KAHL, 1932 (Fig. 35)

1932 Opisthotricha crassistilata KAHL, Tierwelt Dtl., 25: 609.

Nom e n c I at u rea n d t a x 0 nom y: KAHL (1932) classified Opisthotricha as

subgenus of Oxytricha; thus, the correct name in the original description is Oxytricha
(Opisthotricha) crassistilata. Incorrect subsequent spellings: Oxytricha crasistillata
(KAHL) BoRR. (TIRJAKovA & MATIS 1987a, b, P 9, 22); Oxytricha crasistylata KAHL (MA-
TIS 1975, P 586); Opisthotricha crassistylata (STILLER 1974b, P 145, 146). BORROR
(1972a) synonymised Oxytricha crassistilata, O. opisthomuscorum, and O. eugienivora,
because they all have a single micronucleus, long dorsal cilia and caudal cirri, and en-
larged transverse cirri. I consider them three valid species because there are reliable dif-
ferences in the habitat, the body shape, and the shape of the transverse cirri; however, de-
tailed redescriptions are needed. Oxytricha crassistilata sensu ALEKPEROV (1993) from
Russian soil has two micronuclei, which does not match the original description (Fig.
241a, b). Oxytricha crassistilata resembles Tachysoma pellionellum, which, however,
lacks caudal cirri.

M 0 r p hoi 0 g y: In life 90-150 /.lm long, about 3.5 times as long as broad. Irregularly
elliptical; left margin convex, right one straight or convex in well feed specimens. Con-
tractile vacuole slightly above mid-body. Well feed specimens with dark granules. Adoral
zone of membranelIes about 35 % of body length, membranelIes prominent. Buccal area
conspicuously small, anterior portion obviously covered by a conspicuous cytoplasmic
lip. All 18 frontal, ventraL and transverse cirri prominent; especially transverse cirri
strongly enlarged, protrude distinctly beyond posterior end of cell. Marginal cirri "long".
Dorsal cilia 8-10 /.lm long. Three conspicuously elongated, soft caudal cirri.

o c cur r e n c e and e colo g y: Probably confmed to freshwater. KAHL (1932) did

not mention the type location; he found O. crassistilata among decaying blades of grass
and Glyceria, probably somewhere (near Hamburg?) in Gennany. Re-
cords not substantiated by illustrations: at 5-28°C, pH 5.8-6.3, and
6.2-12.4 mg 1-1 0 2 in road pools near Bratislava and in submerged and
wet mosses in Slovensky raj, Czechoslovakia (MATIS 1975, TIRJAKovA
& MATIS 1987a); rare in Maritza River, Bulgaria (DETCHEVA 1981);
Ukraine (KOVALCHUK 1997); China (NING et al. 1993).
Feeds on flagellates. Biomass of 106 individuals about 36 mg.
DETCHEVA (1983c) found O. crassistilata with low frequency (0.3 %)
in an alphamesosaprobic section of the Maritza River, Bulgaria, under

Fig. 35 Oxytricha crassistilata (from KAHL 1932). Ventral view from life, 120 11m.
Note the single micronucleus between the macronuclear nodules, the prominent trans-
verse cirri, and the elongated caudal cirri and dorsal cilia.
136 SYSTEMATIC SECTION

following conditions: pH 7.4, 4.6 mg 1.1 02, 53 % Oz. saturation, 51 mg 1.1 Ca z+, 52 mg 1.1
Mgz+, 224 mg 1.1 HCO]', 34 mg 1.1 SOl', 14 mg 1.1 Cl', 19.2 °dH, 0.6 mg 1.1 NH/-N,
0.05 mg 1.1 NOz·-N.

Oxytricha elongata (SMITH, 1897) KAHL, 1932 (Fig. 36)

1897 Opisthotricha elongata SMITH, Trans. Am. microsc. Soc., 19: 65 (Fig. 36).
1932 Opisthotricha elongata SMITH, 1897 - KAIn., TielWelt Dtl., 25: 609.

Nom e n c I a t u rea n d t a x 0 nom y: KAliL (1932) classified

Opisthotricha as subgenus of Oxytricha; the correct name in his revi-
sion is thus Oxytricha (Opisthotricha) elongata. In the rather similar
Oxytricha longissima all marginal cirri are of about the same length,
whereas they are distinctly elongated posteriorly in O. eiongata. BOR-
ROR (l972a) synonymised O. eiongata with O. simi/is, which is, how-
ever, distinctly smaller (80-100 11m) and has only 2 caudal cirri.
Doubtful species; detailed redescription necessary.

M 0 r p hoi 0 g y: In life 250-320 11m long (incorrect measure-

ment?), about 3 times as long as broad. Body margins slightly con-
verging posteriorly, both ends broadly rounded. Contractile vacuole
about in mid-body. Rapid movement. Adoral zone of membranelles
about 113 of body length. Arrangement of frontal and ventral cirri in-
Fig. 36 Oxytricha
sufficiently observed, probably typically oxytrichid. Five transverse
elongata (after cirri, do not protrude beyond posterior end of cell. Marginal cirri
SMI11I 1897a). Ven- very much longer posteriorly than anteriorly; number very likely dis-
tral view from life, tinctly underestimated in Figure 36 because 11 right marginal cirri
250-320 11m. Ar- seem very few for an about 300 11m long species. Three caudal cirri
rangement of cirri
obviously incor- of about same length as posterior marginal cirri. Dorsal cilia likely
rectly observed; short.
however, because
of large size con- o c cur r e n c e: Locus classicus is a pond in Louisiana, USA (SMITII
sidered as valid 1897a). KAliL (1932) found similar, but somewhat smaller (about
species.
200 11m) specimens in the sapropel (in Germany?). Not found since.

Oxytricha euglenivora KAHL, 1932 (Fig. 37a, b)

1932 Opisthotricha euglenivora KAHL, TielWelt Dtl., 25: 609 (Fig. 37a).
1968 Oxytricha (Opisthotricha) euglenivora KAHL, 1932 - CHORIK, Free-living ciliates, p 135 (Fig. 37b).

Nom en c I at u rea n d t a x 0 nom y: KAHL (1932) classified Opisthotricha as

subgenus of Oxytricha; the correct name in the original description is thus Oxytricha
 Oxytricha 137

(Opisthotricha) euglenivora. BORROR

(1972a) synonymised it with 0. crassistilata
(Fig. 35) and O. opisthomuscorum (Fig. 73a,
b). However, I follow KAHL (1932) and con-
sider them to be three valid species, although
the differences are admittedly not very
prominent (see key). Detailed redescription
necessary.

M 0 r p hoi 0 g y: In life about 80-90 11m

long and 2-2.5 times as long as broad. Well-
feed specimens spindle-shaped or ovoid.
Slightly contractile. Contractile vacuole dis-
tinctly in front of mid-body. Adoral zone of
membranelles about 1/3 of body length. All
18 frontal-ventral-transverse cirri and mar-
ginal cirri distinctly enlarged. Transverse Fig. 37., b Oxytricha euglenivora (a, from KAlIL
1932; b, from CHoRIK 1968). Ventral views from
cirri rather steeply arranged, protrude dis-
life, a = 80 JIm, b = 90 JIm. Note single micronu-
tinctly beyond posterior end of cell. Three cleus; p 136.
conspicuously elongated caudal cirri. Dorsal
cilia about 10 11ID, posteriorly even up to
15 11m long.

o c cur r e n c e and e colo g y: According to KAHL (1932) common and some-

times abundant on old leaves with algae ("oligosapropel"); with reference to the low
number of records, however, obviously very rare. KAHL (1932) did not mention the type
location, but he probably found Oxytricha euglenivora somewhere (near Hamburg?) in
Germany. CHORIK (1968) observed it in small water bodies in Moldova. LoKOT (1987; er-
roneously designated as O. minor KAHL) found it in freshwater bodies of the Lake Baikal
area, Russia. Records not substantiated by illustrations: fishponds in Tomky, Czechoslo-
vakia (MATIS 1973); small, meso- to eutrophic lake in Germany (MOCKE 1979); in plank-
ton of Goloseevo pond in Kiev, Ukraine (KOVALClRJK & SVINTSOVA 1994); Pantanal flood
plain, Brazil (HARooIN & HECKMAN 1996, HEcKMAN 1998). Feeds on algae, especially on
euglenoid flagellates (species name!). Biomass of 106 individuals about 30 mg.

Oxytrichafallax STEIN, 1859 (Fig. 38a-j, m-r, Table 17)

1859 Oxytrichafallax STEIN, Organismus der lnfusionsthiere I, p 189 (Fig. 38d-g).

1882 Oxytrichafallax. STEIN- KOWALEWSKIEGO, Pam. ftzyogr., 2: 413 (Fig. 38c).
1882 Oxytrichafallax, STEIN - KENT, Manual infusoria II, p 787.
1930 Oxytrichafallax STEIN - WANG, Contr. bioI. Lab. Sci. Soc. China, 6: 13 (Fig. 38b).
1932 Oxytrichafallax STEIN, 1859 - KAm., TielWeit Otl., 25: 602.
 138 SYSTEMATIC SECTION

Fig. 38a-c Oxytricha fallax

(a, from CURDS 1969; b, from
WANG 1930; c, after KOWALE-
WSKlEGO 1882). Ventral views
from life, a = 155 Ilm, b =
140 Ilm, c = size not indi-
cated; p 137.

1937 Oxytrichafallax SWIN - SOKOLOFF & ANCONA, An. lost Bioi. Univ. Mex., 8: 174 (Fig. 380).
1961 Oxytrichafallax S1EIN, 1859 - REUTER, Acta zoot. fenn., 99: 19 (Fig. 38h).
1962 Oxytrichafallax SWIN - LIEBMANN, Handbuch der Frischwasser- und Abwasserbiologie I, p 359 (Fig.
38i).
1968 Oxytrichafallax S1EIN, 1859 - CHORIK, Free-living ciliates, p 133 (Fig. 38m).
1969 Oxytrichafallox SWIN, 1859 - CURDS, Wat. Pollut. Res., No. 12: 67 (Fig. 38a).
1972 Oxyrichafa/lax S1EIN - BlcK, Ciliated protozoa, p 180 (Fig. 38j).
1982 Oxytrichafallax SlEIN, 1859 - HEMBERGER, Dissertation, p 142 (Fig. 38n).
1991 Oxytricha fallax SWIN, 1859 - FOISSNER, BLAlTERER, BERGER & KOHMANN, loformationsberichte des
Bayer. Landesamtes ftlr Wasserwirtschaft, 1/91: 279.
1993 Oxytricha hymenostoma STOKES, 1887 - SHIN & KIM, Korean J. Zoot., 36: 223 (Fig. 38p-r).
1994 Oxytricha hymenostoma STOKES, 1887 - SHIN, Dissertation, p 137 (Fig. 38p-r; mixed up the figures of
Rubrioxytricha haematoplasma and O. hymenostoma).

Nom e n c I at u rea n d t a x 0 nom y: The correct name in KAHL (1932) is Oxytri-

cha (Oxytricha) fallax because he divided Oxytricha into subgenera. Incorrect subse-
quent spellings: Oxitrichafallax (BAmsH 1992, p 57; Bauer 1983, p 38); Oxytrichafal-
lox STEIN (CAIRNS & RUTHVEN 1972, P 420; STOUT 1984, P 123); O.falax (BAKOWSKA &
JERKA-DZlADOSZ 1978, P 297).
BORROR (1972a) synonymised several species with O.fallax, for example 0. hymeno-
stoma (70-110 11m long) and 0. bimembranata (probably confmed to edaphic habitats).
This is not justified because only an about 150-200 11m large freshwater Oxytricha with
an oval outline may be identified with 0. fallax. Thus, the popUlations of FOISSNER
(1979a), GRIMES (1972 and his later works), and REYNOLDS (1932) are probably not con-
specific with O. fallax. The same is true for 0. fallax sensu HAsHIMOTO (1961, 1962,
 Oxytricha 139

Fig. 38d-j Oxytrichafallax (d--g, after STEIN 1859b; h, after REUTER 1961; i, from LIEBMANN 1962; j, after
BICK 1972a. d-i, from Iife;j, from life?). d) Ventral view ofa specimen with typical body shape, 148-173 11m.
e) Ventral view of a slightly contracted specimen. 1) Ventral view of a specimen creeping between detritus. g)
Cyst, size not indicated. b-j) Ventral views, h = size not indicated, i = 155 11m, j = 150 11m; p 137.

Fig.38k Insufficient redescription of Oxytrichafallax (after QUENNERSTEDT 1865). Ventral view from life, size
not indicated; p 137.

Fig. 381 Oxytricha hymenostoma (after REYNOLDS 1932). Ventral view, haematoxylin staining, 75-100 11m;
p ISO.
140 SYSTEMATIC SECTION

0
a
..
-
- - ;-

-
,. -:" : -
If."
., "'...
n

Fig.38m-o Oxytrichafallax (m, from CHORIK 1968; n, from HEMBERGER 1982; 0, from SoKOLOFF & ANCONA
1937. m, 0, from life; n, protargo1 impregnation). m,o) Ventral views, m = 160 "un, 0 = size not indicated. 0)
Ventral infraciliature, 195 Jlm; p 137.

1963, 1964), who identified his population according to the descriptions by BISHOP
(1943; classified as indeteIDlinable), HORvArn (1933; no illustration), and LUND (1935;
synonymised with O. hymenostoma); I classify the FOISSNER, GRIMES, REYNOLDS, and
HAsHIMOTO populations as O. hymenostoma. I found a specimen matching the original de-
scription in every respect in a betamesosaprobic brook near the town of Salzburg. Thanks
to its large size I was able to undertake several counts and measurements and the results
agreed very well with the data by HEMBERGER (1982). Oxytricha fallax sensu SCHMALL
(1976; P 38) is very likely a misidentified Histriculus histrio because "the marginal rows
are closed and caudal cirri are lacking" (Fig. 177q).
The Oxytricha hymenostoma STOKES of SHIN & KIM (1993b; see also SHIN 1994) is
distinctly larger (120-190 x 50-85 flm) than the type population described by STOKES
(l887b; about 100 flm). Thus, the Korean population is preliminary classified as O. fal-
lax and briefly described.
The populations by BARKER (1949; Fig. 232i), BATnSH (1992; his Fig. 54), BISHOP
(1943; Fig. 221f), CELA (1972; Fig. 232n), CHARDEZ (1967; Fig. 223p), CONN (1905; Fig.
221c), FORMISANO (1957; Fig. 234b), GREGORY (1923; Fig. 240n), HEBERER (1928; Fig.
229h), KOFFMAN (1926; Fig. 231d), LUNDIN & WEST (1963; Fig. 220k), QUENNERSTEDT
(1865; Fig. 38k), SMITH (1978; Fig. 221e; see also BLOCK 1984), and WEST & LUNDIN
 Oxytricha 141

(1963; Fig. 220k) are insufficiently redescribed. An identification as O. fallax should be

based mainly on the original description (Fig. 38d-f) until an authoritative redescription
is available.

M 0 r ph 0 log y and b i 0 log y: In life 130-180 J.lm long and about 2.5 times as
long as broad; according to HEMBERGER (1982) up to 200 x 110 J.lm. Long oval, posterior
end broadly, anterior often narrowly rounded (Fig. 38d). Rather contractile and very
flexible (Fig. 38f). Macronuclear nodules about 30-35 x 15-20 J.lm, fine granulated.
Usually 2 micronuclei, difficult to discern in life. The amicronucleate race reported by
WOODRUFF (1921) is probably an Oxytricha hymenostoma. Contractile vacuole slightly in
front of mid-body. Cortical granules lacking. Cytoplasm colourless; sometimes packed
with about 25 J.lIll sized food vacuoles causing dark appearance at low magnification.
Swims 2.3 body-lengths per rotation (BULLINGTON 1925, JENNINGS 1899b, LUDWIG 1929).
WOODRUFF (1907) reported on great variation of some characters (size, shape, number of
micronuclei...) during life cycle.
Adoral zone of membranelles about 40-45 % of body length, about 50 membranel-
les. Basis of membranelies about 12-14 J.lm broad. 18 frontal-ventral-transverse cirri ar-
ranged as shown in Figure 38d-f. Frontal and transverse cirri and buccal cirrus enlarged.
Frontal cirri in life about 25 J.lm long; transverse cirri in life about 40 J.lm, according to
HEMBERGER only 25 J.lm long, protrude slightly (left) to distinctly (right) beyond posterior
end of cell (Fig. 38d, e). About 25 left and 28-30 right marginal cirri. Dorsal cilia about
2-3 J.lm, according to HEMBERGER (1982) 4-5 J.lm long. Caudal cirri inconspicuous in life
because only slightly longer (25 J.lm) than marginal cirri (20 J.lm).
Population of SInN & KIM (1993b) and SInN (1994; described as. O. hymenostoma;
Fig. 38p-r): In life 120-190 x 50-85 J.lm; soft and flexible; subelliptical or oval; both
ends rounded; left anterior margin concave; 2 macronuclear nodules, after protargol im-
pregnation 18-20 x 10-11 J.lm; usually 2, rarely 3, micronuclei, in life(?) about 4 J.lm
across; contractile vacuole near mid-body; no cortical granules; adoral zone of mem-
branelles about 41 % (36-56 J.lm) of body length, 26-31 (mean = 28) membranelles; buc-
cal cavity deep; undulating membranes 22-38 J.lm long; almost invariably 18 frontal-
ventral-transverse cirri (sometimes only 4 postoral and pretransverse ventral cirri); ar-
rangement of postoral ventral cirri somewhat unusual (Fig. 38r); 14-20 (mean =17.6) left
and 20-23 (mean = 20) right marginal cirri; 6 (rarely 5) dorsal kineties, kinety 3 (in me-
dian) with about 15 basal body pairs; cilia 5 Ilm long; invariably 3 caudal cirri.
Cyst with smooth wall (Fig. 38g; STEIN 1859b). BRAND (1923) also figured a cyst,
however, the identification is not substantiated by an illustration of an excysted specimen.
REES (1878) described infections of the macronucleus by parasitic algae. SCHNEIDER
(1930) observed the formation of a cell envelope (= Hillle) after the addition of iodine-
ink (not substantiated by an illustration).
Morphogenesis commences with the proliferation of an oral primordium at the left
transverse cirrus (HEMBERGER 1982). TCHANG & HUANG (1983) provided a rather sche-
matic illustration of a middle morphogenetic stage showing 6 frontal-ventral-transverse
 142 SYSTEMATIC SECTION

,.
•

,•

,
,
0 ,
•••, •
f
,

Fig. 38p-r (}xylrichafallax (from SHIN 1994. p, from life; q, r, protargol impregnation). p) Ventral view,
116 Jim. q, r) Dorsal and ventral infraciliature, 120 Jim; p 137.

cirri streaks with 1,3, 3, 3, 4, and 4 cirri respectively (see also DILLER 1965a). TCHANG &
PANG (1979a, b), WOODRUFFF (1913), and ZHANG et aI. (1983) studied the nucleo-
cytoplasmic relationship and the phenomenon of macronucIear regulation in dorsiconju-
gants. A specimens regenerates within 5-6 min after being deeply cut in the middle re-
gion (IsHIKAWA 1912).
Although little is known about the morphology, a considerable number ofbiochemi-
cal, genetic, and other investigations exist. However, I doubt that the identification was
correct in all cases. For instance, the experiments by SCHLEGEL (1985) and SCHLEGEL &
STEINBROCK (1986) revealed that their 0. fallax differs significantly from the type species,
0. granulifera, whereas it agrees rather well with some other, probably Stylonychia spe-
cies. In spite of this uncertainty, some of these works are cited: AoL & BERGER (1995,
1996), CARTINHOUR (1984), CARTINHOUR & HERRICK (1984), DAWSON (1984), DAWSON &
HERRICK (1982, 1984a, b), DAWSON et al. (1984), DIlLER (1962, 1965a), DOAK et al.
(1994, 1997), GATES (1986), GRESLIN et al. (1988), HERRICK et al. (1985, 1987a, b),
HUNTER et al. (1989), JAHN (1966), KAINE (1982), KAINE & SPEAR (1980, 1982, see also
KRAUT et al. 1986), KRAMER (1988), MAUPAS (1889), MURTI (1976), PLUTA & SPEAR
(1981), PLUTA et al. (1982, 1984), PREER & PREER (1979), RAE & SPEAR (1978), RAIKov
(1989), SEEGMILLER et al. (1996, 1997), SPEAR (1980), STEINBROCK (1990), SWAGER et al.
(1990), SWANTON et al. (1982), TCHANG et al. (1981), VERNI & ROSATI (1980), WADA &
 Oxytricha 143

SPEAR (1980), WILLIAMS & HERRICK (1991), WILLIAMS et al. (1993), WITIlERSPOON et al.
(1997).

o c cur r e n c e and e colo g y: According to the faunistic data rather common in

freshwater; however, see taxonomy for uncertainty in species identification. Obviously
prefers eutrophic stagnant and slightly to heavily organically polluted running waters,
where O. fal/ax lives mainly in the aufwuchs, detritus, and in layers of filiform bacteria.
According to own experience much more seldom than, for example, Tachysoma pe/-
lionel/um or species ofthe Stylonychia mytilus complex.
Locus classicus is a brook in the "Procopthal" near Prague, Czechoslovakia, where
STEIN (1859b) discovered Oxytricha fallax together with Gonostomum affine between de-
caying leaves of trees. Records substantiated by illustrations: trout-rearing basins in Ger-
many (HEMBERGER 1982); Poland (KOWALEWSKIEGO 1882); small water bodies in Moldova
(CHORIK 1968); in great numbers in Lake Ho Hu in Nanking, China, during February
(WANG 1930); mesosaprobic water bodies in Mexico (SOKOLOFF & ANCONA 1937). SlllN &
KIM (1993b) and SlllN (1994) found O. fallax in the Han River in Seoul, Korea, in early
June and late August.
Records from freshwater habitats not substantiated by illustrations: Sphaerotilus
community in a river near Vienna, Austria (STRUHAL 1969; further records from Austria
see FOIssNER & FOISSNER 1988 and PErZ et al. 1988); fish farm in Belgium (CHARDEZ
1989); various, often polysaprobic water bodies in Belgium (BERVOErS 1940); rivers and
limnocrene in Bulgaria (DErCHEVA 1979b, 1981, 1991, 1993, DETSCHEWA 1972, RusEvet
al. 1988, RussEv et al. 1984); brook heavily polluted by brewery sewage in Czechoslova-
kia (VASICEK 1964); littoral of a side arm of Danube River and periodical pools with pH
5.8-6.0 in Czechoslovakia (MATIS 1961, 1975; further records from Czechoslovakia:
SRAMEK-HuSEK 1953, MATIS & TIRJAKovA 1994a); cooling system of a conventional
power station supplied with water from the beta- to alphamesosaprobic Rhine River in
Germany (BERNERTH 1982); plankton of Rhine River, Germany (CZERNIN-CHUDENITZ
1958); polysaprobic area in the WeiBe Elster River, Germany, downstream of a sewage
treatment plant (BEER 1958); upper layer of mud in Isebekkanal, Hamburg, Germany
(CASPERS & SCHULZ 1962); floating, foul-smelling Oscil/atoria mats in a side arm of
Mosel River, Germany (LAUTERBORN 1901); betamesosaprobic area of Elbe River and
polluted Hamburg Harbour, Germany (GRIMM 1968, HECKMAN et al. 1990); alphameso-
saprobic Oker River in Braunschweig, Germany (REICHENBACH-KLINKE 1959); throughout
the year with high abundance (30 indo ml- I ) in road pools with putrefying plants in Ba-
varia (DINGFELDER 1962); highly polluted Salzach River at the village of Laufen, Bavaria
(BAUER 1983); meso- to eutrophic lakes and pools near MUnster, Germany (MOCKE
1979); scattered in a polluted lake near Berlin, Germany, in February (MARSSON 1901);
Westphalian caves and mines in the Erzgebirge, Germany (GRIEPENBURG 1934, 1935,
WETZEL 1929); well of a garden on Heligoland, Germany (LAUTERBoRN 1894); oligo- to
betamesosaprobic area and pelagial of a branch of Danube River in Hungary (BERECZKY
1969, 1991, NOSEK & BERECZKY 1981); polluted Zagyva River, Hungary (HAMAR 1971);
144 SYSTEMATIC SECTION

6-8 indo ml- 1 sediment and in detritus-drifts in Lake Balaton, Hungary (GEIl.ERT 1961,
GELLERT & TAMAs 1958, 1959b); eutrophic soda lake (STILLER 1942) and together with
Dexiostoma campy/um, Trithigmostoma cucullu/us, and Paramecium caudatum in a
ditch near Szeged, Hungary (HORVATH & KUHN 1941); stalactite cave in north-east Hun-
gary (BAlOMI 1969; further records from Hungary: BERECZKY 1977b, HORVATH 1933, KRE-
PUSKA 1917, 1930); 48 indo cm-2 in a slightly polluted area ofPanne River, Northern Italy
(MAnoNI 1980, 1983, 1984, MAnoNI & GHETIl 1977); littoral area of Lake Como, Italy
(CATTANEO 1888; further records from Italy: CANELLA 1954, GRISPINI 1938); the Nether-
lands (VERSCHAFFELT 1930); with 33 % frequency in the Sphaerotilus natans community
of Lyna River, Poland (HUL 1986); heavily polluted river in Poland (GRABACKA 1988);
abundant in putrefying water in Warsaw, Poland (WRZESNIOWSKIEGO 1866); polluted
Lobregat River and other freshwater habitats near Barcelona, Spain (GRACIA & IGUAL
1987a, GRACIA et al. 1987a, 1989, MARGALEF LOPEZ 1945); Danube River, Slovakia
(SZENTIvAN¥ & TIRJAKovA 1994); Lake Geneva, Switzerland (ANDRE 1912, 1916, MER-
MOD 1914); ponds near Basle, Switzerland (RIGGENBACH 1922); Save River, Yugoslavia
(PRIMC 1981); reservoirs in the USSR (ALEKPEROV 1984b, c, BELOVA 1988, KRAVCHENKO
1969); Lake Baikal (GAJEVSKAJA 1927, GAJEWSKAJA 1933, ROSSOLIMO 1923; further re-
cords from the USSR: AGAMALIEV 1973, CHORIK & VIKOL 1973, DADAY 1903, 1904, LIEPA
1973, 1983, ZHARIKOV & ROTAR 1992); sediment ofChangjiang River, China (Gu et al.
1988; further records from China: NING et al. 1993, SHEN 1981, SHEN & GONG 1989,
YANG 1989); Tibetan Plateau (WANG 1977); pond in Japan (YOSHINO 1993, uncertain);
Costa Rica (Rmz 1961); Conestoga drainage basin, Pennsylvania, USA (CAIRNS 1965a);
Douglas Lake and Black River, Michigan, USA, at pH 6.5-7.0 and 20-24 °C (CAIRNS &
YONGUE 1966); New Jersey, USA (DILLER 1965a); among algae in Creel Bay, North Da-
kota, USA (EDMONDSON 1920); tree holes in Ohio, USA (LACKEY 1940); on polyurethane
foam in a shallow, eutrophic pond with hard and well-buffered water in Spokane County,
Washington, USA (PRATT et al. 1986); common in decaying Ceratophyllum from bottom
of Put-in Bay Harbour, Lake Erie, USA (JENNINGS 1899a); pond and river in Virginia,
USA (HENEBRY & CAIRNS 1980, YONGUE & CAIRNS 1979; further records from North
America: BOVEE 1960, McCASHLAND 1956, McCORMICK & CAIRNS 1990, MCCoRMICK et
al. 1991, WANG 1928, YONGUE & CAIRNS 1971); tributary of Amazon River, Colombia
(BOVEE 1957); moss infusion from Venezuela (SCORZA & NuNEz MONTIEL 1954); freshwa-
ter habitats in Tahiti (EDMONDSON 1910); Algeria (DADAY 1910); Coast Lake in Victoria
Land, Antarctica (THOMPSON 1972).
REUTER (1961, 1963a) found D./allax in rock pools with up to 20.5 % NaCI(!); how-
ever, he did not mention at which specific salinity it occurred. ZINN (1954) observed it in
a Typha-pond in Massachusetts at a salinity of about 1 %. BUTSCHINSKY (1897) and
ZACHARIAS (1888) found 0. /allax in saline lakes in the USSR and in Germany. KUMSARE
et al. (1974), LACKEY (1936, 1938a), LACKEY & LACKEY (1970), and MERESCHKOWSKY
(1877) recorded it from marine habitats. DIAZ PEREZ & MONTOTo LIMA (1989) found D.
lallax in the polluted Almendares River estuary (average salinity 1-1.5 %) at the north
coast of Havana City, Cuba. However, most of these records are not substantiated by iJ-
 Oxytricha 145

lustrations. Thus, the tolerance to high salinity has to be verified on reliably detennined
populations.
Oxytricha fallax occurred also in activated sludge, trickling filters, and rotating bio-
logical contactors (BAHR 1954, BANINA 1983, 1990, BARKER 1943, CURDS 1969, 1975,
KLIMOWICZ 1970, 1972, KUTIKOWA 1984, LIEBMANN 1962, LUNA-PABELLO et al. 1996, MA-
MAEVA 1983, SCHERB 1968b, STAMM 1979, 1984, WACHS 1968, 1984), and according to
LACKEY (l938a) also in Imhoff tanks. LACKEY (1938a) counted up to 500 indo ml'! in
trickling filters, LIEBMANN (1936) up to 400 indo ml'! in putrid waste water. KLIMOWICZ
(1973) found up to 284 indo ml-! at a sludge loading of about 0.06 kg BODs kg,l dry mass
d,l, Oxytrichafallax usually occurred during the maturation (second to fifth week) of the
sludge (CURDs 1965a, 1966b, HAMM 1964). KIusHNAMooRTIll & SARKAR (1981) found it in
Indian secondaIy stabilisation ponds with a BOD (BODs?) range of 50-80 mg 1'1, and
LuGO et al. (1991) found high numbers in a stabilisation pond in Mexico, SMlTI{ (1982)
recorded O. fallax from guano of penguins on the sub-Antarctic island of South Georgia;
the identification is, however, not substantiated by morphological data,
Records from terrestrial habitats are certainly misidentifications (BHATTACHARYA et
al. 1977, BUITKAMP 1977a, 1979, CAIRNS & RUTIIVEN 1972, KOFFMAN 1934, SMlTI{ 1972,
1973a, b, 1974a, 1975, 1985, STELLA 1948, STOUT 1961, 1963, 1968, 1970, 1978, 1984,
TAKAHASHI & SUHAMA 1991, VARGA 1936); however, the rather similar (synonymous?) O.
bimembranata was discovered in soil. SMlTI{ (1981) recorded O. fallax from turf peat
mosses (PH 4.9, water content about 56 % dry weight) from the Crozet Islands.
Feeds on bacteria (Bacillus suM/is, Pseudomonasfluorescens), diatoms, filamentous
growths, flagellates (Chi/omonas, Peranema trichophorum), ciliates (Uronema nigri-
cans, Dexiostoma campy/urn), and detritus (CURTIS & CURDS 1971, FAURE-FREMIET 1961a,
FOISSNER & SCHIFFMANN 1974, JOHNSON 1933, 1937, TAMAs & GELLERT 1958). Large food
vacuoles can contain more than 40 specimens of Uronema nigricans. Oxytricha fal/ax
may be grown on 0.1 % lettuce medium (GIESE 1959).
WOODRUFF (1905a, b; see also WOODRUFF & BAITSELL 1911) maintained a culture of
O. fallax for· over 860 generations and a shortest generation time of about 7 h was ob-
served. According to BAITSELL (1914), the division rate is usually below 2 divisions per
day (Russian readers, see also ZAn<E 1970). Biomass of 106 individuals about 155 mg
(FOISSNER et at 1991; calculated from average size), according to REUTER (1963b) 91 mg.
Some autecological data are summarised in Table 17. Further data: 13-24 mg 1'1
COD (MAroNI & GHETTI 1981a); betamesosaprobity and alphamesosaprobity, 38 to
72 mg 1"1 Ca2+, 7.3-26 mg I-I Mg2+' 116-348 mg 1'1 HC0 3', 16-104 mg I-I SOl, 10 to
38 mg 1"1 CI-, ~.08 mg 1'1 Fe2+ (DETCHEVA 1983c); 7.0-46.5 °C, pH 5.0-7.3 (MATIS &
STRAKOVA-STRIE~KOvA 1991). PATRICK et at (1967) found Oxytrichafallax twice in clean
areas of the Savannah River, USA, at >6-8 mg I-I O2, 0.5-1 mg 1'1 BODs, 0.005 to
0.04 mg I-I NH3-N, 0.01-0.03 mg I-I POl-Po PRIMc (1983) observed 0. faUax only at
temperatures of 10-15 °C. This agrees with the results by BIERLE (1970) who found opti-
mal growth at 14°C; however, he observed a thennoresistance of up to 35.7 °C. ISSEL
(1901, 1906, see also BRUES 1928, P 160) recorded it from Italian thennal waters at
146 SYSTEMA TIC SECTION

32-45 °C (identification doubtful because specimens only 69-86 ~m long); MATIS (1971)
found 27°C as broadest range between maximum and minimum temperature of occur-
rence.
Hydrogen-sulphide intolerant (WETZEL 1928b). Lethal doses (LDso4sh) of lead acetate
and lead nitrate was 200 mg I-I and 200-250 mg I-I, respectively; 300 mg I-I and 500
mg 1-1 killed O. fal/ax instantly, and 20-100 mg I-I lead acetate and 40-80 mg I-I lead ni-
trate, respectively, slowed down the growth pattern (KHAN & NASREEN 1981, NASREEN &
KHAN 1979). Lead accumulation was evident as deposition of black granules in the body
(FERNANDEZ-LEBBoRANs et al. 1985). 50 mg I-i of three anionic detergents killed most of
the specimens in 24 h, and 400 mg I-I in 1 h, respectively (MORENO-BARON & LoPEZ-
OcHOTERENA 1976). LAIRD (1958) found O. fallax in infusions with 2 ~g mI-1 DDT.
0.25 ~g mI- 1 desiccated venom (PH of solution 7.3, medium = 0.025 % beef extract) of
the Crotalus atrox rattlesnake had an immediate effect on locomotion; after 24 h animals
were alive, but feeble; all were dead after 48 h (PHILPOTT 1930). STATKEWITSCH (1904)
studied the effect of electrical current on 0. fallax.
Oxytrichafallax inhabits polluted waters (MARsSON 1903) and is classified as alpha-
mesosaprobic indicator of water quality: a; b = 1, a = 8, p = 1, 1=4, SI = 3.0 (FOISSNER
1988a, FOISSNER et al. 1991, SLAoECEK et al. 1981, WEGL 1983). The chemical data, how-
ever, indicate that the betamesosaprobic valence should be slightly increased (Table 17).
According to LIEBMANN (1962), Oxytrichafallax avoids water that is contaminated with
liquid manure. The autecological data and the saprobic classification should be checked
on reliably determined populations.

Oxytrichafaurei TucoLEsco, 1962 (Fig. 39a-j, Table 15)

1962 Oxytrichajaurei TUCOLESCO, Annis Spelt!ol., 17: 105 (Fig. 39g-j).

1986 Oxytrichajaurei (fUCOLESCO, 1962)1 - DRAGESCO & DRAGESCo-KEWIS, Faune tropicaIe, 26: 465 (Fig.
39a-f).

Nom e n c I a t u rea n d t a x 0 nom y: TUCOLESCO (1962a) established this species

as "0. faurd' under the headlines "Genre Oxytrichd' and "Sous-genre Opisthotrichd';
the correct name in the original description is thus Oxytricha (Opisthotricha) faurei and
the author's name is not be put in parentheses as suggested by DRAGESCO & DRAGESCo-
KERNElS (1986). These authors neither mentioned nor drew cortical granules. In addition,
their population differs from the original description in some other characters (ratio of
body length to body width, shape and relative length of buccal area, shape of posterior
end of left marginal row). Thus, the conspecifity of these two populations is doubtful.
Conversely, the descriptions agree well in that several elongated, fine caudal "cirri" are
present. Detailed redescription needed; possibly this worm-like species belongs to an-
other taxon.
 Oxytricha 147

,
#

.,, ,
,,
I

, .<
" ~:J

... ;
~ ,:.:
~ :::'1.
,.';;.

" :'..'.~

..
~

.
~

..
III

'"
"", e
# •
~
, •
, •
~
• ~ I

,
••
c

Fig. 39a-f Oxytricha faurei (from DRAGESCO & DRAGESco-KERNEis 1986. a-t: protargol impregnation). a)
Ventral infraciliature, 170 Ilm. This African population differs in some respects from the type population (Fig.
39g-j) so that the conspecifity is uncertain. b, c) Ventral infraciliature of anterior and posterior part of cell. d)
Ventral infraciliature of posterior portion of a specimen with 6 transverse cirri. e, t) Arrangement of fme cau-
dal "cirri" in dorsal view. Bar divisions = 10 Ilm; p 146.

M 0 r p hoI 0 g y and b i 0 log y: In life about 160-180 ~m long and 8.5 times as
long as broad, protargol-impregnated specimens of DRAGESCO & DRAGESCo-KERNEIS
(1986) only about 3.4 times as long as broad. Margins parallel, posterior end more or less
rounded, anterior distinctly intended at right margin (Fig. 39g). Cross-section round (Fig.
39i). Macronuclear nodules after protargol impregnation 13-25 ~m long (mean = 17 ~m;
n = 24). 2-3 small spherical micronuclei. Contractile vacuole slightly in front mid-body.
 148 SYSTEMATIC SECTION

Fig. 39g-j Oxytricha faurei (after TUCOLESCO 1962. g-j, from life). g)
Ventral view, 160-180 ~m. The cortical granules are drawn only in the
anterior portion (arrow). b) Posterior end of cell in ventral view. i) Cross-
section. j) Arrangement of cortical granules in lateral view (left) and as
seen from above (right); p 146.

..::.,..-...

,
"
~

"
Cortical granules rod-shaped, rather regularly arranged in
:::~;:
longitudinal rows. Granules probably colourless because
..:::......
, TUCOLESCO (1962a) did not mention any colour (Fig. 39g, h,
...
'" j); granulation not mentioned by DRAGESCO & DAAGESCO-
KERNElS (1986).
j
Adoral zone of membranelles only about 18 % of body
length in type population (Fig. 39g), about 33 % in African
population (Fig. 39a). Buccal area very small. Five trans-
verse cirri, protrude only slightly, if at all, beyond posterior
end of cell. Marginal rows widely separate in type popula-
tion (Fig. 39g, h); left marginal row distinctly overlapping
on right margin in African population (Fig. 39a, c). Dorsal
cilia shortish than long. 5-6 fine, distinctly elongated caudal
cirri (Fig. 39g, h).

o c cur r e n c e and e col 0 g y: Very rare species.

Locus classicus is a water body in the lalomicioara Cave,
Romania. TUCOLESCO (1962a) also found O. faurei in a
small brook outside the cave. DRAGESCO & DRAGESco-KERNEIS (1986) collected it in fresh-
water habitats near Cotonou, Benin. Feeds on diatoms (DRAGESCO & DRAGESCo-KBRNEIS
1986). Biomass of 106 individuals of type population about 50 mg.

Oxytricha granulosa SCHMITZ, 1986 (Fig. 40a-d)

1986 Oxytricha granulosa SCHMITZ, Dissertation, p 91 (type slides are deposited in the Institut file land-
wirtschaftliche Zoologie und Bienenkunde at the University of Bonn, Germany).

Nom e n c I a t u rea n d t a x 0 nom y: This species is described in a thesis, which

is generally not considered a valid publication (lCZN 1985, Article 9). Conversely, O.
granulosa is included in the Zoological Record. Investigation of other populations rec-
ommended.

M 0 r p hoI 0 g y and b i 0 log y: In life (?) 155-280 x 45-60 ~m (mean = 195 x

50 ~m). Long elliptical, anterior end broadly rounded, posterior portion tapering. Smooth
and flexible, slightly contractile. Two ellipsoidal, 18-25 ~m sized macronuclear nodules
 Oxytricha 149

re;7Jr--AHl , .

\b-.a--Ha

Ek eY

HargR

",,--Harg L
1\-\-N--- TrYe

1-U1~---(e c

frc
i=::::::rr--AHZ
fT-f.:="-Ir--- par H
'1=-~--t-endH

Harg R Harg L Fig.40a-d Oxytricha granulosa (from ScHMrrz 1986. a,

b, from life; c, d, protargol impregnation). a) Ventral
view, 160 ~m. b) Ventral view (?) showing rows of corti-
cal granules. C, d) Dorsal and ventral infraciliature,
160 11m. Explanation of original labels: AMZ = adoral
zone of membranelles, Cc = caudal cirri, cV = contrac-
tile vacuole, OK = dorsal kineties, Ek = inclusion, endM
"--"f-- Trve = endoral, Frc = right frontal cirrus, GR = cortical gran-
ules, Ma = macronuclear nodule, Marg L = left marginal
cirral row, Marg R = right marginal cirral row, Mi = mi-
cronucleus, parM = paroral, Trvc = transverse cirri;
d p 148.

left of median; only one small (2-3 /lm) globular micronucleus between macronuclear
nodules. Contractile vacuole, according to description, in front of mid-body, according to
Figure 40a, however, slightly behind mid-body, during diastole with short collecting ca-
nals. Pellicle colourless. Cortical granules yellow-green, spherical, about 0.7 /lm across,
arranged in 14-18 longitudinal rows per side (Fig. 40b). Cytoplasm colourless, with
many compact food vacuoles containing algae and yellowish, irregularly shaped inclu-
sions rendering cells brownish at low magnification. Rapid movement.
150 SYSTEMATIC SECTION

Adoral zone of membrane lIes 114-113 of body length, consists of 44-50 adoral mem-
branelles; distal end distinctly overlapping on right margin. Paroral and endoral in Oxytri-
cha pattern, composed of basal body pairs each (Fig. 40a, d). Three slightly enlarged
frontal cirri, 1 buccal cirrus, 4 frontoventral cirri, 3-4 postoral ventral cirri (2 adjoin buc-
cal vertex, 1 or 2 slightly to distinctly separated posteriorly), 1-2 pretransverse ventral
cirri, 5-6 about 18 Jlm long transverse cirri, which do not protrude beyond posterior end
of cell; number of frontal-ventral-transverse cirri obviously slightly variably (Fig. 40d).
31-48 (mean = 40) left and 33-48 (mean = 41) right marginal cirri, in life about 15 Jlm
long, each composed of2 x 6 basal bodies; marginal rows not distinctly separated poste-
riorly. Dorsal cilia (r8 Jlm long, arranged in 4-5 dorsal kineties of body length (Fig.
40c), rightmost kinety often terminating near mid-body (dorsomarginal row!). 2-4 incon-
spicuous, about 15 Jlm long caudal cirri, composed of2 x 2 basal bodies.

o c cur r e n c e and e colo g y: Rare. Locus classicus is the Rhine River (Oberer
Niederrhein) in Germany, where SCHMITZ (1986) discovered O. granulosa in the auf-
wuchs at less than 10°C during spring and autumn. Feeds on green algae and diatoms.

Oxytricha hymenostoma STOKES, 1887 (Fig. 381, 4Ia-g, 42a-i, Table 3)

1887 Oxytricha hymenostoma STOKES, Ann. Mag. nat Hist., 20: III (Fig. 41a).
1888 Oxytricha hymenostoma, STOKES - STOKES, 1. Trenton nat Hist. Soc., I: 290 (Fig. 41 a).
1891 Oxytricha ludibunda STOKES, JI R. microsc. Soc., year 1891: 702 (Fig. 4Ie).
1932 Oxytricha hymenostoma STOKES, 1887 - KAHL, Tierwelt Otl., 25: 603.
1932 Oxytricha ludibunda STOKES, 1891 -I<AHL, Tierwelt Otl., 25: 603.
1932 Oxytrichafallax- REYNows, 1. expo Zoo!., 62: 330 (Fig. 381; misidentification).
1935 Oxytricha - LUND, 1. Morph., 58: 258, 275 (Fig. 42g).
1950 Oxytricha elliptica GELEI & SZABADOS, Annis bioI. Univ. szeged., 1: 265, 285 (Fig. 41 f, g).
1966 Oxytricha minor ORAGESCO, Arch. Protistenk., 109: 197 (Fig. 42i).
1970 Oxytricha elliptica GELEI & SZABADOS, 1950 - ORAGESCO, Annis Fac. Sci. Univ. fed. Cameroun
(Numero hors-serie): 122 (Fig. 42h).
1970 Oxytrlcha lundi ORAGESCO, Annis Fac. Sci. Univ. fed. Cameroun (Numero hors-serie): 123 (for the
Oxytricha of LUND 1935).
1970 Oxytricha minima ORAGESCO, Annis Fac. Sci. Univ. fed. Cameroun (Numero hors-sene): 123 (see no-
menclature).
1972 Opistotricha macrostoma ORAGESCO, Annis Fac. Sci. fed. Cameroun, 9: 119 (Fig. 41b-d; incorrect sub-
sequent spelling of Opisthotricha).
1972 Oxytrichafallax STEIN - GRIMES, 1. Protozool., 19: 428 (Fig. 42d; misidentification).
1974 Oxytricha dragescoi STILLER, Annis hist.-nat. Mus. natn. hung., 66: 133 (see nomenclature).
1979 Oxytrichafallax STEIN (1859) - FOISSNER, Int. Rev. ges. Hydrobio!. Hydrogr., 64: 131 (Fig. 42a-c; mis-
identification).
1981 Oxytrichajallax- HAMMERSMITH & GRIMES, J. Embryo!. expo Morph., 63: 19 (Fig. 42e, f; misidentifica-
tion).
1986 Oxytricha hymenostomata STOKES, 1887 - DRAGESCO & ORAGESCo-KERNEIS, Faune tropicale, 26: 461
(Fig. 41 b; incorrect subsequent spelling).
1988 Oxytricha ludibunda STOKES, 1891- FOISSNER, Hydrobiologia, 166: 22.
 Oxytricha 151

..

Fig. 41a-c Oxytricha hymenostoma (a, after STOKES 1887b; b, after DRAGESCO 1972a from DRAGESCO &
DRAGESCO-KERNEIS 1986; c, from DRAGESCO 1972a a, from life; b, c, protargol impregnation). a) Ventral view,
100 )lm. The body shape, the undulating membranes, the length of the adoral zone of membranelies, the poste-
riorly displaced posterior postoral ventral cirrus are reminiscent of Sty/onychia spp. or Sterkiella histriomusco-
rum. However, according to STOKES the body is soft and flexible, strongly indicating that O. hymenstoma is an
Oxytrichinae. b, c) Ventral and dorsal infraciliature, b = 70)lot, c = 75 )lm; p 150.

1991 Oxytricha hymenostoma STOKES, 1887 - FOISSNER, BLATTERER, BERGER & KOllMANN, lnformationsber.
Bayer. Landesarntes filr Wasserwirtschaft, 1/91: 289.

Nom e n c I at u rea n d t a x 0 nom y: Oxytricha minor DRAGESCO, 1966a, is ajun-

ior secondary homonym of O. minor (MAsKELL, 1887b) KARL, 1932 and a primary homo-
nym of O. minor KARL, 1932 (now O. alfredO. This was already recognised by DRAGESCO
(1970), who introduced the replacement name 0. minima for his species. STIlLER (1974a)
was obviously unaware this because she proposed another substitute name, namely O.
dragescoi. Incorrect subsequent spellings: Oxytricha hymatostoma STOKES (SInN & KIM
1993b, P 224); Oxytricha hymenostomata (CAIRNS 1965a, p 12; FOISSNER 1988a, P 22;
GROLrERE 1975, P 491; HORvArn 1950, p 159; LEPSI 1965, P 975); Oxytricha lundibunda
(THOMPSON 1972, P 287).
I agree with HEMBERGER (1982) that the two species of STOKES should be syn-
onymised because the original descriptions do not show significant differences. BORROR
152 SYSTEMATIC SECTION

(
t
..

Fig.41d-g Oxytricha hymenostoma

(d, from DRAGESCO 1972a; e, after
STOKES 1891; t: g, from GELEI and
SZABADOS 1950. d, protargol impreg-
nation; e, from life; 1; g, opalblue
staining after BRESSLAU). d, r, g)
Ventral and dorsal infraciliature, d =
70 ~m, 1; g = 100 ~m. e) Ventral
view, 105 ~m; p 150.

(1972a) assumed identity with O. fallax, which is, however, distinctly larger and has an
oval outline. Oxytricha elliptica, O. lundi, and Opisthotricha macrostoma were already
synonyrnised with the present species by DRAGESCO & DRAGESCo-KERNEIS (1986). The
populations by REYNOLDS (1932), GRIMES (1972), FOISSNER (1979a), and IiAMMERSMITII &
GRIMES (1981) are too small to be reliably identified with 0. fallax. In addition, they do
not show the characteristic oval body shape. Conversely, O. hymenostoma sensu SHIN &
KIM (1993b; Fig. 39p-r) is classified as O. fallax because of its large size (120-190 x
50-85 /lm). HAsHIMOTO (1961, 1962, 1963, 1964) identified his "O.fallax" according to
the descriptions by BISHOP (1943; classified as unidentifiable taxon), HORVATH (1933; no
figure), and LUND (1935; classified as O. hymenostoma), indicating that he did not work
with a true 0. fallax. Thus, I preliminary classify his population as O. hymenostoma.
The preceding paragraph shows how complicated the taxonomy of O. hymenostoma
is and, unfortunately, I am not able to provide a satisfactory solution. The original de-
scription and several redescriptions listed above are reminiscent of Sterkiella histriomus-
corum (for example, Fig. 41f, 42a, d, e) or of a small Stylonychia (Fig. 41a, b, d, e,
42g-i) in many characters: shape, arrangement of postoral ventral cirri, pattern of undu-
lating membranes, length of adoral zone of membranelles. However, according to STOKES
 Oxytricha 153

,/-\ --
, <
l
~
{ { I
--, I
\

,, ,
\
--". \ ~
\ \
',-. '; ') (
~ \\ ( (
-\.... \
,,,
(
--, ')\
-. '\ "
( <
.,
,,
)
(
-. 'l

,,
f

,
,,
a b
,
Fig. 42a-c Oxytricha hymenostoma (from FOISSNER 1979a, a-c, composite after live observation and protar-
gol- and Chatton-Lwoff silver-impregnated specimens), a) Ventral infraciliature, 110 ~m, b) Lateral view,
125 ~m, c) Dorsal infraciliature, 120 ~m, The silverline system is shown only in the left posterior portion;
p ISO,

(1887b), the body is soft and flexible; synonymisation is hence not justified None of the
redescriptions considers both the live aspect and the infraciliature with a detailed mor-
phometric characterisation, thus none can be deemed authoritative. Detailed redescription
necessary.
The population by ESPOSITO et al. (1978) is only schematically illustrated, so that the
identification cannot be verified (Fig. 221j). The populations by CONN (1905; Fig.
22Id), CURDS (1969; Fig. 221g), and NAlDU (1965; Fig. 224m) are insufficiently rede-
scribed.

M 0 r p hoi 0 g y and b i 0 log y: Classification in Oxytricha uncertain, see taxon-

omy. In life 70-110 Jlm long, 2.5 times as long as broad. Elliptical to nearly parallel-
sided. Both ends rounded. Usually 2 micronuclei, however, amicronucleate races are re-
ported (DAWSON 1919, 1920). Contractile vacuole about in mid-body, pulsates about
every 14-15 s (GELEI 1935). Rapid movement.
154 SYSTEMATIC SECTION

·U ••
.Q
II ~ -: \

-.
(
, \: ,:
II ~
-,
• ~
, /I ~ \
-,
{

•
•
I - ': "" I
.• • •
~ '"t

•. •
....
t •
#
d

Fig. 42<1-( Oxytricha hymenostoma (d, from GRIMES 1972b; e, f, after HAMMERSMI1H & GRIMES 1981. d, pro-
largol impregnation; e, f, redrawn from scanning electron micrographs). d) Ventral infraciliature, about 80 J.lm.
Arrow marks posterior postoral ventral cirrus. e, f) Ventral and dorsal infraciliature, e = 55 J.lm, (= 60 J.lm.
Possibly Sterkiella histriomuscorum; p ISO.

Adoral zone of membrane lies about 40-45 % of body length (that is, Stylonychinae-
like!), 22-35 membranelles. Frontal and transverse cirri distinctly enlarged. Buccal cirrus
somewhat behind anterior end of undulating membranes. Third postoral ventral cirrus
distinctly displaced posteriad (Fig. 42d, arrow). According to STOKES (l887b, 1891),
transverse cirri protrude slightly beyond posterior end of cell; according to the redescrip-
tions all transverse cirri protrude. Caudal cirri inconspicuous, arranged at ends of kineties
1,2, and 4 (Fig. 41c, g, 42c, t).
Resting cysts in live about 18-20 ~m across. Wall smooth, about 2-3 ~m thick
(GRIMES 1973d). Ultrastructure and morphogenesis of O. hymenostoma are described by
HASHIMOTO (1961), GRIMES (1972), and GRIMES & ADLER (1976). The formation of the
oral primordium commences very close to the left transverse cirrus, and dorsal infracilia-
ture originates according to the Oxytricha pattern. Conjugation was studied by HAMMER.
SMITH (l976a).
Further literature, mainly concerning experiments on regeneration, doublet forma-
tion, biochemistry, and genetics: BANCHEITI et aI. (l978b, 1980a, b, 1982a), CETERA et aI.
 Oxytricha 155

Fig. 42g-i Oxytricha hymenostoma (g, after LUND 1935; h, from DRA-
GESCO 1970; i, from DRAGESCO 1966a. g, Haidenhain's hematoxylin
stain; h, i, protargol impregnation). Ventral infi'aciliature, g = 75 Jim, h
= 65 Jim, i = about 80 Jim; p ISO.
Fig. 42j Keronopsis /ongicirrata (from GELEI & SZABADOS 1950. Bress-
lau stain). Ventral infraciliature, 100 Jim (nuclear apparatus not
known). A supposed synonym of Oxytricha /ongicirrata, p 163.
156 SYSTEMATIC SECTION

(1978), ESPOSITO et al. (l976a, 1978), GRIMES (1973a-<:, e, 1974, 1976, 1982a), GRIMES &
GAVIN (1987), GRIMES & HAMMERSMITH (1980), lfAMMERSMITH (1976b, 1978), NOBll..I et al.
(1981), REYNOLDS (1932; see also FAURE-FREMIET 1967), RICCI et al. (1978, 1980a).

o c cur r e n c e and e colo g y: In the aufwuchs and detritus of freshwater habitats.

Locus classicus is a hay-infusion from the USA; STOKES (1887b) did not state the specific
locality where he discovered Oxytricha hymenostoma, however, he lived and worlced in
Trenton, New Jersey, USA, and I assume that the type locality is near there. Locus classi-
cus of the synonym 0. ludibunda is a pond with decaying Sphagnum in the USA (STOKES
1891). Locus classicus of the synonym 0. minor is a hay-infusion(?) from Thonon,
France (DRAGESCO 1966a) and the locus classicus of the synonym o. elliptica is a rain
pool in Szeged, Hungary (GELEI & SZABADOS 1950). Locus classicus of the synonym "Op-
istotricha macrostoma" is a plant infusion in Uganda (DRAGESCO 1972a). Further records
substantiated by illustrations: eutrophic alpine ponds in Austria (FOISSNER 1979a); fresh-
water canals near Pisa, Italy (ESPOSITO et al. 1978); river near Antelias, Lebanon (LUND
1935); Japan (HAsIDMOTO 1961); freshwater aquarium in the Osborn Zoological Labora-
tory in New Haven, USA (REYNOLDS 1932); Jordan River, a small stream located on the
campus of Indiana University of Bloomington, USA (GRIMES 1972b, HAMMERsMITH &
GRIMES 1981); Cameroon (DRAGESCO 1970).
Records from freshwater habitats not substantiated by illustrations: slime of polluted
waters in England (CURTIS & CURDS 1971); Italy (GRISPINI 1938); Turiec River in Slova-
kia (TIRJAKovA 1993); water reservoirs in the USSR (KRAVCHENKO 1969); China (NING et
al. 1993, SHEN & GONG 1989); Tibetan Plateau (WANG 1977); debris from a pond near
New Haven, Connecticut, USA (DAWSON 1919); infrequent in soft waters and frequent in
hard waters of south-eastern Louisiana, USA (BAMFORTH 1963); at about 24°C in the
Tahquamenon River, Douglas Lake Region, Michigan, USA (CAIRNS & YONGUE 1966);
various water bodies in Conestoga Basin, Pennsylvania, USA (CAIRNS 1965a); river with
medium-hard water in Eastern United States (PATRICK 1961); Peru (SARMIENTO & GUERRA
1960); mosses from Venezuela (SCORZA & NuNEz MONTIEL 1954); Coast Lake, Victoria
Land, Antarctica (THOMPSON 1972).
HORVATH (1950) found 0. hymenostoma in the sandy soil of the flood plain of the
Tisza River at Szeged, Hungary. The records from Indian soil (BHATTACHARYA et al.
1977), Mississippi deltaic soils, USA (BAMFORTH 1969), and soil from paddy fields in Ja-
pan (TAKAHASll & SUHAMA 1991) are not substantiated by illustrations.
According to literature common in biological sewage-treatment plants, such as acti-
vated sludge, percolating filters, and rotating biological contactors (BANINA 1983, CURDS
1975, KLIMOWICZ 1973, KUTIKOWA 1984, MADaNI & GHE1TI 1981b, MADaNI et al. 1979,
REID 1969). CURDS (1966b) observed o. hymenostoma mainly during maturation of acti-
vated sludge. KLIMOWICZ (1972) counted 2735 indo ml- I in a Polish aeration tank under
following conditions: mean sludge loading = 0.04 kg BODs kg- I dry mass d-I, mean con-
centration of activated sludge = 3.75 g I-I, time of sewage aeration = 24 h, destruction of
organic matter = 90 % BODs, mean oxygen content in aeration tank = 3.4 mg I-I, 5 to
 Oxytricha 157

16°C, mean pH of affluent sewage = 6.9; abundances were higher at low sludge loading
than at high loading (KLIMOWICZ 1970).
Feeds on bacteria, autotrophic flagellates (Chlamydomonas sp.), algae, diatoms, fila-
mentous growths, flagellates, ciliates (CURTIS & CURDS 1971, FOISSNER 1979a); also can-
nibalistic (DAWSON 1919). Average generation time over a four month period (presuma-
bly at room temperature) about 11 h, shortest generation time 5 h (DAWSON 1919). HASm-
MOTO (1961) estimated 6.8 hat 25-27 °C in a wheat infusion and with Chilomonas para-
maecium as food. GRIMES (1972b) maintained the organisms in phosphate-buffered baked
lettuce medium inoculated with Enterobacter aerogenes. Biomass of 106 medium-sized
individuals about 30 mg (ForSSNER et al. 1991).
PATRICK et al. (1967) found Oxytricha hymenostoma in the Savannah River, USA,
under following conditions: >4-20°C, 7-<11 mg 1-\ O2, 0.5-1.0 mg 1'\ BODs, 0.009 to
0.08 mg 1'\ NH3-N, <0.007 mg 1'\ NOi-N, >0.05-0.2 mg 1'\ NOl'-N, >0.01-0.05 mg 1'\
POl·-p. ForSSNER et al. (1982) provided following autecoiogical data for populations
from small, Ientic, dystrophic alpine water bodies from Austria (56 analyses): 0.2 x 106 to
20 X 106 bacteria mI'\ (direct counting), 0.2-23°C, pH 4.7-6.2, 3.0-12.9 mg 1'\ O2
(37-191 % saturation), 9-100 mg 1'\ KMn04-consumption, 0.0-6.2 mg}"\ NH/-N,
0-3 mg 1'\ NOi-N, 0-0.5 mg }.\ NOl'-N, 0-0.57 mg 1'\ POl'-P, 0-1.2 mg}"\ FeJ+, up to
0.8 mg }.\ H2S.
Oxytricha hymenostoma is classified as polysaprobic indicator of water quality: p; a
= 2, P = 8, 1 = 4, SI = 3.8 (SLADECEK et al. 1981, WEGL 1983, FOISSNER et al. 1991). This
distribution agrees with the fact that it occurs in sewage treatment plants mainly during
the maturation phase. However, I have some doubt that all other sample sites mentioned
above were alphamesosaprobic or polysaprobic. Thus, the saprobic classification has to
be confirmed on reliably determined populations.

Oxytricha kahlovata nom. nov. (Fig. 43a-c)

1932 Opislhotricha ovala KAHL, Tierwelt Dtl., 25: 609 (Fig. 43a).
1971 Opislolricha ovala KAHL - CHARoEZ, Revue verviet. Hist. nat., 28: 38 (Fig. 43c; incorrect subsequent
spelling).
1974 Oxytricha ovala (KAHL) - PATSCH, Arb. lnst. landw. Zool. Bienenk., 1: 61 (Fig. 43b).

Nom e n c I at u rea n d t a x 0 nom y: KARL (1932) classified Opisthotricha as

subgenus of Oxytricha; the correct name in the original description is thus Oxytricha
(Opisthotricha) ovata. However, this is a junior primary homonym of Oxytricha ovata
DUMAS, 1929. BORROR (1972a) synonymised O. kahlovata with Cyrtohymena sapropeli-
ca, which also has distinctly anteriorly displaced transverse cirri and a single micronu-
cleus (Fig. 96h). However, they differ in body-shape and shape of undulating membranes.
The arrangement of the frontal and ventral cirri is doubtful in Oxytricha ovata sensu
PATSCH (1974), so that the following description is based mainly on KARL's data. The re-
description by BIERNACKA (1967; Fig. 226e) is insufficient.
158 SYSTEMATIC SECTION

M 0 r p hoi 0 g y: In live about 100 to

120 ~m long. Anterior end narrow, poste-
rior very broadly rounded, that is, outline
oval. KAlIL (1932) did not observe a micro-
nucleus, but he supposed one, which is con-
firmed by the redescription by PATSCH
(1974; Fig. 43b). Contractile vacuole dis-
tinctly in front of mid-body. Cytoplasm of-
ten darkly granulated. Adoral zone of mem-
braneHes about 113 of body length. Frontal
cirri distinctly, transverse cirri probably not
enlarged. Dorsal cilia in life about 6 ~m
long. Three elongated caudal cirri.

Occurrence and ecology:KAlIL

(1932) discovered O. leahlovala among de-
caying blades of grass in a freshwater habi-
tat, probably somewhere in Gennany. Possi-
b ble polysaprobic. Further records: freshwa-
ter in Belgium (CHARDEZ 1971); scattered in
some brooks near Bonn, Gennany (PATSCH
1974); Latvian rivers (LIEPA 1978). AGA-
MALIEV (1975) recorded it from the western
Fig. 43a-c Oxytricha kahlovata (a, from KAHL
1932; b, from PATSCH 1974; c, from CHARDEZ coast of the Caspian Sea (see also AGAMALI-
1971. a, c, from life; b, protargol impregnation). a, YEV 1976b). ROSA (1962; Tafel 1lI, 16)
c) Ventral views, a = 110 Jim, c = 105 Jim. b) found 0. kahlovala in Czech forests. Feeds
Ventral infraciliature, 70 Jim; p 157. on bacteria and flagellates. Biomass of 106
individuals about 70 mg. DETCHEVA (1979c)
found it once in the Strouma River, Bulgaria, at 20 °C, pH 7.7, 10 mg 1-1 0 2(97 % satura-
tion), 2.4 mg I-I BODs, 51 mg I-I Ca2+, 16 mg I-I Mg2+, 98 mg I-I SOlo, 11 mg I-I CI-,
0.08 mg I-I NH/-N, 1.3 mg I-I NOl--N, 0.11 mg I-I N02--N, 0.11 mg 1-1 Fe2+.

Oxytricha longa GELEI & SZABADOS, 1950 (Fig. 44a-q, 45a, b,

Tables 3, 4, 15)

1933 Oxytricha pelionella O. F. MOLLER - HORVAlH, Arch. Protistenk., 80: 295 (Fig. 45a, b; misidentifica-
tion; incorrect subsequent spelling ofpellionella).
1950 Oxytricha longa GELEI & SZABADOS, Annis bioI. Univ. szeged., I: 266, 286 (Fig. 44a).
1984 Oxylricha tonga GELEI & SZABADOS - NIESSEN, Diplomarbeit, p 71 (Fig. 44o--q).
1987 Urosomoida agilijormis FOISSNER, 1982 - GANNER, FOISSNER & ADAM, Annis Sci. nat., 8: 199, in part:
populations P3 and P4; not populations PI and P2 (Fig. 44b-n; authoritative redescription).
 Oxytricha 159

4-- -j-\
3-- --I J (
2 - - -;. / (

I
I

,,
,I

, /
,,
,,
, )
,
,, )
I
, I

• d

Fig.44a-d Oxytricha longa (a, from GELEI & SZABADOS 1950; b-d, population 3 from GANNER et aI. 1987b. a,
sublimate fixation; b, from life; c, d, protargol impregnation). a, b) Ventral views, a = 140 Jim, b = 60 Jim. Ar-
rows in (b) mark 2 caudal cirri. d) Ventral and dorsal infraciliature, 60 Jim. Oxytricha longa should not be
confused with Urosomoida agiliformis, which has only 1-4 transverse cirri inclusively pretransverse ventral
cirri. 1-4 = dorsal kineties 1-4; p 158.

T a x 0 nom y: I am rather certain that the populations P3 and P4 by GANNER et al.

(I987a, b) are conspecific with the population described by GELEI & SZABADOS (1950),
especially since they have the same nwnber of dorsal kineties. I don't know why GANNER
et al. (I987b, Table II) asswned that Oxytricha longa has 5 kineties, although in the
original description (p 286) it says: "so dass wir in den meisten Praparaten nur 4 Borsten-
reihen (= dorsal kineties) antreffen". The most important character for separation from U.
agiliformis is the swn of the transverse cirri and pretransverse ventral cirri (see key and
Table 15). Furthennore, morphogenesis of Oxytricha tonga proceeds as in Oxytricha
granulifera, that is, primordia V and VI of the proter do not originate de novo as in Uro-
somoida spp., but from cirrus IV/3 and V/4 respectively (Table 4).
Oxytricha longa is also rather similar to O. similis, especially concerning the 2 cau-
dal cirri which point towards the right side. However, according to the authoritative rede-
scription by KAm.. (1932), O. similis has 5 transverse cirri (Fig. 57a-d). SONG & WILBERT
(1989, P 162) provided no figure but included a morphometric characterisation of Oxytri-
 160 SYSTEMATIC SECTION

Fig.44e-h Oxytricha longa (population 3 from GANNER et aI. 1987b. e-h, protargol impregnation). e) Ventral
infraciliature of anterior half of cell, 30 11m. f-h) Ventral infraciliature of early morphogenetic stages, f =
60 11m, g, h = 55 11m. Arrow in (f) denotes first proliferation of basal bodies for oral primordium of opisthe;
p 158.

cha longa. Their population usually has 3 (mean = 2.86; n = 7) caudal cirri, like the NIES-
SEN(1984, Fig. 44q) population, indicating that further species exist.

M 0 r p hoi 0 g y and b i 0 log y: In life 60-100 x 25-40 J.1m. Elliptical. Macronu-

clear nodules about 12 x 5 J.1m after protargol impregnation. Usually 2 (rarely 3), after
protargol impregnation, 1.5-3.0 x 1.4-2.3 J.1m sized micronuclei. Posterior portion of
narrow buccal area covered by a very hyaline pellicular projection (Fig. 44a, b). Buccal
cirrus at anterior end of undulating membranes. Membranes slightly curved, but not inter-
secting. Dorsal kinety 2 anteriorly and kinety 3 posteriorly slightly shortened. Kinety 4
terminates somewhat in front mid-body. Caudal cirri on kineties 1 and 2, distinctly longer
than marginal cirri (Fig. 44a, b, d, m, see taxonomy).
Morphogenesis commences with proliferation of basal bodies close to the postoral
ventral cirrus IV/2 (Fig. 44f; GANNER 1991, GANNER et al. 1987b). Immediately
afterwards, a second field of basal bodies develops apokinetaly in front of the left trans-

Fig. 44i-n Oxytricha Zonga (population 3 from GANNER et aI. 1987b. i-n, morphogenetic stages after protargol ~
impregnation). i-k) Ventral infraciliature of middle stages, i = 65 Ilm,j, k = 60 11m. Arrows in (i) mark disper-
sal of buccal and anterior postoral ventral cirrus. Arrows in G) denote development of marginal primordia. Ar-
rows in (k) mark origin of dorsal kinety 4 (dorsomarginal row) in proter and opisthe. I-n) Ventral and dorsal
infraciIiature of late stages, I, m = 60 11m, n = 70 11m. Dotted lines in (I) connect cirri which originate from
same streak. Arrow in (m) marks parental dorsal kinety 4. New cirri black, parental white; p 158.
 Oxytricha 161

""
"
"
"
"
"" 0 :-
"
"

0 ~
" "oo
"
0

"0
0
0
"
"
0 " Q
<> () 0 () () 0

~D () (][J0
<>
¢

#.~
o ~ ..

',' ~II~f~'"
# I
~ ~
~ ff
r~ ~
~:
....

·•.· c '#~I\I
'"
<> .\ ~:
,
'"

I ~.
~
'\111 :
•, ~f.
·•· ~
.
.: .
.
\\-
···
\
~ ~\~ :
••\'
0

i
· ·.~""
\

"~~~ ..
:,;>
" ..
... -
O()
"0

'(
,;>
.*.t:.O
0 "
" ...... 0

t /~ ~,
;:. ••....
',JI ~§Sl
~"
....•...:',• I. s§;:: -....
1'1'
I\ ~" ' ... ,
.. S
§ ..
• •
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• \\1 \~ : •
\ #~ ~ • ..
.. I ~ s. :
''. .......:
.
 162 SYSTEMATIC SECTION

.•
,::
0 ' .

'"

"
" (J

Fig.44o-q Oxytricha longa (from NIESSEN 1984.0, from life; p, q, protargol impregnation). 0) Ventral view,
80 11m. p, q) Ventral and dorsal infraciliature, 80 11m. This population from Australian saline soils deviates in
some characters (invariably 3 caudal cirri, arrangement of frontoventral and postoral ventral cirri,S transverse
cirri) from the populations ofGELEI & SZABADOS (1950) and GANNER et aI. (1987b), indicating that it is possi-
bly another species; p 158.

verse cirrus (Fig. 44g). The formation of the frontal-ventral-transverse streaks of the pro-
ter proceeds simultaneously with those of the opisthe (Fig. 44h-j). The streaks V and VI
of the opisthe extend towards the frontal field and generate the streaks V and VI of the
proter (Fig. 44i). The streaks I-VI generate the following numbers of cirri both in the
proter and in the opisthe: 1,2, 3, 3, 4, and 4. This holds for the large number of speci-
mens with 4 transverse cirri. About 8 % of the specimens investigated by GANNER et al.
(l987b) had 5 transverse cirri. In specimens with only 3 transverse cirri (8 % of speci-
mens) cirrus 11111 is lacking. Morphogenesis of dorsal ciliature is in Urosomoida pattern,
that is, fragmentation of dorsal kinety 3 does not occur.

o c cur r e n c e and e colo g y: In the aufwuchs and detritus of freshwater habitats

and in soil. Locus classicus is the Ski Istvan Square in Szeged, Hungary, where GELEI &
 Oxytricha 163

SZABADOS (1950) discovered 0. longa in a slightly polluted,

well-oxygenated rain puddle. High abundances occurred 10
days after sampling, indicating that it is resistant to
pollution. This is supported by the sample site of population
4 by GANNER et al. (1987b), namely the bank of the Salzach
River in Salzburg, Austria, which was polysaprobic by
wastes of pulp and paper industry, but well-oxygenated;
population P3 was isolated from a meadow soil in Israel.
HORVATH (1933) found Oxytricha longa in a pond in Sze-
ged, Hungary.
Further records: brooks and rivers in Upper Austria
(BLATIERER 1994, own observations); rarely in a eutrophic
pond (Poppelsdorfer Weiher) in Bonn, Germany (SONG &
WILBERT 1989); salt polluted running waters in Germany b
(MrnAILoWITSCH 1989); La Jarosa Reservoir, Spain
(FERNANDEZ-LEBORANS et al. 1990); alkaline saline soils from
arid regions in South Australia (NIESSEN 1984, POMP & WIL-
BERT 1988).
Feeds on bacteria, small flagellates, and ciliates. Can be
cultured in table water (Eau de Volvic, France) with some
Fig. 45a, b Oxytricha longa
squashed wheat grains to support bacterial growth (GANNER
(after HORvArn 1933. a, tolu-
et al. 1987b). Grew well also in plant infusion and in broth idinblue staining; b, from
(HORVATH 1933). Biomass of 106 individuals about 30 mg. life?). Ventral and right lateral
view, 110 J.lm; p 158.

Oxytricha longicirrata KAHL, 1932 (Fig. 46a, by

1932 Urosoma longicirrata KAHL, Tierwelt Dtl., 25: 607.

Nom e n c I at u rea n d t a x 0 nom y: I<AHL (1932) classified Urosoma as subge-

nus of Oxytricha; the correct name in the original description is thus Oxytricha (Uro-
soma) longicirrata. The overall morphology and the nuclear apparatus is reminiscent of
that of Ancystropodium maupasi (Fig. 198a-g). However, this species has a vel)' long re-
tractable tail-like process which was not observed in Oxytricha longicirrata.
Furthermore, Ancystropodium has a ventral infraciliature which agrees with that of Gas-
trostyla. The large buccal area and the position of cirrus 1II12 (distinctly behind level of
cirrus VII4) do not justify the classification in Urosoma. Oxytricha longicirrata differs
from Urosoma emarginata (Fig. 127a-y), which has also a notch on the right posterior
margin, not only in the arrangement of the frontoventral cirri, but also in that it has only
one micronucleus, conspicuously long cirri, and a short tail-like process. Pseudostrom-
1 Keronopsis /ongicirrata GEl.EI & SZABADOS, 1950, Annis bioI. Univ. szeged., I: 271,289 (Fig. 42j), a some-
what superficially described species from a rain puddle in Szeged, Hungary, is possibly ajunior synonym, as
indicated by body shape and long cirri (names!). 32 membranelles, 5 transverse cirri. Feeds on flagellates.
 164 SYSTEMATIC SECTION

bidium planctonticum is pelagic and has two micronuclei

and numerous caudal cirri (Fig. 147a-j). Detailed rede-
scription necessary.

M 0 r p hoi 0 g y and b i 0 log y: In life about

Fig. 46a, b Oxytricha [ongicir-
rata (from KAHL 1932). Left lat-
eral and ventral view from life,
180 J.lm. Note single micronucleus
(large black dot in b); p 163.
inconspicuous caudal cirri. Dorsal cilia in life about 5 Jim long.
150-200 Jim long. Broadest (about 1/3 of body length)
near anterior end. Only slightly or not flattened. Posterior
end with a short, conspicuous tail-like process pointing in
a right-dorsal direction (but not as long and not as retract-
able as in Ancystropodium maupasl). Macronuclear nod-
ules almost adjoining, spherical to slightly ellipsoidal.
Contractile vacuole about at level of cytostome. Slow ro-
tating movement. Adoral zone of membranelles about
38 % of body length. Frontal lip collar-shaped. Frontal,
ventral, and transverse cirri distinctly enlarged. All cirri
conspicuously long, marginal cirri posteriorly longer than
anteriorly. Likely typical Oxytricha cirral pattern. Trans-
verse cirri distinctly displaced anteriorly, that is, do not
(?) protrude beyond posterior end of cell. Probably three

o c cur r e n c e and e colo g y: Very rare species. Locus classicus is a pond near
Hamburg, Germany, where KAHL (1932) discovered O. longicirrata in the flocculent
sapropelic detritus. The mud was 10-15 cm deep and consisted largely of decaying
plants, such as Lemna and StratiDles (KAHL 1927a). KAHL (1932) always found it there
with low abundance and mainly during the winter. The records from a Polish pond filled
with sugar factory wastes and from Latvian rivers are not substantiated by illustrations
(GRABACKA 1973, LIEPA 1978). MATIS & TIRJAKovA (1994a) found it in running waters in
Slovakia. Feeds on rhodobacteria. Biomass of 106 individuals about 175 mg.

Oxytricha /ongissima DRAGESCO & NJINE, 1971 (Fig. 47)

1971 Oxytricha [ongissima DRAGESCO & NJlNE, AnnIs Fac. Sci. Univ. fed. Cameroun, 7-8: 134 (Fig. 47).
1986 Oxytricha [ongissima DRAGESCO & NJlNE, 1971 - DRAGESCO & DRAGESCo-KERNEIS, Faune tropicale, 26:
469.

T a x 0 nom y: Perhaps identical with the poorly known Oxytricha elongala (Fig. 36). In
life it can be confused with Urosoma giganlea (Fig. 134c-g), but after protargol impreg-
nation can be unequivocally separated by the number of dorsal kineties and the position
of the transverse cirri. Some morphometric data in the original description of Oxytricha
longissima are obviously wrong. Most of them are corrected in DRAGESCO & DRAGESCo-
KERNElS (1986). Detailed reinvestigation necessary.
 Oxytricha 165

Fig. 47 Oxytricha [ongissima (after DRAGESCO & NJINE 1971

from DRAGESCO & DRAGESCo-KERNEIS 1986). Ventral infracilia-
ture of a specimen with 4 postoral ventral cirri, protargoi im-
pregnation, 220 f.lm; p 164.

M 0 r p hoI 0 g y and b i 0 log y: In life about

250 !lm long, after protargol impregnation 210-235
x 96-120 !lm. Macronuclear nodules after protar-
gol impregnation 18-34!lm long (mean = 26 !lm);
each nodule with 1 spherical micronucleus, after
protargol impregnation 5.5-7.5 !lm across.
Adoral zone of membranelles about 113 of
body length. According to the DRAGESCO &
DRAGESCO-KERNEIS (1986) text, Oxytricha longis-
sima has only 35-38 (mean = 37) adoral mem-
branelles, according to Figure 47, however, about
46 membranelles are present; the higher value is
more likely for such a large hypotrich. Undulating
membranes almost in Stylonychia pattern, indicat-
ing that classification in Oxytricha is incorrect.
Frontal, ventral, and transverse cirri distinctly to
slightly larger than marginal cirri. 3-4 postoral ven-
tral cirri. Five distinctly anteriorly displaced trans-
verse cirri, do not reach posterior end of cell.
40-54 (mean = 46) right and 37-42 (mean = 40)
left marginal cirri. Dorsal cilia 2-4 !lm long, ar-
ranged in 5 kineties. Caudal cirri inconspicuous.

o c cur r e n c e and e colo g y: Locus classicus is a stagnant water in a track near

Dovala, Cameroon (DRAGESCO & NJINE 1971). NJINE (l977a) found Oxytricha longissima
with a frequency of7.1 % and at 23-27 °C. Not found since. Biomass of 106 individuals
about 570 mg.

Oxytricha matritensis RAMIREz-MON1ESINOS & PERFZ-SILVA, 1966 (Fig. 48)

1965 Oxytricha matritensis ALONSO, lnt. Congr. Protozool., 2: 230 (nomen nudum).
1966 Oxytricha matritensis RAMlREZ-MONTESINOS & PEREZ-SILVA, Microbioiogia esp., 19: 194 (Fig. 48).

Nom e n c I at u rea n d t a x 0 nom y: The name published by ALoNSO (1965) is

not accompanied by a description sufficient for differentiation from other taxa, so that it
must be considered as nomen nudum. RAMiREz-MONTESINOS & PEREZ-SILVA (1966) did not
 166 SYSTEMATIC SECTION

observe caudal cirri, indicating that this species possibly be-

longs to Tachysoma. However, since it cannot be excluded that
they overlooked them due to the inadequate stain method, I pre-
liminarily avoid changing the generic classification. A syn-
onymisation with Oxytricha aeruginosa as suggested by BORROR
(l972a) is not justified because no coloured cytoplasm is de-
scribed for O. matritensis. BORROR (1972a) also suggested iden-
tity with O. muitiseta, which has, however, more than 5 trans-
verse cirri. Poorly known, doubtful species which needs detailed
reinvestigation.

M 0 r p hoi 0 g y and b i 0 log y: In life about 100 to

Fig. 48 Oxytricha matri-
tensis (from RAMIREz-
MONTESINOS & PEREZ-SIL-
VA 1966. Stained with
crystal violet and iodine-
glycerine after PEREZ-SIL-
VA & RAMIREz-MoNTE-
SINOS 1964). Ventral in-
fraciliature, length about
110 /lm; p 165.
120 Ilm long, 3 times as long as broad. Elliptical. Macronuclear
nodules about 19 x 7 Ilm. Usually 2 micronuclei, about 3 Ilm
across. Adoral zone of membranelles about 113 of body length.
Ventral infraciliature presumably without specialities. Dorsal
cilia 2-4 Ilm long. Caudal cirri possibly absent (see taxonomy).
Cyst spherical, about 24-26 Ilm in diameter. AwNSO (1965) and
AwNSO & PEREz-SILVA (1966a) studied the polytene chromo-
somes.
o c cur r e n c e: Locus classicus is Madrid, Spain, where
RAMrnREZ-MoNTESINOS & PEREZ-SILVA (1966) discovered Oxytri-
cha matritensis in a moss habitat at a spring in the playground
of the "Centro de Investigaciones BioI6gicas", presumably a
freshwater habitat. Not found since. Biomass of 106 individuals
about 37 mg.

Oxytricha minor (MASKELL, 1887) KAHL, 1932 (Fig. 49a~)

1887 Opisthotricha parallela, ENGELM, var. minor MAsKELL, Trans. Proc. N. Z. Inst, 20: 17 (Fig. 49a).
1932 Opisthotricha parallela var. minor MAsKELL, 1888 -KAm.., Tierwelt Dtl., 25: 609 (Fig. 49b).
1975 Opisthotricha parallela (ENGElMANN, 1862) - GROUERE, Protistologica, 11: 493 (Fig. 49c; authoritative
redescription).

Nom e n c I a t u rea n d t a x 0 nom y: According to Article 45 (g) of the leZN

(1985) the rank of MAsKELL's (1887b) taxon "var. minor" is subspecific since it was pub-
lished before 1961. Thus, the change in rank from subspecies to species level does not af-
fect the authorship (Article 50 (c». The generic change was already undertaken by KAm.
(1932) when he classified Opisthotricha as subgenus of Oxytricha; the correct name in
his revision is thus Oxytricha (Opisthotricha) para/ella var. minor. However, Oxytricha
minor (MASKELL, 1887) KAm., 1932 is the senior secondary homonym of Oxytricha minor
 Oxytricha 167

Fig. 49a--c Oxytricha minor (a, after MASKELL 1887b; b,

from KAHL 1932; c, from GROLIERE 1975. a, b, from life; c,
protargol impregnation). a) Ventral surface in dorsal view,
70 Ilm. b) Ventral view, 100 Ilm. c) Ventral infraciliature,
60 Ilm; p 166.

KARL, 1932; the KARL (1932) species thus re-

quires a substitute name (see Oxytricha alfredl).
The original description is rather super-
ficial, so that each identification must be some-
what arbitrary. The population described by
GROLrERE (1975) is much more similar to the
KARL (1932) population than to O. parallela,
which is about 200 ~m long and has 4 distinct
caudal cirri. The conspecifity of KARL's and
GROLffiRE'S population is mainly indicated by the
elongated body which is slightly but distinctly
bent to the left in mid-body (compare with Fig.
49b, c). Redescription needed, especially live
observation and morphometric characterisation.

M 0 r p hoI 0 g y and b i 0 log y: In life

about 100 ~m long, after protargol impregnation
50-70 x 10-12 ~m (mean = 58 x 10.5 ~m). b
c
Elongated with parallel margins, both ends
rounded. Very flexible. Macronuclear nodules
after protargol impregnation 8-10 ~m long,
each nodule with 1 micronucleus. Contractile vacuole in front of mid-body. Cytoplasm
colourless, very transparent.
Adoral zone of membrane lIes about 1/3 of body length, 20-25 membranelles. Undu-
lating membranes rather straight. Arrangement of frontal-ventral-transverse cirri as in
many other Oxytricha species; according to GROLrERE (1975) only 3 frontoventral cirri
and sometimes only 1 pretransverse ventral cirrus. Five transverse cirri, protrude dis-
tinctly beyond posterior end of cell. 25-30 right and 18-21 left marginal cirri. Dorsal
cilia short, arranged in 3 kineties (uncommon, possibly a short row was overlooked?).
Three caudal cirri, distinctly longer than marginal cirri (Fig. 49b, c).

o c cur r e n c e and e colo g y: Very likely confmed to freshwater. Locus classi-

cus is a freshwater habitat in the Wellington District, New Zealand (MAsKElL 1887b).
KARL (1932) provided no information about the sample site. GROLrERE (1975) found Oxy-
tricha minor in temporary pools near Besse-en-Chandesse, France. Feeds on bacteria.
Biomass of 106 individuals about 4 mg.
168 SYSTEMATIC SECTION

Oxytricha monspessulana (CHATION & SEGUELA, 1940) BORROR, 1972 (Fig.

50a, b)

1939 Opislhotricha monspessu/ana CHArrON & SEGUELA, C. r. hebd. Stanco Acad. Sci., Paris, 208: 869 (no-
men nudum).
1940 Opisthotricha monspessu/ana CHArrON & SEGuELA, Bull. bioI. Fr. Belg., 74: 365,417 (Fig. 50a, b).
1972 Oxytricha monspessulana CHArroN & SEGuELA, 1940 - BoRROR, J. Protozool., 19: 14.

Nom e n c I a t u rea n d t a x 0 nom y: The name published by CHATTON & SEGUELA

(1939) is not accompanied by a description sufficient for differentiation from other taxa,
so that it must be considered a nomen nudum. Although BORROR (1972a) did not use the
phrase ''nov. comb.", as he did in other cases, I consider him as combining author. Some
characters, for instance, the long adoral zone of membraneHes and the arrangement of the
postoral ventral cirri, are reminiscent of Sterkiella histriomuscorum, with which it is pos-
sibly synonymous. The population described by GROLIERE (1969) is obviously not con-
specific with Oxytricha monspessulana because it has 4 macronuclear nodules and an in-
flexible body. GROLIERE'S population and that studied by FERNANDEZ-LEBORANS (1985) are
classified as Sterkiella cavicola (Fig. 181j-I).

M 0 r p hoI 0 g y and b i 0 log y: In life 90--120 x 55-65 11m. Flexible, which is the
only reason why I do not synonymise it with Sterkiella histriomuscorum. Macronuclear
nodules ellipsoidal, each with I spherical micronucleus. Contractile vacuole slightly in
front of mid-body.
Adoral zone of mem-
braneHes almost half
of body-length (indi-

- cating that this species

is a Stylonychinae, see
taxonomy), about 40
membranelles. Frontal,

I•
til
til
til
til Fig. 50a, b Oxylricha mon-
til spessulana (after CHArroN
,.,-
til & SEGuELA 1940. a, from
life; b, wet silver impregna-

,.
,- tion). a) Ventral view,

,.
90-120 )lm. Possibly a
til Sterkiella. b) Ventral infra-
,,
~ ciliature of an very early
morphogenetic stage, size
I not indicated. Only a small
part of the silverline system
is shown (arrow). OP = oral
b primordium.
 Oxytricha 169

ventral, and transverse cirri distinctly larger than marginal cirri. Five transverse cirri, pro-
trude slightly (left) to distinctly (right) beyond posterior end of cell. Dorsal cilia 1-2 !lm
long, arranged in 6 kineties. Three rather long and flexible caudal cirri.

o c cur r e n c e and e colo g y: Locus classicus is Montpellier, France, where

CHATION & SEGuELA (1940) discovered 0. monspessulana in a basin with algae (Spiro-
gyra sp., Characeae) and decaying leaves in the "Jardin des Plantes", a botanical garden.
The record by GROLIERE & NJINE (1973) is obviously identical to that OfGROLIERE (1969),
classified as Sterkiella cavicola. Oxytricha monspessulana feeds on bacteria. Biomass of
106 individuals about 90 mg.

Oxytricha multiseta
DRAGESCO, 1966 (Fig. 51a, b)

1961 Opisthotrieha similis ENGELMANN,

1862 - REuTER, Acta zool. fenn.,
99: 19 (Fig. 51 b; misidentifica-
tion?).
1966 Oxytrieha multiseta DRAGESCO,
Protistologica, 2: 87 (Fig. 51a).

T a x 0 nom y: The population

by RElTfER (1961) has 3 caudal
cirri, so that the identification as
O. similis (2 caudal cirri) is not
justified. The body-shape and the
ventral infraciliature of his popu-
lation are reminiscent of that of
Oxytricha multiseta. However,
REUTER (1961) described 2 pre-
transverse ventral and 5 transverse
cirri, whereas O. multiseta obvi-
ously has 8 to 9 pretransverse
Fig. 51a, b Oxytri-
ventral and transverse cirri. Thus,
eha multiseta (a,
an identification as O. multiseta from DRAGESCO
should be primarily based on the 1966b; b, after REu-
original description (Fig. 51a). TER 1961. a, protar-

BORROR (1972a) synonymised O. gol impregnation; b,

from life). a) Ven-
multiseta with 0. aeruginosa -
tral infraciliature,
which has, however, a reddish, llO J.lm. b) Ventral
granulated cytoplasm - and the view, size not indi-
poorly known 0. matritensis (Fig. cated.
170 SYSTEMATIC SECTION

48). Oxytricha multiseta and O. matritensis differ only in the number of transverse cirri,
which, however, has to be confmned by detailed redescriptions. The increased number of
transverse cirri in 0. multiseta is reminiscent of 0. variabilis (Fig. 62), and the overall
morphology agrees with that of 0. longa (4 transverse cirri) and Urosomoida agiliformis.
However, Urosomoida species usually have a lower number of pretransverse ventral and
transverse cirri.

M 0 r p hoi 0 g y and b i 0 log y: According to DRAGESCO (l966b) in life (?) about

120 ~m long, population by REUTER (1961) in life 80-90 ~m long and 3.6 times as long
as broad. Long elliptical, distinctly converging posteriorly, both ends rounded. Adoral
zone of membranelies about 30 % of body length, 26-29 membranelles. All postoral ven-
tral cirri arranged very close to cytostome. Distance between posterior postoral ventral
cirrus and pretransverse ventral cirri rather large. According to original description 6-7
transverse cirri; Figure Sla shows a specimen with 6 transverse cirri and 2 pretransverse
ventral cirri. About 33 right and 29 left marginal cirri. Dorsal cilia short, caudal cirri in-
conspicuous; distinguishable from marginal cirri in live specimens according to REUTER
(1961; Fig. SIb).

o c cur r e n c e and e colo g y: Locus classicus is a small pool in Haute-Savoie,

France (DRAGESCO I966b). Not found since. REUTER (1961) found his population in a
small rock pool (0.06 % NaCl") on Ungskar Island, Finland. Biomass of 106 individuals
about 30 mg.

Oxytricha paraha/ophila (WANG & NIE, 1935) comb. nov. (Fig. 84c)

1935 Opisthotricha paraholophila WANO & NIE, Sinensia, 6: 503 (incorrect original spelling).
1935 Opisthotrichaparahalophila WANO& NIE, Sinensia, 6: 504, 505.

Nom e n c I a t u rea n d t a x 0 nom y: In accordance with Article 32b (i) of the

leZN (l98S) I chose the species name "parahalophila" as the correct original spelling.
"Opithotricha parahalophild' in WANG & NIE (1935, P 440) is an incorrect subsequent
spelling of Opisthotricha.
Almost all characters, excluding the freshwater habitat, of Oxytricha parahalophila
agree very well with that of 0. halophila KAHL, 1932; especially 0. halophila from saline
inland waters (Fig. 84h) is rather similar. But because of the different biotops (freshwater
against marine) I preliminarily consider both 0. parahalophila and O. halophila as indi-
vidual species; however, their validity has to be checked by detailed redescriptions.

M 0 r ph 0 log y and b i 0 log y: In life 70-1 IS x 30 ~m. Both ends rounded, wid-
est at or slightly in front of mid-body. Macronuclear nodules ellipsoidal, not widely sepa-
rated. Single spherical micronucleus between macronuclear nodules, not, however, shown
in Figure 84c! Contractile vacuole slightly in front of mid-body. Cortical granules absent.
 Oxytricha 171

Cytoplasm colourless, granulated, usually transparent, slightly opaque in well-fed speci-

mens. Adoral zone ofmembranelles about 113 of body length. Frontal, ventraL and trans-
verse cirri slightly enlarged. Transverse cirri long and slender. Dorsal cilia "long and
fine", according to the illustration, however, only about 4 ~m. Three long and straight,
but rather thin caudal cirri, usually directed slightly to the right.

o c cur r e n c e and e colo g y: Locus classicus is a pond in Siao-Ying, China,

where WANG & NIE (1935) discovered Oxytricha parahalophila in an old standing cul-
ture among submerged mosses. Not found since. Feeds on bacteria and diatoms. Biomass
of 106 individuals about 20 mg.

Oxytricha paralleia ENGELMANN, 1862 (Fig. 52a-c)

1862 Oxytricha paral/ela ENGELMANN, Z. wiss. Zoo!., 11: 388 (Fig. 52a, b).
1882 Opisthotricha parallela, ENG. sp. - KENT, Manual infusoria ll, p 785.
1931 Opisthotricha parallela KENT - TAl, Sci. Rep. nato. Tsing Hua Univ., 1: 51 (Fig. 52c).
1932 Opisthotricha (Oxytricha) parallela ENGELMANN, 1862 - KAIn., Tierwelt Otl., 25: 609.

Nom e n c I a t u rea n d t a x 0 nom y: This species was unequivocally established

in Oxytricha, and not in Opisthotricha as erroneously assumed by BORROR (1972a). KAlIL
(1932) classified Opisthotricha as subgenus of Oxytricha; the correct name in his revi-
sion is thus Oxytricha (Opisthotricha) parallela. Incorrect subsequent spellings: Opistho-
tricha parallelis EN-
GELMANN (for example,
WEBB 1956, P 153),
Opistotricha parallela
ENGELMANN (MOCKE
1979, P 274). The

Fig. 52a--c Oxytricha par-

allela (a, b, after ENGEL-
MANN 1862; c, from TAl
1931. a-c, from life). a)
Dorsal view, 125/lm. Note
the 4 elongated caudal cirri
(arrows) and the long dorsal
cilia. b, c) Ventral views, b
= 160 /lm, c = 74/lm.

Fig. 52d Opisthotricha pa-

ralellis (from KAHL 1928).
Insufficient redescription,
100 /lm. Note two buccal
cirri; p 172.
172 SYSTEMATIC SECTION

specimen measured by TAl (1931; Fig. 52c) was only 74 J.lm long. However, since the
other characters (4 caudal cirri, long dorsal cilia) agree rather well with the original de-
scription, I accept the identification. The population described by KAHL (1928; Fig. 52d)
is distinctly smaller and clearly has 2 buccal cirri; it is preliminarily classified as insuffi-
cient redescription, as in Opisthotricha parallela sensu SCHUSTER (1899; Fig. 240i). The
small population of GROLIERE (1975) is fixed as the authoritative redescription of O. mi-
nor (MAsKELL, 1887) KAHL, 1932 (Fig. 49c). Detailed redescription of O. parallela
needed.

M 0 r ph 0 log y and b i 0 log y: According to the text by ENGELMANN (1862), in

life about 200 J.lm long, according to the illustrations, only 120-160 J.lm, TAl (1931)
population, even only 74 J.lm (see taxonomy); about 4 times as long as broad. Body mar-
gins nearly parallel, both ends rounded. Contractile vacuole about in mid-body; TAl
(1931) described 2 vacuoles (misobservation?). Rapid movement.
Adoral zone of membranelles only about 114 of body length. Frontal and transverse
cirri distinctly enlarged. According to Figure 52b, only 3 frontoventral cirri present. Five
transverse cirri, protrude distinctly beyond posterior end of cell. Dorsal cilia "long",
probably about 10 J.lm. Four prominent, widely spread caudal cirri.

o c cur r e n c e and e colo g y: Locus classicus is a freshwater habitat near leip-

zig, Germany (ENGELMANN 1862). TAl (1931) found Oxytricha parallela in a pond on
west campus of Tsing Hua University, China.
Records not substantiated by illustrations: Lansersee, a small lake in Tyrol, Austria
(DALLA TORRE 1891); rare in Maritza River, Bulgaria (DETCHEVA 1981); Moldavia River
in Prague, Czechoslovakia (KALMUS 1928, SRAMEK-HUSEK 1953); freshwater habitats near
Perth, England (CRAIGIE 1921); ponds and infusion of willow twigs, Estonia (JACOBSON
1928); mouth of Schussen River, Germany (WETZEL 1928a); brook near Bonn, Germany
(JUTRCZENKI 1982); small, eutrophic lake in Germany (MOCKE 1979); polluted Tisza River
in Hungary (J6SA 1974, SmLER 1942); stagnant water in Italy (CUNEO 1891); Latvian riv-
ers (LIEPA 1978, 1990); stagnant water with Lemna minor in Warsaw, Poland (WRZE-
SNIOWSKIEGO 1866); cooling plant in Moldova (CHORIK & VIKOL 1973); inland salt waters
of the USSR (BOUTCHINSKY 1895); soil in Japan (TAKAHASHI & SUHAMA 1991); St. John's
Bayou, a fairly large body of brackish water in Louisiana, USA (BORROR 1962, PATTER-
SON et al. 1989, SMITH 1904).
DETCHEVA (1983c) found 0. parallela in the Maritza River, Bulgaria, under follow-
ing conditions: alphamesosaprobity, 0.3 % frequency, pH 7.4,23 °C, 6.1 mg 1.1 0 2 (72 %
saturation), 89 mg I-I Ca2+, 8_8 mg I-I MgZ+, 235 mg I-I HC03-, 59 mg I-I SOlo, 20 mg I-I
CI-, 19 °dH, 0.8 mg I-I NH/-N, 0.085 mg 1-1 N0 2--N, 0.05 mg I-I Fe2+. WEBB (1956) re-
corded it throughout the year from brackish water at Parkgate, Cheshire, near the Dee
River estuary, Wales, at a salinity of up to 1.7 %. BARNES et al. (1976) found it in the
sediment of a sheltered and organically rich site on Scolt Head Island, Norfolk, England.
 Oxytricha 173

Feeds on bacteria, diatoms, and detritus (WEBB 1956). Biomass of 106 individuals about
67 mg (when 160 flm long).

Oxytricha pseudofurcata nov. spec. (Fig. 53a, b)

1954 Tachysomafurcata KAHL - GELEI, Acta bioI. hung., 5: 335 (misidentification).

Tax 0nom y: The determination OfGEIEI (1954b) is certainly wrong because Tachy-
soma bicirratum (the replacement name for Tachysoma furcata) has long dorsal cilia,
whereas they are short in Oxytricha pseudofurcata. Furthermore, the latter species has 2
caudal cirri and not 2 elongated marginal cirri as in Tachysoma bicirratum. The rather
similar Oxytricha setigera has long dorsal cilia. Detailed redescription necessary.

D i a g nos i s: In life about 66 x 25 flm; 1 mi-

cronucleus between the 2 macronuclear nodules;
about 18 adoral membranelles; 4 transverse
cirri; usually 4, rarely 5 dorsal kineties with
2-4 flm long dorsal cilia; 2 elongated caudal
cirri.

T Y pel 0 c a lit y: Hypertrophic ponds on a

pasture in the Borzsony mountains, Hungary.

M 0 r p hoi 0 g y and b i 0 log y: Ellipti-

cal, both ends rounded. Macronuclear nodules
arranged very closely. Contractile vacuole about
in mid-body, during diastole with longitudinal
collecting canals. Anterior and posterior portion
of cell packed with greyish cytoplasmic crystals.
a b
Rapid movement. Adoral zone of membranelles
about 37 % of body length. Buccal area rather
Fig. 53a, b Oxytricha pseudofurcata (from
wider than narrow. Frontal, ventral, and trans- GELEI 1954b. a, sublimate fixation; b, opal-
verse cirri slightly enlarged; arrangement of blue stain after BRESSLAU). Ventral and doisal
these cirri obviously genus specific. Transverse infraciliature, a = 55 Ilm, b = 50 Ilm.
cirri protrude far beyond posterior end of cell.
Right marginal row terminates adjacent to right
transverse cirrus, usually consists of 13 (sometimes up to 17) cirri, left marginal row usu-
ally with 12 (sometimes up to 16) cirri. Probably 3 dorsal kineties of about body length
and 1 distinctly shortened kinety. Caudal cirri rather long, especially the right one.
174 SYSTEMATIC SECTION

o c cur r e n c e and e colo g y: Probably confined to freshwater habitats. GELEI

(l954b) found this population in 4 hypertrophic, ephemeral ponds in Hungary, described
in detail by GELEI (l954a). According to GELEI et al. (l954), Oxytricha pseudofurcata
was highly abundant in pond III during summer when the water temperature was about
34 °Cl Feeds on bacteria, hyphae, and autotrophic flagellates. Biomass of 106 individuals
about 10 mg.

Oxytricha saprohia KAHL, 1932 (Fig. 54a-c)

1932 Oxytricha saprobia KAHL, Tierwelt Dtl., 25: 603 (Fig. 54a).
1972 Oxytricha saprobia KAHL - BICK, Ciliated protozoa, p 21 (Fig. 54b).
1977 Oxytricha saprobia KAHL 1935 - BERECZKY, Annis Univ. Scient bpest Rolando BOtvOs, 18-19: 169
(Fig. 54c).
1991 Oxytricha saprobia KAHL, 1932 - FOISSNER, BLAlTERER, BERGER & KOllMANN, lnformationsberichte des
Bayer. Landesamtes rurWasserwirtschaft, 1191: 292.

Nom e n c I a t u rea n d t a x 0 nom y: Incorrect subsequent spelling: Oxytricha

saprobica KAHL, 1932 (FOisSNER & FOISSNER 1988, P 90). The illustration in BICK (1972a,
Fig. 54c) looks like a redrawing from KAHL (l932, Fig. 54a). Oxytricha saprobia sensu
VUXANOVICI (1963; Fig. 2201) is insufficiently described. Detailed redescription
necessary.

M 0 r p hoi 0 g y and b i 0 log y: In life about 100 !lm long and 2.7 times as long as
broad. Asymmetrical, margins distinctly converging posteriorly, right one nearly straight,
left convex. Contractile
vacuole slightly in front of
mid-body. Adoral zone of
membrane lies about 37 %
of body length. Distal ado-
ral membranelles con-
spicuously long. Fronto-
ventral cirri distinctly en-

Fig. S4a-c Oxytricha saprobia

(a, from KAHL 1932; b, from BICK
1972a [possibly a redrawing
from KAHL 1932]; c, from BEREC-
ZKY 1977a. a, b, from life; c,

method not indicated). Ventral

views, a, b = 100 J-lm, c = size
not indicated. Note single micro-
b nucleus (arrow).
 Oxytricha 175

larged. Transverse cirri prominent, protrude slightly (left) to distinctly (right) beyond
posterior end of cell.

o c cur r e n c e and e colo g y: Common in freshwater habitats, sometimes abun-

dant according to KAHL (1932) who found O. saprobia between decaying blades of grass,
probably somewhere (near Hamburg?) in Germany. TIRJAKovA (1993) recorded it from
the Turiec River in Slovakia and BERECZKY (l977a) counted 15-25 ind.I·1 in the spring
and autumn plankton of the Danube River, Hungary, under the following conditions:
10-12 °C, pH 7.7-8.0, 7.5-10.2 mg I-I O2, 5.1-7.9 mg I-I Orconsumption, 0.35-1.2
mg 1-1 NH/, 0.03-0.44 mg I-I NOi.
Records from freshwater habitats not substantiated by illustrations: moderately to
heavily polluted miming waters in Austria (AUGUSTIN et al. 1987, BLATIERER 1994, FOISS-
NER & FOISSNER 1988); beta- to alphamesosaprobic Main River and Agger River, Ger-
many (BERNERTII 1982, NusCH 1970); mesosaprobic Danube River, Hungary (BERECKY
1979, BERECZKY et al. 1983, NOSEK & BERECZKY 1981); China (NING et al. 1993, YANG
1989); south-eastern Louisiana, USA (BAMFORTII 1963). VARGA (1953b) recorded it from
fermented farmyard manure. According to PRIMC (1983),0. saprobia prefers low water
temperatures (about 5°C). Terrestrial records not substantiated by illustrations (BAM-
FORTII 1969, LUZZATII 1938).
Feeds on bacteria and flagellates. Biomass of 106 individuals about 34 mg (FOISSNER
et al. 1991). Oxytricha saprobia is classified as alphameso- to polysaprobic indicator:
a-p; a = 6, P = 4, I = 3, SI = 3.4 (FOISSNER 1988a, FOISSNER et al. 1991, SLAoECEK. et al.
1981, WEGL 1983). However, O. saprobia is little known and confusion with other, more
common species (for example, 0. setigera, see below) cannot be excluded. This classifi-
cation thus has to be verified on reliably determined populations.

Oxytricha setigera STOKES, 1891 (Fig. 55a-s, 56a-n, Tables 3, 15, 16)

1891 Oxytricha setigera STOKES, JI R. microsc. Soc., year 1891: 701 (Fig. 55a).
1932 Oxytricha setigera STOKES, 1891- KAHL, Tierwelt Otl., 25: 604.
1932 Steinia balladynula KAHL, Tierwe1t Otl., 25: 614 (Fig. 55b).
1950 Oxytricha plana GELEI & SZABADOS, Annis bioI. Univ. szeged., 1: 268,287 (Fig. 551, m).
1950 Tachysomajusiformis GELEI & SZABADOS, Annis bioI. Univ. szeged., 1: 272, 290 (Fig. 55n).
1963 Oxytricha setigera STOKES - WEST & LUNDIN, Pap. Mich. Acad. Sci., 48: 108 (Fig. 55j).
1977 Oxytricha setigera STOKES - BUlTKAMP, Acta Protozool., 16: 272 (Fig. 55k).
1977 Oxytricha setigera STOKES 1891 - BERECZKY, AnnIs Univ. Scient. bpest. Rolando EOtvOs, 18-19: 169
(Fig. 56n).
1978 Oxytricha setigera- SMITH, Br. Antarct. Surv. Sci. Rep., No. 95: 75 (Fig. 55c).
1982 Oxytricha setigera STOKES, 1891 - FOISSNER, Arch. Protistenk., 126: 83 (Fig. 55d-i; authoritative rede-
scription).
1986 Oxytricha buitkampi ORAGESCO & ORAGESco-KERNEIS, Faune tropicale, 26: 472.
1989 Cyrtohymena balladynula (KAHL, 1932) - FOISSNER, Sber. Ost. Akad. Wiss., 196: 239.
1989 Oxytricha setigera STOKES, 1891- SONG & WILBERT, Lauterbornia, Heft 3: 164 (Fig. 550-s).
176 SYSTEMATIC SECTION

,,
~

, •
•
_t
.,•
•
•
:." . : "."'.\

• . ........ i >!
•.. • •
g .. " ..............
: ", : ....
 Oxytricha 177

1991 Oxytricha setigera STOKES, 1891 - FOISSNER, BLATTERER, BERGER & KOHMANN, Infonnationsberichte des
Bayer. Landesamtes fur Wasserwirtschaft, 1/91: 294.
1991 Oxytricha setigera STOKES, 1891-GSCHWIND, Diplomarbeit, p 59 (Fig. 56a-rn).

Nom e n c I a t u rea n d t a x 0 nom y: KAHL (1932) classified Steinia as subgenus

of Oxytricha; the correct name of S. balladynula in the original description is thus Oxy-
tricha (Steinia) balladynula. Incorrect subsequent spellings: Oxytricha setifera (STOUT
1978, P 17); O. (Steinia) balladynulla (DRAGESCO & DRAGESCo-KERNEIS 1986, P 473);
Steinia balladina KAHL, 1930 (CHARDEZ 1987, P 13).
There are some differences between the original description and the redescription by
FOISSNER (1982), namely in the body shape, the nuclear apparatus, and the number of
marginal cirri. Nevertheless, I agree with FOlSSNER (1982) that his redescription should be
considered as authoritative. It is conceivable that STOKES (1891) made some misobserva-
tions because of the very small size of 0. setigera. Oxytricha balladynula is very likely
synonymous with O. setigera because the size, the long dorsal cilia, the low number of

Fig. 55k--n Oxytricha setigera (k, from BUITKAMP 1977b; I--n, from GELEI & SZABADOS 1950. k, protargol im-
pregnation; I, m, opalblue stain after BRESSLAu; n, sublimate fIxation). Ventral and dorsal infraciliature, k =
40 11m, I, n = 55 11m, m = 35 11m; p 175.

~ Fig. 55a-j Oxytricha setigera (a, after STOKES 1891; b, from KAHL 1932; c, from SMIlH 1978; d-i, from FOlSS-
NER 1982; j, from WEST & LUNDIN 1963. a--t: j, from life; g-i, protargol impregnation). a~, j) Ventral views, a,
c, d,j = 50 11m, b = 60 11m. Should not be confused with Oxytricha balladyna (p 126). e, 1) Dorsal and lateral
view, e = 70 11m, f= 55 11m. g. b) Ventral and dorsal infraciliature, g = 45 11m, h = 40 11m. i) Ventral infracilia-
ture of anterior half of cell, 25 11m; p 175.
 178 SYSTEMATIC SECTION

marginal cirri, and the nuclear apparatus agree rather well with the authoritative rede-
scription; the rectangular outline agrees better with the original description (compare with
Fig. 55a, b, d). KAHL (1932) classified o. balladynula in the subgenus Steinia because the
peristomiallip is distinctly curved anteriorly (Fig. 55b, d, 0), and FOISSNER (1989) trans-
ferred it to Cyrtohymena. Furthermore, Oxytricha plana and Tachysoma fusiformis have
to be synonymised with the present species because there is no character which allows a
reliable separation from Oxytricha setigera. Oxytricha buitkampi was established by
DRAGESCO & DRAGESCo-KERNEIS (1986) for the popUlation by BUITKAMP (1977b; Fig. 55k;
see also FOISSNER 1987d). Oxytricha setigera sensu SONG & WILBERT (1989) agrees more
or less completely with the redescription by FOISSNER (1982). The populations by WEST &
LUNDIN (1963) and SMITII (1978) are rather superficially described, however, the small
sizes and the long dorsal cilia indicate that they in fact observed something like 0.
setigera.
In life O. setigera can be easily confused with the very similar O. balladyna (com-
pare Fig. 55a with Fig. 33a). Thus, an identification should always be confirmed by de-
tailed live observations or protargol impregnation, where the significant difference,
namely the position of the buccal cirrus, is recognisable.

M 0 r ph 0 log y and b i 0 log y: In life 40--60 x 16-21 !lm. Long-oval; according

to FOISSNER (1982) and SONG & WILBERT (1989) posteriorly often somewhat tapered, after
STOKES (1891) and KAHL (1932) body margins nearly parallel. Flattened about 1.5-2 to 1,
according to GSCHWIND (1991) loaf-shaped in lateral view (Fig. 56b). Very flexible,
slightly contractile, especially anterior portion of cell. Macronuclear nodules in life about
10 x 5 !lID, after protargol impregnation 8-12 x 4-7 !lm (mean = 9.1 x 5.3 !lm; n = 10).
Micronucleus in life about 3 !lm across. Contractile vacuole about in mid-body, during
diastole with very inconspicuous collecting canals. Cortical granules absent, cytoplasm
colourless. Rapid movement, sometimes slightly jumping.
Adoral zone of membranelles 25-33 % of body length. Bases of largest mem-
branelles in life about 6 !lm wide. Buccal area deep, in life anteriorly curved like a hook.
Undulating membranes distinctly curved, crossing, that is, in typical Oxytricha pattern.
Frontal and transverse cirri slightly enlarged. Posterior frontoventral cirri arranged be-
hind level of cytostome (Fig. SSg, p, arrow). 8-10, usually 9 frontoventral and postoral
and pretransverse ventral cirri. Invariably 5 (n = 10), in life about 20 !lm long transverse
cirri which protrude far beyond posterior end of cell. Marginal rows begin somewhat be-
hind level of buccal vertex, distinctly separate posteriorly. Marginal cirri in life about
15 !lm long. Dorsal cilia in life 10-15 !lm long, slightly curved, almost perpendicularly
spread in resting animals. Constantly 4 (n = 10) dorsal kineties, kinety 1 distinctly short-

Fig. 55(}-s Oxytricha setigera (from SONG & WILBERT 1989. 0, q, from life; p, r, s, protargol impregnation). 0, ~
q) Ventral and lateral view, 0, q = 45 11m. p, r, s) Ventral and dorsal infraciliature, 45 11m. Arrow in (P) marks
posterior frontoventral cirrus. AZM = adoral zone of membranelIes, BC = buccal cirrus, CC = caudal cirri, CV
= contractile vacuole, FC = right frontal cirrus, PVC = posterior postoral ventral cirrus; p 175.
 Oxytricha 179

AZM= =-"_f~. .• §~
.
FC--t ;
BC--'.S
'(~
1/ t/, '
, ,I ---,
---.!,,(. /, \ I \
, .• \ ,)--CV

.
'--'
•

....
pVC-----"'-'-t .
,
f •

, '
,

CC- ~ "," p
550

," ."

;\
:.,", .'

..•...
.'\ I ...
y\: .. .
•... . 1r
...
 180 SYSTEMATIC SECTION

,'X
,~

~'(-~
. ./§.
:'
.~

#
,:

:?
,
~
f

#
;
, f

.'• ,
t ,
I
#

b • , . #.

# t1
#

# f ,
••t ,
t # I

• '"
• '" d e

Fig_ 56a--e Oxytricha setigera (originals kindly supplied by W. FOISSNER [from GsCHWIND 1991). a, b, from
life; c--e, protargol impregnation). a) Ventral view, 47 )lm. b) Lateral view. Note loaf-shaped outline and fine
dorsal cilia. c, d) Ventral infraciliature of various specimens, c, d = 35 Ilm. Note position of buccal cirrus at
posterior end of undulating membranes. e) Dorsal infraciliature, 35 )lm; p 175.
 Oxytricha 181

0
"0
00 0
0
0

0 h

\,\ .•
I

., '.''.
• •
k • I

Fig. 56f-m Oxytricha setigera (originals kindly supplied by W. FOISSNER [from GsCHWIND 1991). f-m, mor-

phogenetic stages after protargol impregnation; parental cirri white, new cirri black). f-i) Ventral and dorsal
infraciliature of two middle stages, t: g, = 48 11m, h, i = 51 11m. j, k) Ventral and dorsal infraciliature of a late
stage, 48 11m. I, m) Ventral and dorsal infraciliature of a very late stage, 47 11m. 1-6 = frontal-ventral-
transverse streaks; p 175.
 182 SYSTEMATIC SECTION

ened anteriorly, kineties 2 and 3 about of body length, kinety 4 ter-

minates about in mid-body (dorsomarginal row), sometimes con-
sists of2 basal body pairs only (Fig. 55s). 3-4 (mean = 3.5 and 3.9
in 2 populations; n = 10 each) fme, in life about 18 ~m long caudal
cirri.
Morphogenesis of cell division is described by GSCHWIND
(1991; Fig. 56f-m). It commences with the proliferation of basal
bodies between the left transverse cirrus and the left pretransverse
ventral cirrus. This and later stages are reminiscent of the type spe-
cies, 0. granulifera. Dorsal morphogenesis is in Urosomoida pat-
tern. Some comments on morphogenesis, but no figures, are given
by TARTAR (1941).

Fig. 560 Oxytricha 0 c cur r e nee and e colo g y: Widely distributed through-
seligera (from BERE- out the year in lentic and lotic freshwater habitats and in soil, often
CZKY 1917a). Ventral abundant. Locus classicus is a pond near Trenton in New Jersey,
view from life, size USA (STOKES 1891). Records largely substantiated by illustrations:
not indicated; p 175.Austria (AUGUSTIN et al. 1987, FOISSNER 1987b, FOISSNER & FOISSNER
1988); beta- to alphamesosaprobic brook in Bavaria (GSCHWIND
1991); Poppelsdorfer Weiher, a eutrophic pond in Bonn, Germany (SONG & WILBERT
1989); mesosaprobic Danube River (35 ind.I>I) and rain puddle in Szeged, Hungary
(BERECZKY 1977a, b, BERECZKY et al. 1983, GELEI & SZABADOS 1950); freshwater habitats
in Michigan, USA (LUNDIN & WEST 1963, WEST & LUNDIN 1963); soil of a "savane
briHee", Plateau du Grand Nord, Ivory Coast and soil of a deciduous forest near Bonn,
Germany (BUITKAMP 1977a, b, 1979); various terrestrial habitats in Australia (BLATTERER
& FOISSNER 1988); soil from South Georgia, South Shetland, South Orkney, Signy, and
Deception Islands, Antarctica (SMITH 1972, 1973a, 1974a, 1978, 1985).
Records from terrestrial and freshwater habitats not substantiated by illustrations:
mesosaprobic running waters in Upper Austria (AOOLR 1994b, BLATTERER 1994, FOISSNER
& MOOG 1992, own observations); soil in Belgium (CHARnEZ 1967); Bulgarian rivers
(DETCHEvA 1979b, 1981, RussEv et al. 1976); forest and agricultural soils and freshwater
in Czechoslovakia (MATIS & TIRJAKovA 1994a, ROSA 1957a, b, 1962, TIRJAKovA 1988,
1992b, 1993); soil of a beech wood in Denmark (STOUT 1968); soil from East Greenland
(STOUT 1970); sediment of Loch Leven, England (BRYANT & LAYBOURN 1974); clean and
beta- to alphamesosaprobic brooks and rivers in Bavaria (FOISSNER 1997a, FOISSNER et al.
1992a, b); aufwuchs of a mesotrophic Eifel Maar lake, Germany (PACKROFF 1992, PACK-
ROFF & WILBERT 1991); unpolluted foothill stream (Breitenbach) near the village of
Schlitz, Germany (PACKROFF & ZWICK 1996); beech-forest and limed spruce forest in Ger-
many (LEHLE 1989, 1993, 1994); Italy (GRISPINI 1938); Latvian river (LIEPA 1973); La Ja-
rosa Reservoir, Spain (FERNANDEZ-LEBORANS et al. 1990); Danube River, Slovakia (SZEN-
TIVANy & TIRJAKovA 1994); soil influenced by herbicide, USSR (GEL'CER & GEPTNER
1976); stream in India (RANGANATHAN & SHREEDHARAN 1986); leaf-litter in a pond near
 Oxytricha 183

Tokyo and soil in Japan (HATANO & WATANABE 1981, TAKAHASID & SUHAMA 1991); lakes,
rivers, eutrophic ponds, upper soil horizon of a Delta hardwood bottomland forest, soil of
a cedar glade, and Mississippi deltaic soils, USA (BAMFORTH 1967, 1969, 1995, BOVEE
1960, CAIRNS 1965a, CAIRNS & YONGUE 1966, MARTIN & SHARP 1983, McCASHLAND 1956,
PATRICK 1961, PRATT et al. 1986); New River in Virginia, USA, at l3-19 °C, pH 7.0-7.3,
and 7-8 mg I-I O2 (YONGUE & CAIRNS 1979); limnetic habitats on the campus of the Uni-
versity of Costa Rica (Rmz 1961); soil in Peru, Brazil, Costa Rica, and Australia (FOISS-
NER 1995b, 1997b); beech wood soils, inundated soil under grassland, and burnt sites in
New Zealand (STOUT 1961, 1963, 1984); bryophyte peat and angiosperm soil from the
sub-Antarctic island of South Georgia (SMITH 1982).
Some autecological data of freshwater populations are listed in Table 16. BUITKAMP
(1979) found 0. setigera mainly in soil samples incubated at 30°C and 35 °C. Detailed
autecological data of various terrestrial populations are given in FOISSNER & PEER (1985)
and FOISSNER et al. (1985).
Feeds on bacteria and heterotrophic flagellates. STOUT (1956) cultured Oxytricha
setigera in peat extract. Biomass of 106 individuals after FOISSNER (1987a) about 8 mg, af-
ter NESTERENKO & KovALCHUK (1991) about 12 mg.
Oxytricha setigera is classified as alpha- and betamesosaprobic indicator of water
quality: a-b; b = 4, a = 6, I = 3, SI = 2.6 (FOISSNER 1988a, FOISSNER et al. 1991, MO-
RAvcovA 1977, SLADECEK et al. 1981, WEGL 1983). This largely agrees with the available
autecological and faunistic date. However, since it can not be excluded that it was some-
times mixed up with the rather similar O. balladyna, further autecological investigations
on reliably determined populations are needed.

Oxytricha simi/is ENGELMANN, 1862 (Fig. 57a-d, Tables 16, 17)

1861 Oxytricha pel/ionella (EHR. STEIN) - WRZESNIOWSKI, AnnIs Sci. nat., 16: 333 (Fig. 57d; misidentifica-
tion).
1862 Oxytricha similis ENGELMANN, Z. wiss. Zool., II: 388 (no figure).
1882 Opisthotricha simi/is, ENG. sp. - KENT, Manual infusoria II, p 785.
1932 Opisthotricha (Oxytricha) similis ENGELMANN, 1862 - KAHL, Tierwelt Dtl., 25: 611 (Fig. 57a; authorita-
tive redescription).
1935 Opistholricha similis (ENGELMANN) KAHL 1932 - WANG & NIE, Sinensia, 6: 502 (Fig. 57b).
1972 Opisthotricha simi/is ENGELMANN - BlcK, Ciliated protozoa, p 21 (Fig. 57c).
1991 Oxytricha similis ENGELMANN, 1862 - FOISSNER, BLATTERER, BERGER & KOHMANN, lnformationsberichte
des Bayer. Landesamtes fur Wasserwirtschait, 1191: 300.

Nom e n cIa t u rea n d t a x 0 nom y: KAHL (1932) classified Opisthotricha as

subgenus of Oxytricha; the correct name in his revision is thus Oxytricha (Opisthotricha)
similis, which is a secondary homonym of O. (Histrio) simi/is in KAHL (1932, P 615).
Oxytricha similis is a slender, medium-sized species with 2, more or less prominent cau-
dal and 5 transverse cirri. The 2 caudal cirri are arranged rather symmetrical in the popu-
lation of WANG & NIE (1935), whereas both point slightly to the right in the other popula-
184 SYSTEMATIC SECTION

Fig. 57a-d Oxytricha simi/is (a,

from KAm. 1932; b, after WANG
& NIE 1935; e, after 81CK 1972a;
d, after WRZESNlOWSKI 1861. a, b,
d, from life; e, from life?). Ven-
tral views, a = 90 Jim, b = 65 Jim,
e = 105 Jim, d = 80 Jim; p 183.

tions, suggesting that they

are not conspecific. The
population by REuTER
(1961) has 3 caudal cirri
and is classified as O.
multiseta (Fig. 51 b). The
populations by CuRDs
b (1969; Fig. 221h) and
PATSCH (1974; Fig. 230h)
are insufficiently rede-
scribed. Oxytricha similis
should not be confused
with Tachysoma pellionellum, which has long dorsal cilia and a single micronucleus be-
tween the macronuclear nodules and lacks caudal cirri. The rather similar Oxytricha
longa and Urosomoida agiliformis have fewer transverse cirri. Identification should be
confmned on protargol-impregnated material. Detailed redescription necessary.

M 0 r p hoI 0 g y and b i 0 log y: In life about 80-100 J,lm long, 3.8 times as long as
broad (KAHL 1932); according to WANG & NIE (1935) in life 60-90 x 22-27 J,lm. Long el-
liptical, according to ENGELMANN (1862) margins almost parallel. Contractile vacuole
slightly in front of mid-body.
Adoral zone of membranelles about 27 % of body-length. Arrangement of frontal,
ventral, and transverse cirri obviously genus-specific. Five transverse cirri, protrude dis-
tinctly beyond posterior end of cell. Dorsal cilia 2-4 J,lm long. Some biochemical data,
for example, DNA content of micronucleus, are given by STEINBROCK et al. (1981; see
also KRAUT et aI. 1986) and SCHLEGEL & STEINBROCK (1986); however, the identifications
are not substantiated by illustrations.

o c cur r e n c e and e colo g y: In the aufwuchs and detritus of mesosaprobic wa-

ter bodies; not as common as, for instance, Tachysoma pellionel/um. Locus classicus is a
freshwater habitat near Leipzig, Germany (ENGELMANN 1862). WRZESNIOWSKI (1861)
found "0. pellionelld' in a pond in Mokotow near Warsaw, Poland. KARL (1932) ob-
served Oxytricha similis repeatedly in mesosaprobic detritus, and WANG & NIE (1935)
collected it from a pond near Tsing-Liang-Shan in Nanking, China.
 Oxytricha 185

Table 16 Autecological data of Oxytricha setigera (columns 1-5) and O. simi/is (columns 6, 7; see also col-
umn 7 ofTable 17). References: column 1, from PATRICK et al. (1967; 9 analyses from Savannah River, USA);
column 2, from CAIRNS & YONGUE (1973a; 2 areas of Cape Fear River in near Fayetteville, North Carolina,
USA); column 3, from DETCHEVA (1983c; rarely in Maritza River, Bulgaria); column 4, from BERECZKY
(1977a; high abundance in the summer plankton of Danube River in Hungary); column 5, from FOISSNER et al.
(1991; various running waters in Austria, n = 12-14, I1coo = 6, nN02-N = 9); column 6, from MAooNi & GHErn
(1981a; few records from Torrente Stirone, Northern Italy); column 7, from DETCHEVA (1983c; 1 record from
Maritza River, Bulgaria)

Parameter' Reference
2 3 4 5 6 7
Saprobity a a
Frequency (%) 0.6 0.3
Temperature (0C) 14.5-29 23-27 22.8 22 3.3-10.7 18.5-19 25
pH 6.5-7.0 6.6--7.3 6.6--7.4 7.9 7.0-8.7 7.8-8.5 7.6
O2 (mgl·') 6.3-10.7 6.6--9.5 3.5-6.2 8.9 7.8-13.1 7.3-8.6 4.6
O2 (% saturation) 41-73 74-113 57
BODs (mg I·') 0.5-<3 1-6 11-12 7.1 0.7->7.8 7.2
Total hardness edH) 3.0--12.8
CO2 (mgl-') >1-10
HC03• (mg I·') 213-284 277
KMnO.-consumption 2.4-25
COD (mgl·') 7-15 44-56
NH/-N (mg I·') 0.001-0.17 0.005-0.7 0.04 0.23 0.008-0.95 0.55-1.8 0.04
NOi-N (mg I·') 0.07 <0.001-0.01 0.11-2.2
N03·-N (mg I·') 0.07-<0.5 0.25-0.84 0.4-1.1 0.61 0.5-6.6 0.7-1.2 0.18
POl--p (mg I·') 0.012-0.1 0.05-0.28 0.008-0.66 0.35-1.0
C1" (mg 1-') 1-<10 12.5-34 17-28 0.9-90 88.5-1094 24.5
Fel +(mg I·') 0.1-0.6 0.07-1.5 0.04
Mg2+(mgl·') <3-3 1.7-5 21 22
Cal +(mg I·') 4-10 60-69 70
SOl- (mg 1-') 2-11 5.7-18 75-167 131

, The figure "0" denotes only that the parameter in question could not be detected with the analytical method
used.

Records from freshwater habitats not substantiated by illustrations: mesosaprobic

brooks and rivers in Upper Austria (AooLR 1992, 1993a-c, 1994a, 1995a-c, BLATIERER
1994); rare in Maritza River, Bulgaria (DETCHEVA 1981); moss samples with vatying wa-
ter content and running waters in Czechoslovakia (MATIS & TIRJAKovA 1994a, TIRJAKovA
& MATIS 1987a); underground water in Czechoslovakia (MoRAvcovA 1986); slime of pol-
luted waters in England (CURTIS & CURDS 1971); sewage fishponds in Bavaria, Germany
(KAUFMANN 1958); Illach River, Germany (FOISSNER 1997a); freshwater basins with or-
ganic material from the vicinity of Bonn, Germany (BICK 1967a); alphamesosaprobic
river near Krefeld, Germany (REuss 1976); very abundant in sandy detritus in Hamburg
Harbour, Germany (BARTSCH & HARTWIG 1984); aufwuchs and detritus drifts in Lake
Balaton, Hungary (GELLERT & TAMAs 1958, TAMAs & GELLERT 1958); rock pool with
186 SYSTEMATIC SECTION

Table 17 Autecological data of Oxytrichajallax (columns 1-5), Gastrostyla muscorum (column 6), and Oxy-
tricha similis (column 7; see also columns 6, 7 of Table 16). References: column t, from LACKEY (1938a; vari-
ous Iimnetic and marine habitats in the USA, n = 14); column 2, from CAIRNS & YONGUE (1973a; Cape Fear
River near Fayetteville, North Carolina The values are from "Station 2, year 1970"; no detailed information on
sample size); columns 3 and 7, from BICK & KUNZE (1971; summary of literature data); column 4, from MA-
DONI & GHETII (1981a; Torrente Stirone in Northern Italy. The values are from "Station 8, I.F."); column 5,
from DETCHEVA (1972a, 1975a-c, 1976b, 1977, 1979a, c, 1983a; many analyses from Bulgarian running wa-
ters. The data of DETCHEVA 1971, 1975a are identical); column 6, from DETOIEVA (1971, 1975a; 1 analysis
from a Bulgarian running water. The data of these tVlO references are identical). Some further data of these ref-
erences are given in the text

Parameter' Reference
2 3 4 5 6 7
Frequency (%) 7.3 I.1-S.8 0.9
Temperature COC) 25-27 2-S6 1.8-7 7-24 12
pH 4.6-7.5 6. 6-fJ. 7 6.8-7.5 7.6-7.9 6.8-8.3 7.2 8.9-9.8
O2 (mg I·') 6.9-11.4 3-S 11.5-12.7 0.8-10.7 2.3 0-6.3
O2 (% saturation) 9-97 21
BODs (mg I·') 3.5-10.3 2-27 S8
NH:-N (mg I·') 0.03-0.1 0.23-0.4 0.03-1.1S 0-12.4
N02·-N (mg I·') 0.02-0.04 0-0.34 O.ll
NO,·-N (mg I·') 0.2S-O.4 1.6-3.2 O-ll 4.27
POl--p (mg I·') 0.06-0.07 0.17-0.2

, The figure "0" denotes only that the parameter in question could not be detected with the analytical method
used.

0.34 % salinity near the Biological Station Espegrend, Norway (REuTER 1963a); Warsaw,
Poland (WRZESNIOWSKIEGO 1966); 54 specimens per 25 ml sediment in a eutrophic reser-
voir near Barcelona, Spain (SALvA06 CABRE 1993, SALVADO & GRACIA 1991; further re-
cords from Spain: FERNANDEZ-LEBORANS et al. 1990, FRANCO et al. 1995, GRACIA & IGUAL
1987b, GRACIA et al. 1989, MARGALEF LOPEZ 1945); Turiec River in Slovakia (TIRJAKovA
1993); Danube River, Slovakia (SZENTIVANY & TIRJAKovA 1994); low salinity lagoons of
the Caspian Sea (AGAMALIEV 1973, 1986); lakes in Azerbaijan (ALIEV 1982b); China
(SHEN 1980, SHEN & JIANG 1979, SHEN et al. 1995, WANG et al. 1992, YANG 1989); lakes
and ponds in the USA (CAIRNS et al. 1973, HENEBRY & CAIRNS 1980, YONGUE & CAIRNS
1971); Des Plaines River wetlands and New River, USA (HENEBRY & GORDEN 1989,
YONGUE & CAIRNS 1979).
Terrestrial records not substantiated by illustrations (possibly confused with Uroso-
moida agiliformis or Oxytricha longa): soils in Szeged, Hungary (HORVATH 1950); dry
mosses in Bratislava, Czechoslovakia (TIRJAKovA & MATIS 1987b).
Feeds on bacteria, diatoms, filamentous growths, and small colourless flagellates
(BICK 1968a, BICK & KUNZE 1971, CURTIS & CURDS 1971). Biomass of 106 individuals
about 14 mg (FOISSNER et al. 1991). Some autecological data are shown in Tables 16, 17.
Oxytricha similis is abundant at alkaline conditions and at high bacterial abundances, es-
 Oxytricha 187

pecially in spring and autumn; tolerates 3.7 % salinity (inland natron lake water; BICK
1964).
MATIS (1975) found 0. similis in road pools in Bratislava, Czechoslovakia, at 1 to
11.5 °C, pH 5.5-6.4, and 5.8-8.4 mg 1.10 2 and MATIS & STRAKOVA-STRIESKOvA (1991) re-
corded it from a brook and 2 thennallakes in Czechoslovakia at 20-38 °C and pH 5-7.3.
LEGNER (1975) counted up to 39 indo mI·1 twelve days after the addition of 0.2 g 1"1 glu-
cose and 0.3 g I-I peptone. Eurythennic (1-38 0c) with an optimum at 10°C (BICK &
BERTRAM 1973, MATIS 1975, MATIS & STRAKOVA-STRIESKOvA 1991, PRIMC 1983).
Oxytricha similis also occurs in percolating filters and activated sludge plants, but is
not recorded from Imhoff tanks (BANINA 1983, CURDS 1975, CURDS & COCKBURN 1970a,
KUTIKOWA 1984). According to CURDS & COCKBURN (1970b), it is characteristic for
activated-sludge plants having effiuents of high quality with a BODs <20 mg 1-1.
Oxytricha similis indicates beta- and alphamesosaprobic conditions: b-a; b = 5, a =
5, I = 3, SI = 2.5 (FOISSNER 1988a, FOISSNER et al. 1991, SLADECEK et al. 1981, WEGL
1983). This classification largely agrees with the faunistic records and the ecological
data. However, the autecology and saprobiology of this species has to be confirmed on
reliably determined populations.

Oxytricha siseris VUXANOVICI, 1963 (Fig. 58a-h, Table 15)

1963 Oxytricha siseris VUXANOVICI, Studii Cerc. BioI., 15: 215 (Fig. 58a).
1974 Tachysoma siseris (VUXANOVICI, 1963) - STILLER, Annis hist-nat. Mus. nato. hung., 66: 133.
1982 Oxytricha siseris VUXANOVICI, 1963 - FOISSNER, Arch. Protistenk., 126: 85 (Fig. 58b--h; authoritative re-
description; I slide of protargol-impregnated specimens is deposited in the OberOsterreichische Landes-
museum in Linz, Upper Austria).

T a x 0 nom y: The original description is somewhat superficial. Thus, the redescription

by FOISSNER (1982), which largely agrees with the original description, should be consid-
ered as authoritative. Oxytricha siseris differs from O. setigera, among others, in the
number of micronuclei and in the shape and arrangement of the undulating membranes,
which is not in the typical Oxytricha pattern. Cannot be synonymised with Tachysoma
pellionellum as suggested by BORROR (l972a).

M 0 r p hoI 0 g y and b i 0 log y: In life 80-120 x 35 J.lm; after VUXANOVICI (1963)

only 60-80 J.lm long in life. Slenderly oval, margins often converging posteriorly, thus
posterior end usually narrowly rounded. Macronuclear nodules 11-13 x 5-9 J.lm (mean =
11.7 x 7.0 J.lm; n = 11) after protargol impregnation, arranged slightly left of median, al-
ways distinctly separated. Each macronuclear nodule with a single, after protargol im-
pregnation about 2.3-2.7 J.lm (n = 11) sized, globular micronucleus. Contractile vacuole
only slightly in front of mid-body, during diastole with 2 long collecting canals. Cortical
granules absent. Cytoplasm colourless, posteriorly often with many cytoplasmic crystals.
 188 SYSTEMATIC SECTION

Fig. 58a~ Oxytricha siseris (a, from VUXANOYlCI 1963; b-e, from FOISSNER 1982. IK, from life; d, e, protar-
gol impregnation). a, c) Ventral views, a = 60 Jim, c = 85 Jim. b) Lateral view, 85 Jim. d, e) Ventral and dorsal
infiaciliature, d, e = 75 J.1m; p 187.

Rapid, jerky movement, also swims very fast by rotation about main body axis, rarely
gliding.
Adoral zone of membranelies about 113 of body length, middle portion slightly dis-
placed to left margin (Fig. 58d, h). Buccal area very small, distinctly excavated, anteri-
orly hooked. Peristomiallip motile, with peculiar short appendages which do not impreg-
nate with protargol (Fig. 58h). Paroral moderately long, slightly bent, consists of2 rows
of basal bodies, arranged in middle portion of buccal area, distinctly in front of endoral.
Endoral almost straight to distinctly curved, probably consists of 2 rows of basal bodies
too. Frontal and ventral cirri only slightly, transverse cirri distinctly larger than marginal
cirri. According to VUXANOVICI (1963) and FOIsSNER (1982), only 3 frontoventral cirri pre-
sent (Fig. 58a, d); in my opinion, however, the anterior-most cirrus on the postoral area
corresponds to the posterior-most frontoventral cirrus of other Oxytricha species. Post-
oral ventral cirri somewhat displaced posteriorly (Fig. 58g, arrowheads). 1 or 2 pretrans-
verse ventral cirri (Fig. 58d, g). Figure 58g is identical to Figure 58d except that the ante-
rior pretransverse ventral cirrus located to the left at about the level of the posterior-most
 Oxytricha 189

#
,
,
,
,

9 h

Fig. 58f-b Oxytricha siseris (f, h, from FOISSNER 1982; g, from FOISSNER 1982 slightly modified. f, from life;
g, h, protargo! impregnation). t) Dorsal view, 90 J.lIl1. g) Ventral infraciliature as in Figure S8d. The arrow-
heads mark the 3 postoral ventral cirri; further details, see text. b) Ventral infraciliature of anterior half of cell,
bar = 20 ~m. e = endoral, PL = peristomiallip, p = paroral; p 187.

postoral ventral cirrus is shown (reconstructed after unpublished microphotographs

kindly supplied by W. ForSSNER). Usually 5, rarely 6, in life about 40 11m long transverse
cirri which protrude far beyond posterior end of cell. Both marginal rows begin at about
level of buccal vertex. Marginal cirri in life about 30 11m long; distance widens posteri-
orly. Dorsal cilia in life 13 11m (anterior half) to 20 11m (posterior half) long, motile. Usu-
ally 5, rarely 6 dorsal kineties. Kinety 1 distinctly shortened anteriorly, consists of 18-30
basal body pairs. Constantly 3 (n = 11), in life up to 40 11m long, prominent motile caudal
cirri on dorsal kineties 1,2, and 3 (Fig. 58e); however, morphogenesis will possibly show
that they originate from kineties 1,2, and 4, as in many other species.

o c cur r e n c e and e col 0 g y: Rare, in freshwater and terrestrial habitats. Locus

classicus is a small basin in the botanical garden of Budapest, Romania (VUXANOVICI
1963). ForSSNER (1982) found Oxytricha siseris in the alluvial soil of two xerothermic,
uncultivated grassland sites and in two wheat fields in the Tullner Feld, Lower Austria;
for detailed descriptions ofthese sites and autecological data, see FOISSNER & PEER (1985,
 190 SYSTEMA TIC SECTION

"Profile 1, 3, 4, and 6"). FERNANDEZ-LEBORANS & ANTONIO-GARCIA

(1988) and FERNANDEz-LEBORANS et al. (1987) provided some data on
the effects of the heavy metals lead, zinc, mercury, and cadmium. They
obtained their samples from the Manzanares River, La Pedriza, Spain.
TIRJAKovA (1988) found 0. siseris in 4 of 145 soil samples from differ-
ent fields in Czechoslovakia. Soil in Brazil (FOISSNER 1997b). Probably
feeds on bacteria. Biomass of 106 cells about 30 mg (FOISSNER 1987a).

Oxytricha sphagni KARL, 1932 (Fig. 59)

1932 Opisthotrieha sphagni KAHL, Tierwelt Dtl., 25: 608.

Nom e n c I a t u rea n d t a x 0 nom y: KAHL (1932) classified

Opisthotricha as subgenus of Oxytricha; thus the correct name in the
original description is Oxytricha (Opisthotricha) sphagni, which is a
junior secondary homonym of Oxytricha (Histrio) sphagni in KAHL
(1932). Detailed redescription necessary.
Fig. 59 Oxytri-
eha sphagni
(from KAHL M 0 r ph 0 log y and b i 0 log y: In life about 100-120 Ilm long.
1932). Ventral Long-elliptical. Contractile vacuole slightly in front mid-body. Adoral
view from life, zone of membranelles about 27 % of body length. Frontal and trans-
100-120 Ilm. verse cirri only slightly enlarged, transverse cirri distinctly protruding
Note single mi-
cronucleus.
beyond posterior end of cell. Marginal cirri displaced inwards. Dorsal
cilia in life about 15 Ilm long, where not spread but trailing behind.
Three distinctly elongated caudal cirri.

o c cur r e n c e: Obviously vel)' rare. KAlIL (1932) discovered O. sphagni in

Sphagnum, probably somewhere (near Hamburg?) in Germany. Records not substanti-
ated by illustrations: Tibetan Plateau (WANG 1977); Yuelushan Area, China (YANG 1989).
Biomass of 106 individuals about 17 mg.

Oxytricha sp. (Fig. 60, Table 15)

1986 Oxytrieha sp. - DRAGESCO & DRAGESCo-KERNilIS, Faune tropica/e, 26: 468.

Nom e n c I at u rea n d t a x 0 nom y: Vel)' probably this form is not a true Oxytri-
cha because the ventral infraciliature differs significantly from the typical Oxytricha pat-
tern. The somewhat increased number of transverse cirri is reminiscent of O. multiseta
and 0. variabilis, from which it differs, however, in the arrangement of the ventral cirri.
Redescription (live observation, morphometric characterisation, dorsal infraciliature)
necessary.
 Oxytricha 191

Fig. 60 Oxytricha sp. (from DRAGESCO & DRAGESCO-KERNEIS 1986).

Ventral infraciliature, protargol impregnation, 105 flm; p 190.

M 0 r p hoi 0 g y and b i 0 log y: After protargol im-

pregnation about 95 x 32 !lm. Both ends rounded. Macro-
nuclear nodules after protargol impregnation 14-19!lm
(mean = 15 !lm; n = 12) long, micronuclei not observed.
Adoral zone of membranelles about 36 % of body length.
Buccal area large. Buccal cirrus at about level where un-
dulating membranes cross. Arrangement of frontal-ventral- I'·:~
transverse cirri rather unusual. 6 transverse cirri; 2 small '01
pretransverse ventral cirri at about level of anterior-most ':.:1
transverse cirrus. Four not conspicuously elongated caudal f':J
cirri in gap between ends of marginal rows. Dorsal infra-
ciliature not described, dorsal cilia probably short.
i'l
: :.~

..~.
o c cur r e n c e: Freshwater in Benin (Africa).
• ••
Oxytricha ten ella SONG & WILBERT, 1989 • •
(Fig. 61a-e, Table 15) f , ,· ,.;j/!

1989 Oxytricha tenella SONG & WILBERT, Lauterbomia, 3: 166 (the

slide ofholotype specimens is deposited in the College of Fisher-
ies, Ocean University ofQingdao, China).

Nom e n c I a t u rea n d t a x 0 nom y: Oxytricha tennella in SONG & WILBERT

(1989, P 219) is an incorrect original spelling. The body shape is reminiscent of O.fallax,
which is, however, at least twice as long and has no cortical granules. The terrestrial spe-
cies O. proximata is 70-140 !lm long and also lacks cortical granules. The terrestrial 0.
granulifera is somewhat longer (80-130 !lm) and has differently arranged frontoventral
cirri.

M 0 r ph 0 log y and b i 0 log y: In life 50-70 x 30-40 !lm. Posterior end broadly,
anterior distinctly more narrowly rounded (Fig. 61b, c). Macronuclear nodules in life
about 10 x 15 !lm, after protargol impregnation 9-15 x 7-11 !lm (mean = 10.9 x 8.2 !lm;
n = 14 and 10, respectively), nodules very closely arranged (Fig. 61a, e). 2-3 (mean =
2.2; n = 13) micronuclei. Contractile vacuole in about mid-body or slightly in front; dur-
ing diastole with short, inconspicuous collecting canals. Cortical granules irregularly ar-
ranged (apparently in several layers); colour, shape, and size of granules not mentioned.
Cytoplasm slightly yellowish, often dark greyish due to inclusions. Moderately rapid
movement.
 192 SYSTEMATIC SECTION

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~~'/) .• 0o"o·.oJJD';';. ' ~ ;;'0' ~" .

a •• ~. 0.:' .~t:"; ~; ".~' ,;',.>~:;~ :
~1J';'-,,?'o:o;oo , .'"
- --(~;;':~'.~:;r!'
c

..
...
,.
.., , ~,I
4

",, , !
\. ,
, ,
, \-
(., .,. .•
,,
~

'- .. \
'\. I.. !
,/

, fl' ,
,,
,, , • . -X ,
\

d
,~~
cc r-\\ e

Fig. 61a4! Oxytricha tene/fa (from SONG & WILBERT 1989. a-c, from life; d, e, protargol impregnation). a-c)
Lateral, ventral, and dorsal view, a = 65 11m, b = 70 11m, c = 75 11m. Cortical granules are only shown in (c). d)
Ventral infraciliature, 55 11m. Arrow marks posterior frontoventral cirrus. e) Dorsal infraciliature, 55 11m,
 Oxytricha 193

Adoral zone ofmembranelles about 37 % of body length. Undulating membranes al-

most straight, crossing at about middle portion. Buccal cirrus at anterior end of undulat-
ing membranes. Constantly (n = 9) 18 frontal, ventral, and transverse cirri. Frontal and
transverse cirri enlarged, all other cirri rather fme and short (8-12 ~m). Posterior fronto-
ventral cirrus (Fig. 61d, arrow) distinctly separate from the other 3 cirri. Transverse cirri
protrude only slightly beyond posterior end of cell. Marginal rows nearly uninterrupted
posteriorly (Fig. 61d). Dorsal cilia in life 2-4 ~m long, according to SONG & WILBERT
(1989) constantly (n = 8) arranged in 4 rows, each of body length, kinety 4 often irregu-
larly interrupted; however, in my opinion, kinety 4 in Figure 61 e in fact consists of 2 ki-
neties, namely an anterior dorsomarginal row and a posterior fragment of kinety 3. In-
variably 3 (n = 9), in life only 5-7 ~m long caudal cirri, probably at end of kineties 1,2,
and 4.

o c cur r e n c e and e colo g y: Locus classicus is Bonn, Germany, where SONG &
WILBERT (1989) discovered Oxytricha tenella in a brook (Melbbach) about 50 m above
the Poppelsdorfer Weiher, a eutrophic pond. Feeds on small algae and flagellates. Bio-
mass of 106 individuals about 20 mg. Not found since.

Oxytricha variabilis GROLIERE, 1975 (Fig. 62)

1975 Oxytricha variabilis GROUERE, Protistologica, 11: 490.

Nom e n c I a t u rea n d t a x 0 nom y: BORY DE SAINT-VINCENT in LAMOUROUX et

al. (1824, p 597) used the almost homonymous binomen Oxitricha variabilis, which is
obviously only an unjustified replacement name for Trichoda praeceps MOLLER, 1786
(see species indeterminata and EHRENBERG 1834, P 1203). Because the original spellings
of BORY's and GROLffiRE'S name deviate in one letter of the genus name I do not replace
GROLIERE'S name. Oxytricha variabilis is described only after protargol impregnation;
thus live observations (for example, cortical granules lacking or present?) and mor-
phometric characterisation necessary.

M 0 r p hoI 0 g y and b i 0 log y: After protargol impregnation 88-115 x 27-40 ~m

(mean = 100 x 32 ~m; n = ?). Body margins nearly parallel, both ends broadly rounded.
Macronuclear nodules ovoid or spherical, about 15-17 ~m long, usually each nodule
with 2 micronuclei.
Adoral zone of membrane lIes about 40 % of body length, 28-32 membranelles. Ac-
cording to microphotographs by GROLIERE (1975), endoral distinctly shorter than paroral.

~ According to SONG & WILBERT (1989), dorsal kinety 4 is interrupted (arrowhead and arrow) in mid- body. In
my opinion, arrow marks anterior end ofkinety 4 and arrowhead denotes posterior end ofkinety 5. Kinety 5 is
probably a dorsomarginal kinety. CC = caudal cirri, X = gap between posterior end of marginal rows, 4 = dor-
sal kinety 4; p 191.
 194 SYSTEMA TIC SECTION

Fig. 62 Oxytricha variabilis (from GROUERE 1975). Ventral infi'aciIiature, pro-

targol impregnation, 100 11m. Arrowhead marks frontoventral cirrus 1M. Note
supernumerary ventral cirri; p 193.

Buccal cirrus at anterior end of undulating membranes. Frontal

cirri only slightly enlarged. Number of ventral and transverse
cirri rather variable. Cirrus IIII2 (Fig. 62e, arrowhead) arranged
on left, between migratory cirri, similarly to in the type species
0. granulifera. Up to 5 postoral ventral cirri. ~ transverse
cirri, do not protrude on left, protrude slightly beyond posterior
end of cell on right. 5 dorsal kineties, dorsal cilia 2-4 /lm long.
Caudal cirri apparently inconspicuous.

o c cur r e n c e and e col 0 g y: Locus classicus is a

pond in the forest between Randan and St-Yorre, France. A de-
tailed description of this site is given by GROuERE & NJINE
(1973). Feeds on bacteria. Biomass of 106 individuals about
27 mg. FERNANDEZ-LEBORANs et al. (1987) provided some data
on the effects of the heavy metals lead, zinc, mercury, and cad-
mium; they obtained their samples from the Manzanares River,
La Pedriza, Spain. SlRODER-KypKE (1996) recorded Oxytricha variabilis from the auf-
wuchs of dystrophic lakes in Germany.

Oxytricha auripunctata BLATIERER & FOISSNER, 1988 (Fig. 63a--e,

Table 15)

1988 Oxytricha auripunctata BLATIERER & FOISSNER, Stapfia, 17: 61 (1 slide ofholotype specimens is depos-
ited in the Oberosterreichische Landesmuseum in Linz, Upper Austria).

T a x 0 nom y: The decreased number and the small size of the transverse cirri and the
arrangement of the frontoventral cirri is reminiscent of Urosomoida. BLATIERER & FOlss-
NER (1988) could not count the number of dorsal kineties exactly, but they supposed that
it has 5. Since all Urosomoida species invariably have 4 kineties, I agree with the original
classification. Oxytricha auripunctata differs from other soil species with cortical gran-
ules (0. granulifera, 0. rubripuncta, O. longigranulosa) in morphometric characters and
the shape and colour of the cortical granules. Oxytricha auripunctata should not be con-
fused with the terrestrial Urosomoida dorsiincisura, which also has yellowish cortical
granules, but ~ macronuclear nodules.

M 0 r ph 0 log y and b i 0 log y: In life 80-130 x 25-35 /lm. Elliptical, both ends
moderately narrowly rounded, right margin usually straight, left distinctly convex. Two
 Oxytricha 195

, # ,
, ,

e
, ,
,
, .-.'
'.':'-'
, ...:-.-,'
:~
, ,
,
,

c e

Fig.63a---e Oxytricha auripunctata (from BLAITERER & FOISSNER 1988. a---d, from life; e, protargol impregna-
tion). a) Ventral view, 90 11m. b) Part of pellicle with irregular rows of cortical granules. C, d) Lateral and dor-
sal view. e) Ventral infraciliature, 90 11m. Arrow marks right frontal cirrus; p 194.

macronuclear nodules, after protargol impregnation 12-16 x 3.8---6.3 ~m (mean = 14.5 x

4.9 ~m; n = 14). 1-5 (mean = 2.3; n = 14) spherical micronuclei, after protargol impreg-
nation 1.4-2.1 ~m (mean = 1.5 ~m; n = 14) across. Contractile vacuole about in mid-
body, during diastole with 2 collecting canals. Cortical granules about 0.5 ~m across,
orange-yellow, stain with protargol, arranged around cirri and in rather disorganised rows
on ventral surface (Fig. 63b). Cytoplasm filled with 2-4 ~m sized, greasy globules and
many 3-7 ~m sized, yellowish crystals; these inclusions and the cortical granules give O.
auripunctata an orange-yellow colour at low magnifications. Slow movement.
Adoral zone of membrane lIes about 28 % of body length. Frontal cirri slightly en-
larged, bases of all other cirri of about same size. Frontal, ventral, and marginal cirri in
life about 12 ~m long; 3-4 (mean = 3.8; n = 20), in life about 18 ~m long transverse cirri,
protrude distinctly beyond posterior end of cell. Postoral ventral cirri arranged about in a
line in median of cell. In 1 of 20 specimens a surplus cirrus between postoral and pre-
transverse ventral cirri. Marginal rows distinctly separate posteriorly, right one slightly
196 SYSTEMATIC SECTION

shortened anteriorly. Distance between left marginal cirri becomes wider toward rear
(Fig. 63e). Dorsal cilia in life about 2 J.lm long, probably arranged in 5 kineties (difficult
to discern because cortical granules stain with protargol!). Usually 3, sometimes (4 of20
specimens) 4-5 caudal cirri dorsally arranged in gap between marginal rows.

o c cur r e n c e and e colo g y: Likely confined to terrestrial habitats. Locus clas-

sicus is Tailem Bend near Adelaide, Australia, where BLATIERER & FOISSNER (1988) dis-
covered O. auripunctata in mosses of a natural pine forest, and also found it in litter and
roots under mosses of the same site. Also recorded from Costa Rica (FOisSNER 1997b).
Feeds on fungi, heterotrophic flagellates, and ciliates. Biomass of 106 cells 25 mg.

Oxytricha bimembranata SHIBUYA, 1929 (Fig. 64)

1929 Oxytricha bimembranata SIllBlNA, Proc. imp. Acad. Japan,S: ISS.

1930 Oxytricha bimembranata SIllBlNA - SlllBlNA, J. imp. agric. Exp. Stn Nishigahara, 1: 200 (Fig. 64).
1932 Oxytricha bimembranata SIllBlNA, 1929 - KAm., Tierwelt DtI., 25: 602.

T a x 0 nom y: The drawings ofSIDBUYA (1929, 1930) are more or less identical. I do
not agree with BORROR (1972a) that this terrestrial species is synonymous with the fresh-
water species O. fallax although it has about the same size and a similar body shape.
Identity with 0. proximata SIDBUYA, 1930 cannot be excluded
(Fig. 75h). The MATIS & DANISKovA (1972) population is classi-
fied as Sterkiella histriomuscorum. Detailed redescription nec-
essary.

M 0 r p hoi 0 g y and b i 0 log y: In life 150-200 J.lm

long, almost three times as long as broad. Elongate-elliptical,
margins slightly converging anteriorly and rather oblique on an-
terior left and posterior right borders. Contractile vacuole at
level of buccal vertex. Cortical granules absent. Adoral zone of
membranelles about 113 of body length. Frontal and transverse
cirri only slightly enlarged. Pretransverse ventral cirri displaced
somewhat anteriad. Five transverse cirri, the left 4 protrude
slightly beyond posterior end of cell. Marginal rows distinctly
separate posteriorly. Caudal cirri and dorsal cilia not
mentioned, probably inconspicuous and, respectively, short.

o c cur r e n c e: Probably confined to terrestrial habitats. Lo-

cus classicus is Tokyo, Japan, where SIDBUYA (1929) discov-

Fig. 64 Oxytricha bimembranata (from SlllBlNA 1930). Ventral view from

life, 150-200 Ilm.
 Oxytricha 197

ered 0. bimembranata in the soil of the experimental fann at Nishigahara. NIKOUUK &
GELTZER (1972; the illustration is from KAHL 1932) found it in soils of the USSR. BAM-
FORTH (1963; without description and illustration!), however, recorded it from freshwater
in south-eastern Louisiana, USA. Biomass of 106 individuals about 150 mg.

Oxytricha granulifera FOISSNER & ADAM, 1983 (Fig. 21a, 65a-z, 66a-g,
67a-g, Tables 3, 4, 15, 18)

1983 Oxytricha granulifera FOISSNER & ADAM, Zoo\. Ser., 12: 1 (Fig. 65a-z, 66a-g; 1 slide ofholotype speci-
merrs and 1 slide of paratype specimerrs are deposited in the OberOSterreichische Landesmuseum in
Linz, Upper Austria).
1989 Oxytricha granulifera FOISSNER and ADAM, 1983 - FOISSNER, Sber. Ost. Akad. Wiss., 196: 235.

Nom e n c I at u rea n d t a x 0 nom y: FOISSNER (1989) proposed O. granulifera as

the type species of Oxytricha (for details, see the relevant chapter in the genus section).
The subspecies, O. granulifera quadricirrata BLATTERER & FOISSNER, 1988, is classified
as separate species (Fig. 78a, b).

M 0 r p hoI 0 g y and b i 0 log y: In life 80-130 x 35-50 ~m. Outline rather con-
stant, broad to slender oval, posterior end usually broadly rounded, very rarely slightly ta-
pered, anterior portion usually distinctly narrowed. Right margin slightly convex to
slightly concave, left always distinctly convex (Fig. 65a, b). Macronuclear nodules in life
about 22 x 9 ~m, after protargol impregnation 10-20 x 7-9 ~m (mean = 14.4 x 7.7 ~m;
n = 25). Distance between macronuclear nodules after protargol impregnation 7-17 ~m
(mean = 10.7 ~m; n = 25). Constantly 2 (n = 25), in life about 5 x 4 ~m sized micronu-
clei. Contractile vacuole about in mid-body, during diastole with 2 short collecting
canals. Cytopyge on right posterior margin. Pellicle and cytoplasm colourless. Cortical
granules colourless to slightly yellowish, absent only along dorsal kineties and marginal
rows (Fig. 65b, 67a, b, f). Granules stain slightly red with methyl green-pyronin, are not
ejected, and render specimens a weak yellow-brown shade. Cells, especially in posterior
portion, packed with yellowish, 2-4 ~m sized crystals. Moderately rapid movement, of-
ten stands almost still, attaches closely to soil particles.
Adoral zone of membrane lIes about 1/3 of body length. Bases of largest mem-
branelles in life about 7 ~m broad. Buccal cavity moderately deep. Frontal, ventral, and
transverse cirri only very slightly larger than marginal cirri. Frontoventral cirrus nV2 lo-
cated on left, between migratory cirri VV3 and VV4 (Fig. 65f). Cirrus IV/3 displaced dis-
tinctly posteriad. Anterior pretransverse ventral cirrus (cirrus V/2) arranged more anteri-
orly than in 0. quadricirrata (compare Fig. 65f with Fig. 78a). Transverse cirri in life
about 22 ~m long, protrude distinctly beyond posterior end of cell. Number of frontal,
frontoventral, postoral ventral, and pretransverse ventral cirri constantly 3, 4,3, and 2, re-
spectively (n = 25). Marginal cirri in life about 12 ~m long, bases posteriorly distinctly
smaller than anteriorly. Caudal cirri in life about 15 !lm long, difficult to distinguish from
 \0
00
-

C/l
. ~
-<
C/l
~ --l
. tIl
~

... ~
~
--l
.......
" (J
" C/l
tIl
.... " - I
!\
I (J
--l
.......
()
, ...... q I) :1 I; ~
Q/)
• l;
•
e .. .,
~ ~g
C

Fig. 65a-g Oxytricha granulifera (from FOISSNER & ADAM 1983b. a-d, from life; e, wet silver impregnation; f, g, protargol impregnation). a) Ventral, dorsal, and lateral
view, a, c = 110 J.lm, b = 130 J.lm. The colourless, 0.5-1.0 J.lm sized, spherical cortical granules are arranged in longitudinal rows. d) Cyst, 32 J.lm. e) Part of silverline
system of ventral surface, bar = 18 J.lm. C, g) Ventral and dorsal infraciliature, f, g = 85 J.lm. Arrows in (f) mark pretransverse ventral cirri, arrowhead denotes frontoven-
tral cirrus IW2. CW = cyst wall, G = granules on cyst Wall, Ma = macronucleus, S = mucous layer; p 197.
 Oxytricha 199

D
o
. 01 ~
~
.~
1( §;;

.
<'q

" : fif:iS
\~
'~"
,
~ J'= '
j~ ,

~/IY"
io "
,
•
,
D
"
0 •, 0
,.
--oP ,.
", "
0

"
"
0
0
~ •, '"
• " "
"
0
"
0
0
~} • •
" ", " "
i-
"" "
t~

" " ""

" " ~ " 0

~ " .~
"
•" /I , ~~"', "
", 0 Q'
,
q a
c Q
0 p ()
a
•
.•..• . ""
0° alJ t)
0
••
.. h .•
.... DO
<>

k m

Fig. 65b-m Oxytricha granulifera (from FOiSSNER & ADAM 1983b. h-m, ventral [h, j, I] and dorsal [i, k, m]
infraciliature of morphogenetic stages after protargol impregnation). b, i) Very early stage, 90 Jim. j-m) Early
stages,j, k = 85 Jim, I, m = 80 Jim. OP = oral primordium; p 197.
 200 SYSTEMATIC SECTION

- -AM
"
"
" "
" " "
" "" "
"
" " "
" " '"
'"
'"
'"
'"
"
"
"
"
"
..

o q s
Fig. 650-5 Oxytricha granulifera (from FOISSNER & ADAM 1983b. Ventral en, p, r] and dorsal [0, q, s] infra-
ciliature of morphogenetic stages after protargol impregnation). 0,0) Early stages, 90 )lm. p-r) Middle stages.
AM = new adoral membranelles, PO = primordium of cirri and undulating membranes of opisthe; p 197.
 <> cl
\>0 0
~O<' "",,'

:;, ~ i ! //J co 0 '. ~

.."~~
.~ : ~
: '() ~ . ~
,l.;04:! V ~ !I
" 65\ : d / : ! .i~
0' i n~
CI
!',;
i:'~' ~
".." \".\Iir~f1 : _ / /';.:..1fj;I.7 P !I ;,.!! ,II';'~ I
~ ,! fU:/I~'
";!'r~ l\
"" h4\' 1/5\ .~,/.. ~ ," \:l'" .. ... ': X,I .-
, ~". ..
1,6.. \0.:if I§ : jty/ ~ .. I
" \~ \J I S "
\: , .. //rW \ I --
~. ~
;f;/. , '\ ';:. ..
\I ~/"" \ ~
, (!, : " f } I§. :
,.; i!!!i!!:!
Iii! ... l
. §'!' .. .,i,l} ~ .. I
,,!, " IiJli!!::: ..
e!ir l \ ~ \'<~\ ~ \ ,- !
!'[ 1'1 11 '1' :
.;/,.,.,1, #
"":'/~ .
-II')
... .... I :i!; ;fiJ/J» ~ ,
\! . :I 8
~
I
, 1\ \ II
" '~
!.~. I ,~ ! §:
s::.
"..
,r ~ ~-..", " l, \ i
\
.J
" ~\[f
t<lr
:
" \
\...
~
D •
.. Q t::J
... " "
~ ~ <>
.. .. ¢C;J " <>
Q

u
. <> v w

Fig.6St-w Oxytricha granulifera (from FOISSNER & ADAM 1983b. t-w, ventral [t, v] and dorsal [u, w] infraciliature of morphogenetic stages after protargol impregna-
tion), t, u) Middle stage, 85 11m, v, w) Late stage, 70 11m. 1-6 = primordia of the frontal-ventral-transverse cirri ofproter and opisthe respectively; p 197.

N
o
 202 SYSTEMATIC SECTION

,,
o

... o

.• 00
o
o0

..
..
IE
~
~
;
\
\
\•
....
•
a b c

Fig. 65x-z, 66a-c Oxytricha granulifera (from FOISSNER & ADAM 1983b. x-z, a-c, ventral [x, a] and dorsal
[y, Z, b, c) infraciliature of morphogenetic stages after protargol impregnation). x-z) Late stages, x, y = 85/lm,
z = 75 /lm. a-c) Very late stages, a, b = 85/lffi, c = 100 /lm. Parental cirri white, new cirri black; p 197.
 Oxytricha 203

."
II

.
to
to

...,.
.
.
to

,
." , ••
...
....
••
~

••• .., .'

, .
.
. f
• • 0

•• • 0
.,
"

~
,
:'~-~':''''.'-~~
. -if: :-.,'-
,
.
'0-::.'-,> .
~;~(":'::';'.....
.. ' '.' ' .
""
•. .•... •.'.'...•...•.." •.
'.'
'---'--'~' ' '.'-'

.•....:&:~': '--, •
,-. _._,'_.-_r...•..•.

-~~~~:/
"" : '

",
"
"", "•
~ ".'• q
f
•
••
f
0
~

00
0

0 0 d •• D' ~ •• 9

Fig.66d-g Oxytrichagranulifera (from FOISSNER & ADAM 1983. d-g, ventral [II, f] and dorsal [e, g] inftacilia-
ture of morphogenetic stages after protargol impregnation). d, e) Very late stage, 110 jlm. r, g) Opisthe, 55 jlm.
Parental cirri white, new cirri black; p 197.

cirri ofleft marginal row; usually 1 cirrus each on dorsal kineties 1,2, and 4. Dorsal cilia
in life about 3 ~m long. Invariably 5 (n = 25) dorsal kineties (Fig. 65g, 67g). Kinety 1 an-
teriorly distinctly and kinety 3 posteriorly slightly shortened. Kinety 4 begins somewhat
behind mid-body. Kinety 5 (= dorsomarginal row) terminates slightly in front of mid-
body.
 204 SYSTEMATIC SECTION

Figo 67a-c Oxytricha granulifera (original SEM micrographs, kindly supplied by W. FOISSNER). a) Ventral
view. The small, bright dots are extruded cortical granules. b) Cortical granules. c) Ventral view of a very early
morphogenetic stage showing longitudinal oral primordium (arrow; cpo Fig. 65j); p 197.
 Oxytricha 205

Cysts in life about 32 Ilm across, colourless to slightly yellow-brown. Cyst wall
smooth, thin, and rather densely covered with 1.4-2.8 Ilm sized, slightly orange coloured
granules embedded in a mucous layer surrounding the cyst (Fig. 65d); presumably, these
granules originate by fusion of the cortical granules of the non-encysted specimens. Cyto-
plasm densely granulated, with some about 5 Ilm sized, shining inclusions. Macronucleus
irregularly spherical.
Morphogenesis of cell division is described minutely by FOISSNER & ADAM (1983b).
For details, see Figures 65h-z, 66a-g, 67c, and Tables 3, 4. The anlagen formation pro-
ceeds identically to in Allotricha antarctica (Table 4), strongly indicating a close rela-
tionship of Oxytricha and Allotricha (Fig. 25a, b).
Some biochemical and genetic data on O. granu/ifera are provided by GREENWOOD et
al. (1991a, b), PALMEDO & AMMERMANN (1997a, b), SCHLEGEL (1985, 1991), and SCHLEGEL
& STEINBROCK (1986; see also SCHLEGEL et al. 1996). The small subunit rRNA sequence is
1758 bases long, the G + C content is 46 % (LEIPE et al. 1994, SCHLEGEL et al. 1991).

o c cur r e n c e and e colo g y: Common; very likely confmed to terrestrial habi-

tats. Locus classicus is the soil of a deciduous forest (Fagus sy/vatica, Quercus robur,
Galium odoratum, Convallaria maja/is), about 1 Ian north of the village of Baumgarten
in Lower Austria (FOISSNER & ADAM 1983b). A detailed description of this site and aute-
cological data are given by FOISSNER et al. (1985, "Profil T'). They found O. granulifera
also in the soil of wheat fields, xerothermic uncultivated grassland, and humid lowland
sites. Detailed autecological data of an alpine population are given in FOISSNER & PEER
(1985). Further records: soil of a paddy field in Japan (W. FOISSNER, pers. comm.); bark
of Acacia trees in Costa Rica (FOIssNER 1994c, 1997b); Peru (FOISSNER 1997b); various
soil samples from Australia (BLATIERER & FOISSNER 1988); soil from Marion Island,
South Atlantic Ocean (FOISSNER 1996).
Feeds on bacteria, fungi, and heterotrophic flagellates. Can be cultured in lettuce me-
dium diluted 1: 1 with table water (Bau de Volvic, France) and with some wheat grains to

Table 18 Metal accumulation and distribution in cadmium-treated cells of Oxytricha granu/i/era, Sty/onychia
/emnae, and S. notophora (Ilg g" dry weight; from PiCCINNl et al. 1992). Sty/onychia notophora was obtained
from G.R. SAPRA; this population is classified as S. pustu/ata in this book)

Species Cd Zn
Supernatant Pellet Supernatant Pellet
Treatment Total (%total (% total Total (%total (%total
11M of Cd homogenate metal) metal) homogenate metal) metal)
O. granu/ifera day 1 1400 6 94
day 3 1700 3 97
O. granulifera 50 day 1 2900 40 60 2500 7 93
day 3 5500 10 90 3000 20 80
S. /emnae 2.5 day I 185 25 75 2765 30 70
day 2 635 6 94 2625 10 90
S. notophora IS day I 1780 12 88 1790 5 95
206 SYSTEMATIC SECTION
 Oxytricha 207

support bacterial growth. Biomass of 106 individuals about 72 mg (FOISSNER 1987a). Oxy-
tricha granulifera is much more sensitive to copper than to cadmium (!RATO et al. 1993).
50 flM cadmium (as CdChHzO) did not damage the cells, whereas 20 flM copper killed
them in one day and 10 flM in three days. One-day exposure to cadmium induced an ac-
cumulation of 3000 flg Cd g') of dry weight, 30 % of which was present in the cytosol
(!RATO et al. 1991, Table 18). Detoxification mechanisms, see !RATO et al. (1995), PIC·
CINNI (1992), and PICCINNI et al. (1992).

Oxytricha islandica BERGER & FOISSNER, 1989 (Fig. 68a-f, Table 15)

1989 Oxytricha islandica BERGER & FOISSNER, Bull. Br. Mus. nat. Hist., 55: 37 (I slide of holotype
specimens, reference number 1988:2:1:15, and 1 slide of paratype specimens, reference number
1988:2:1:16, are deposited in the British Museum of Natural History in London).

T a x 0 nom y: The 4 dorsal kineties are reminiscent of Urosomoida. Indeed, at superfi-

ciallive observation it can be easily confused with U dorsiincisura, which has the same
size and nuclear apparatus. However, this species has cortical granules and only 2-3
transverse cirri (including the pretransverse ventral cirri). In 0. islandica the frontoven-
tral cirri are arranged as in the type species of Oxytricha, O. granulifera, and 2 pretrans-
verse ventral and 5 transverse cirri are invariably present. Confusion with other quadrinu-
cleate oxytrichids, for example, Cyrtohymena quadrinucleata or Sterkiella cavicoia, is
unlikely.

M 0 r p hoI 0 g y and b i 0 log y: In life 100-140 x 35-45 flm. Long elliptical, right
margin sometimes straight, both ends rounded. Slightly contractile. Macronuclear nod-
ules in life about 13 x 10 !lIn, after protargol impregnation 9-17 x 7-8 !lm (mean = 13.1
x 7.4 flm; n = 10), arranged in line almost in median of cell (Fig. 68a, f). 1 or 2 (mean =
1.8; n = 10) micronuclei, in life about 4 flm across. Contractile vacuole somewhat in front
of mid-body, during diastole without distinct collecting canals. Cortical granules lacking.
Cytoplasm colourless, posterior portion packed with cytoplasmic crystals and food vacu-
oles. Rapid movement.
Adoral zone of membranelles about 31 % of body length. Bases of largest mem-
branelles in life about 7 flm wide. Buccal area flat, undulating membranes nearly straight.
Typical Oxytricha cirral pattern with constantly (n = 10) 3 frontal cirri, 1 buccal cirrus, 4
frontoventral cirri, 3 postoral ventral cirri, 2 pretransverse ventral cirri, and 5 transverse
cirri. Frontal and marginal cirri in life about 15 flm, transverse cirri about 22 flm long.
Usually 4 (as in Urosomoida!), sometimes (1 of 10 specimens) 5 dorsal kineties. Kineties

~ Fig. 67d-g Oxytricha granulifera (original SEM micrographs, kindly supplied by W. FOISSNER). d, g) Ventral
and dorsal view of anterior cell portion showing frontoventral cirrus lIll2 (arrow) and dorsal bristles. e} Left
lateral view. t) Detail showing some marginal cirri, and ejected cortical granules arranged in short, longitudinal
rows; p 197.
 tv
oOQ

.....
!IIo...
-: ~ /'- ) ,-
\ I
....
. - • t. ~
/,/''- \'
,J
,i
• \' ,-
f. ,J
•,, , ,-
, l" " ,- 'f
,, f ,- T
,- \/1
, , {' ,-
T -<
\/1
,, • f. r >-j

,
,
-..... f,
... ,..
'~-.- t~ (')
\/1
'T ~
\ r j
\' '\ T @
, \
T'
'" \
"• ~ i
"-.
' ./ f

Fig. 68a-£ Oxytricha islandica (from BERGER & FOISSNER 1989a &-d, from life; e, f, protargol impregnation). a, b) Ventral and lateral view, a = 120 l.un, b = 110 llm. C,
d) Shape variants in ventral view, c, d = 130 llm. e, 1) Ventral and dorsal infraciliature, e, f= 102 llm; p 207.
 Oxytricha 209

1-3 distinctly bent, kinety 1 slightly shortened anteriorly, kinety 4 tenninates roughly in
mid-body. Dorsal cilia in life about 3 ~m long. Caudal cirri on dorsal kineties 1-3.

o c cur r e n c e and e colo g y: Very probably confined to terrestrial habitats. Lo-

cus classicus is the soil of a pasture with Deschilmpsia caespitosa and moss (PH 5.8),
about 100 m from the shore on the Vatnsney peninsula, "Hritserkur", Iceland (BERGER &
ForSSNER 1989a). FOISSNER et al. (1990) found O. islandica in the soil ofa meadow in Ai-
gen in Styria, Austria. Feeds on cysts of amoebas. Biomass of 106 individuals about
50 mg.

Oxytricha lanceolata SHIBUYA, 1930 (Fig. 69a-l, 70a-c, Tables 3, 15)

1930 Oxytricha lanceolata SIfiBUYA, J. imp. agric. Exp. Stn Nishigahara, 1: 210,214 (Fig. 69a).
1935 Oxytricha lanceolata SIfiBUYA, 1930 - KAHL, TierweltDtl., 30: 841.
1987 Oxytricha lanceolata SIfiBUYA, 1930 - BERGER & FOISSNER, Zool. Jb., 114: 219 (Fig. 69g-1; authorita-
tive redescription; 1 slide of neotype specimens is deposited in the Oberosterreichische Landesmuseum
in Linz, Upper Austria).
1989 Oxytricha lanceolala SIfiBUYA, 1930 - BERGER & FOISSNER, Bull. Br. Mus. nat. Hist, 55: 39 (Fig. 69b--f;
1 slide of protargol-impregnated specimens, reference number: 1988:2: 1: 17, is deposited in the British
Museum of Natural History in London).
1996 Oxytricha lanceolata SHIBUYA, 1930 - FOISSNER, Acta Protozool., 35: 116 (Fig. 70a-c).

Nom e n c I a t u rea n d t a x 0 nom y: KAlIL (1932, 1935) divided Oxytricha into

subgenera; the correct name in his revision is thus Oxytricha (Oxytricha) lanceolata. The
habitat and the morphology of the populations described by BERGER & ForSSNER (1987a,
1989a) correspond very well with the original description, which is, however, based on
living observations only. The Antarctic population described by ForSSNER (1996b) agrees
very well with the Eurasian populations (BERGER & ForSSNER 1989a, SHIBUYA 1930); only
the frontoventral cirrus IIII2 and pretransverse cirrus V12 are somewhat differently ar-
ranged. Both cirri are displaced slightly posteriad in the Antarctic population (cp. Fig.
69c, j, 1 with 70c). Very likely, Oxytricha lanceolata is confined to terrestrial habitats.
Hence it may not be synonymised with the coloured freshwater species, O. aeruginosa, as
suggested by BORROR (1972a).

M 0 r p hoI 0 g y and b i 0 log y: In life about 90-110 x 30-50 Ilm. Outline long
elliptical or lanceolate, equally rounded at both ends. Anterior portion sometimes slightly
narrowed (Fig. 69b, f, h). Distinct dorsal furrow anteriorly. Macronuclear nodules in life
21 x 10 Ilm, after protargol impregnation 13-21 x 7-10 Ilm (mean = 15.3 x 7.8 Ilm; n =
12). 1-3, usually 2, in life about 3 Ilm sized spherical micronuclei. Contractile vacuole
slightly in front of mid-body, during diastole with 2 inconspicuous collecting canals. Cy-
toplasm colourless, opaque by small «1 Ilm), greasy granules (no cortical granules!);
posterior portion of cell packed with 3-8 Ilm sized, colourless globules and 2-7 Ilm sized
crystals. Very rapid movement, sliding hastily to and fro.
210 SYSTEMATIC SECTION

': ';
\ '\
! '.
': '.
\ "-
't \
{.." '.
k \'\()
!: \
~/~~
t \' '. \
,
,
l' "-:.,..... \
{'
I \ ....._- '\

.-, 1" '

\,
, \
, l' \
,
, 1 \
,
, r
1
l' '
.-
L
\'
\t~ ".
( \
~ \
'. "

Fig. 69a-f Oxytricha lanceolata (a, from SHIBUYA 1930; b-f, from BERGER & FOISSNER 1989a a, b, e, f, from
life; c, d, protargol impregnation). a, b) Ventral views, a = 90-100 11m, b = 110 11m. C, d) Ventral and dorsal
infraciliature, c, d = 105 11m. e, f) Lateral and dorsal view, e, f= 130 11m; p 209.

Adoral zone of membranelles about 30 % of body length, cilia of distal mem-

branelles about 13 /lm long. Undulating membranes almost straight. Frontal cirri slightly
enlarged, in life about 16 /lm long. Usually 4, rarely 3 (Fig. 69j) or 5 frontoventral cirri.
All postoral ventral cirri immediately behind buccal vertex. Usually 5, rarely (l of 12
specimens) 6 enlarged transverse cirri, slightly cup-shaped, in life about 28 /lm long.
Marginal cirri 13-15 /lm long; left row J-shaped, terminates at posterior end of cel~ right
one terminates at level of posterior transverse cirrus. Dorsal cilia in life 3-4 /lm long.
Constantly (n = 12) 4 dorsal kineties, kinety 1 slightly shortened anteriorly, kinety 4 ter-
minates in mid-body. Usually 1 caudal cirrus each on dorsal kineties 1-3, sometimes 2 on
each of these kineties; caudal cirri not conspicuously elongated but rather motile.
Antarctic population (Fig. 70a-c) in life 80-120 /lm long; invariably (n = 11) 2 mi-
cronuclei; 4-7 (mean = 4.6; n = 11) frontoventral cirri; usually 2, rarely 3 (mean = 2.1; n
= II) pretransverse ventral cirri; usually 5, rarely 6 (mean = 5.2; n = II) transverse cirri;
usually 4, rarely 5 (mean = 4.1; n = 11) dorsal kineties.
Stomatogenesis begins with the formation of a long and narrow oral primordium
which extep.ds from left transverse cirrus to posterior postoral ventral cirrus (Fig. 691).
 "'-":."
""
". \' \

,/
/
'j
~ 'f~~'j
r--, ";;0",,.,
i.:.
I, ':aL'.,_1
,.,.
""
".- r.
···il ''"" ,,.
L/·· .,'" ,.,. ....... ,.''"" ''""
": .. : '-.
,.,. 1\"~"'; f) '.
:,,' "'~
.... " ~~
,.''"" ,.,. §:
I::l
,.,.'" 1 ''"'""
'"
1:\' tI~·" . ~.\
., j
\
""'" ''""
'" #
l' "'",-. \ , 1
i .,'"
" .,'" \
\' ". !J } .-
.- ;
"
""
#
• .. \ _, ~.'-.... If1
.
.}
" ; • "'- '" •
# ~ -" ............. ' ~
~
t. ... ~ -~, f/ k '" -
~1i.'
-.··.•··
Fig. 69g-1 OxytricluJ lanceolata (from BERGER & FOISSNER 1987a. g-i, from Iife;j-I, protargol impregnation). g-i) Ventral, dorsal, and right lateral view, g = 100 Jlm, h,
i = 110 Jlm. j. k) Ventral and dorsal infraciliature of a specimen with only 3 frontoventral cirri and only 2 caudal cirri, j, k = 90 Jlm. 1) Ventral infraciliature of a very
early morphogenetic stage, 85 Jlm. Arrowheads mark 3 caudal cirri; p 209.
tv

--
 212 SYSTEMATIC SECTION

\
\ I /

\ \ I /

\:
\...
('
I
./
/
~ :

I
/

I
/

\ /

Itr
/

r
\
\'
/
:/

'7
'l-.\

~ !
t ) 7
'" ) T
#
\' 1 IT
••• ••
• # •
~ I /
\~ ),~
a • ••• b ~c
Fig. 70a~ Oxylricha Ianceo/ala (from FOISSNER 1996b. a, from life; b, c, protargol impregnation). a) Ventral
view, 97 11m. b, c) Ventral and dorsal infraciliature, 107 11m. Arrow and arrowhead in (b) marks frontoventral
cirrus III12 and pretransverse ventral cirrus V/2, respectively, which are slightly displaced posteriad as com-
pared to the Eurasian populations (cp. with Fig. 69c,j, I). Arrow in (c) marks a short, fifth dorsal kinety found
in few specimens; p 209.

o c cur r e n c e and e colo g y: Common in terrestrial habitats. Locus classicus is

the soil from a field at Miyako-mura near Chiba, Japan (SHIBUYA 1930). BERGER & FOISS-
NER (1987a) found o. lanceolala in the upper soil layer (0-5 cm) of a conventionally
farmed pasture near the village of Seekirchen, Austria. Autecological data of this popula-
tion are given by FOISSNER et al. (1987a, Versuchsflache F). BERGER & FOISSNER (1989a)
found it in a reddish soil ofa heath with grass, about 1800 m above sea-level on the Pico
de Arieiro, Madeira, Portugal. FOISSNER (1996b) found 0. lanceolala in a bulked sample
 Oxytricha 213

(wet Bryum moss from edge ofmeltstream; algae from meltstream; Bryum moss from dry
area near stream) from Garwood Valley, South Victoria Land. Further records: upper soil
layer of a flood plain dominated by Alnetum incanae (PH 7.2) near the Rotmoos in the
Fuscher Tal, Austria (FOISSNER 1987b); soil from the islands Corsica, France, and Born-
holm, Denmark (W. FOISSNER, pers. comm.); soils in Belgium (CHARDEZ 1967, CHARDEZ &
KRIzEu 1970); subtropical soils in China (SHEN et al. 1992); Brazil and Tasmania (FOIss-
NER 1997b); soil from Gough and Marion island, South Atlantic Ocean (FOISSNER 1996c).
Feeds on bacteria, fungi, globular green algae, and heterotrophic flagellates. Biomass
of 106 individuals about 70 mg (FOISSNER 1987a).

Oxytricha /ongigranu/osa BERGER & FOISSNER, 1989 (Fig. 71a-h,

Tables 3, 15)

1988 Oxytricha longigranulosa BERGER & FOISSNER, 1989 - BLAlTERER & FOISSNER, Stapfia, 17: 65 (I slide
of protargol-impregnated specimens is deposited in the OberOsterreichische Landesmuseum in Linz,
Upper Austria).
1989 Oxytricha longigranulosa BERGER & FOISSNER, Bull. Br. Mus. nat. Hist, 55: 39 (Fig. 71a-c, e-h; 1 slide
of holotype specimens, reference number 1988:2:1:18, and 1 slide of paratype specimens, reference
number 1988:2:1:19, are deposited in the British Museum of Natural History in London).

M 0 r ph 0 log y and b i 0 log y: The BLATIERER & FOISSNER (1988) population is

somewhat smaller than the type population; however, the important characters agree
rather well (Table 15). In life about 135 x 55 ~m, sometimes only 80 x 35 ~m. Elliptical,
right margin straight, left one distinctly vaulted at level of contractile vacuole. Both ends
rounded. Macronuclear nodules in life 18-21 x 10-11 ~m, after protargol impregnation
12-17 x 7-9 ~m (mean = 14.8 x 7.6 ~m; n = 13), according to BLATIERER & FOISSNER
(1988) 9-11 x 5-7 ~m (n = 10, protargol impregnation). 1-2, according to BLATIERER &
FOISSNER (1988) 2-6, after protargol impregnation 2-3 ~m sized, almost spherical micro-
nuclei; if only one micronucleus is present, it is not located between the 2 macronuclear
nodules as, for example, in Oxytricha setigera. Contractile vacuole in front of mid-body,
during diastole without collecting canals (possible rather inconspicuous). Cortical gran-
ules colourless, anterior with an inconspicuous knob, stain red with methyl
green-pyronin, but are not ejected (Fig. 71b-d). Cytoplasm colourless, with many about
5 ~m sized greasily shining globules and many food vacuoles. Gliding movement, some-
times nearly jumping.
Adoral zone of membranelIes about 1/3 of body length. Bases of largest mem-
branelles in life about 7 ~m broad. Buccal area flat, undulating membranes almost
straight. Bases of all cirri of nearly same size. Typical Oxytricha cirral pattern (Fig. 71f).
Type material constantly with 5 (n = 13) transverse cirri, population of BLATIERER &
FOISSNER (1988) sometimes (3 of 10 specimens) with only 3 transverse cirri. Marginal and
transverse cirri in life 12-15 ~m and 18-22 ~m long, respectively. Constantly 6 dorsal
kineties. Kineties 1 and 2 of about body length, kinety 3 with a bend to the right at poste-
214 SYSTEMATIC SECTION

.
~ 0 0
0" 0 0 d
o 0 • 0
oo 0 •
o o0 0 0

00°"'0 0
0
o a:

8:
°0 0 0

% ~ :
o() 0 g0" II

71a

"
....
.. #

..

, , ,
, , .
, ..
,
, .
~ ,
, n'!.
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., ,
(

,~
e ..
#
.
•
•
:•
•• t
•
• ••
••
•
h
 Oxytricha 215

rior end and kinety 4 distinctly bent to the left anteriorly so that the connection (before
fragmentation) of these 2 kineties is still recognisable (Fig. 71g, dotted line). Kinety 5
half of body length, kinety 6 consists of about 4 basal body pairs only. Dorsal cilia in life
3-4 pm long. Caudal cirri on kineties 1,2, and 4.
Stomatogenesis commences with the proliferation of basal bodies near the transverse
cirri and near the postoral ventral cirri (Fig. 7lh). Dorsal morphogenesis very likely in
Oxytricha pattern (with 2 dorsomarginal kineties).

o c cur r e n c e and e col 0 g y: Rather certainly confmed to terrestrial habitats.

Locus classicus is a brown soil of a deciduous forest with fragments of leaves and roots
(PH 3.5) from Mt Kura-Take, Amakusa Kumamoto Prefecture, Japan (BERGER & FOISS-
NER 1989a). BLATIERER & FOISSNER (1988) found O. longigranulosa in barks overgrown
with lichens and mosses from a secondary pine forest (Scots Pine, Pinus silvestris) on the
area of a tropical forest between Cairns and Innisfail, Australia. FOISSNER (1997b and
pers. comm.) found it in soil samples from Kenya, Panama, and Brazil, FOISSNER (1996c)
in soil from Marion Island (South Atlantic Ocean). BONKOWSKI (1996) observed it in leaf
litter of a beech forest in Germany, indicating that Oxytricha longigranulosa is a cosmo-
politan. Feeds on bacteria, fungal spores, heterotrophic flagellates, naked amoebas, and
ciliates. Biomass of 106 individuals about 100 mg (type material).

Oxytricha naup/ia BERGER & FOISSNER, 1987 (Fig. 72a-d, Table 15)

1987 Oxytricha naup/ia BERGER & FOISSNER, Zool. Jb. Syst., 114: 219 (l slide ofholotype specimens and 1
slide of paratype specimens are deposited in the OberOsterreichische Landesmuseum in Linz, Upper
Austria).

T a x 0 nom y: Differs from other small to medium sized Oxytricha species mainly in
the body shape, the ventral cirral pattern, the number and arrangement of the dorsal kine-
ties, and the habitat. Identification, however, has to be proved on protargol-impregnated
specimens.

M 0 r p hoi 0 g y and b i 0 log y: In life 80-100 x 40 J-lm. Elliptical to almost paral-

lel sided, anterior portion converging. Macronuclear nodules in life about 21 x 10 J-lm, af-
ter protargol impregnation 10-14 x 6-9 J-lm (mean = 11.1 x 7.8 J-lm; n = 10). Distance
between macronuclear nodules after protargol impregnation 3-9 J-lm (mean = 5.4 J-lm; n =

~ Fig. 71a-h Oxytricha [ongigranu[osa (a--c, e-h, from BERGER & FOISSNER 1989a; d, original, kindly supplied
by W. FOISSNER. a--c, e, from life; d, methyl green-pyronin staining; f-h, protargol impregnation). a) Ventral
view, 130 11m. b) Colourless, rod-shaped extrusomes (about 2.0 x 0.5 11m) in lateral view. c, d) Dorsal and
ventral view. Extrusomes (cortical granules) arranged in short rows. Arrow in (d) marks an extrusome in lateral
view, c = 150 11m. e) Lateral view. C, g) Ventral and dorsal infraciliature, f, g = 100 11m. Dotted line in (g) con-
nects posterior end of dorsal kinety 3 and anterior end of dorsal kinety 4. h) Ventral infraciliature of an early
morphogenetic stage, 100 11m; p 213.
216 SYSTEMATIC SECTION

·.. f ~
:.
... t }
f
~
~
~

,
..
.)
..
..
...
..
~ ~
.
....
..

....
.. ..
.. ...
..
.. ..
..
.. •
.. • •• ..
.... ••
• .
" • c d

Fig.72a-d Oxytricha naup/ia (from BERGER & FOISSNER 1987a a, b, from life; c, d, protargol impregnation).
a, b) Ventral and dorsal view, a = llO J,lm, b = 125 J,lm. c, d) Ventral and dorsal infi"aciliature, c, d = 70 J,lm;
p 215.

10). 2-3 in life 2-3 !lm sized spherical micronuclei. Contractile vacuole about in mid-
body. Cortical granules lacking. Cytoplasm colourless, packed with 2-5!lm sized
crystals, shining globules, and food vacuoles. Rapid movement.
Adoral zone of membranelles about 1/3 of body length, bases of largest mem-
branelles in life about 7 !lm broad. Buccal area flat and very narrow. Bases of frontal cirri
only slightly. enlarged. Right frontal cirrus between distal end of adoral zone of mem-
branelles and anterior end of right marginal row. Invariably 4 frontoventral cirri which
are, however, not arranged in typical Oxytricha pattern (Fig. 72c). Constantly 5 trans-
verse cirri displaced somewhat anteriad so that they protrude only slightly beyond poste-
rior end of cell; bases only slightly enlarged, arranged in an almost longitudinally orien-
tated hook. All cirri in life about 20 !lm long. Dorsal cilia in life 2 !lm long, constantly ar-
ranged in 5 rows (n = 10). Kineties 1-4 unshortened, kinety 5 terminates at about level of
buccal vertex (Fig. 72d). Usually 3, rarely 4 very motile caudal cirri.

o c cur r e n c e and e colo g y: Probably confmed to terrestrial habitats. Locus

classicus is the upper soil layer (0-5 cm) of a salt soil with rushes, about 50 m from the
 Oxytricha 217

sea in the Nauplia Bay, Peloponnesus, Greece (BERGER & FOISSNER 1987a). Also recorded
from Costa Rica (FOISSNER 1997b). Feeds on fungal spores and flagellates, like Aniso-
nema sp. and Euglena viridis. Biomass of 106 cells about 26 mg (FOISSNER 1987a).

Oxytricha opisthomuscorum FOISSNER, BLATIERER, BERGER & KOHMANN,

1991 (Fig. 73a, b, 244a-j, Table 52, Addendum 1 on p 925)

1932 Opisthotricha muscorum KAHL, Tierwelt Dtl., 25: 608 (Fig. 73a).
1980 Oxytricha muscorum (KAlIL, 1930-35) - FOISSNER, Ber. Nat.-Med. Ver. Salzburg, 5: 110 (Fig. 73b).
1991 Oxytricha opisthomuscorum FOISSNER, BLATIERER, BERGER & KOllMANN, Infonnationsberichte des
Bayer. Landesamtes filr Wasserwirtschaft, 1191: 311.
1997 Oxytricha opisthomuscorum FOISSNER and others, 1991- PE1z & FOISSNER, Polar Record, 33: 318 (Fig.
244a-j; description see p 925).

Nom e n c I at u rea n d t a x 0 nom y: KAHL (1932) classified Opisthotricha as sub-

genus of Oxytricha; the correct name in the original description is thus Oxytricha (Opistho-
tricha) muscorum, which is a primary homonym of 3 other Oxytricha muscorum described
by KAHL (1932; for details, see Cyrtohymena muscorum). The nomenclatural rectification
was done by FOISSNER et al. (1991). Both KAHL (1932) and FOISSNER (1980a) described this
species after live specimens; this explains some minor differences in the cirral pattern. The
FOISSNER (1980a) population is reminiscent of O. setigera, which is, however, smaller and
which has fewer marginal cirri. FERNANDEZ-LEBORANS (1985, p 376) described the fine struc-
ture of the undulating membranes of
"Oxytricha muscorum". The micrograph
(his Fig. 13), however, shows an in-
creased number of cirri on the frontal
area so that the identification is doubtful;
according to this paper the anterior por-
tion of the paroral is composed of 6 rows
with 46-48 basal bodies, whereas the
posterior portion consists of 2 adjoining
and parallel rows of 20-22 basal bodies
each. Detailed redescribed by PETZ &
FOISSNER (1997a, see p 925).

Morphology and biology:

After KAHL (1932) in life 80-100/lm,

Fig. 73a, b Oxytricha opisthomuscorum (a, from

KAlIL 1932; b, from FOISSNER 1980a). Ventral
views from life, a = 90 flm, b = 75 flm. Note sin-
gle micronucleus between macronuclear nodules. b
218 SYSTEMATIC SECTION

after FOISSNER (1980a) in life 65-80 ~m long. About 3 times as long as broad. According
to KAHL (1932), right margin rather more straight than convex, left anteriorly slanted.
Rather contractile (FOISSNER 1980a). Macronuclear nodules arranged in median or even
slightly right of it Contractile vacuole about in mid-body. Hasty, jerky movement.
Adoral zone of membranelles about 38 % of body length. Peristomial lip anteriorly
distinctly bent to the left, which is reminiscent of Cyrtohymena species. Arrangement of
frontoventral cirri not as in many other Oxytricha species, especially posterior-most cir-
rus distinctly displaced posteriad (Fig. 73a and 244c, large arrow). Five transverse cirri
which protrude rather far beyond posterior end of cell. Dorsal cilia stiff, after KAHL
(1932) in life 10-15 ~m, according to FOISSNER (l980a) 9-12 ~m long, spread in resting
animals, during swimming, however, trailed behind like fringes. Three long (about 113 of
body length, according to KAHL 1932) and soft caudal cirri, energetically motile.

o c cur r e n c e and e colo g y: Locus classicus are mosses on walls and rocks
near Hamburg, Germany (KAHL 1932). FOISSNER (1980a, c) found O. opisthomuscorum
on irrigated rocks and in pasture pools in the Glockner area, Austrian Alps. Records not
substantiated by illustrations: in road pools near Bratislava, Czechoslovakia, during
autumn at 10-21 °C, pH 5.5--6.6, and 7.3-12.5 mg 1'\ O2 (MATIS 1975); submerged, wet,
moist, and dry mosses in the area of Slovensky raj, Czechoslovakia (TIRJAKovA & MATIS
1987a); in two thermal lakes in Czechoslovakia at 7.0--40.5 °C and pH 5.0-7.4 (MATIS &
STRAKOVA-S1RIESKOvA 1991); Turiec River in Slovakia (TIRJAKovA 1993); frequent in
mosses and clear water from a spring in Italy (STEllA 1947); freshwater habitats in the Ti-
betan Plateau (WANG 1977); soil near Manaus, Brazil (FOisSNER 1997b); various soil sam-
ples (damp soil; wet mosses) from South Victoria Land and grass sward from Deschamp-
sia antarctica patch (PH 4.4) of a sheltered north-facing slope on Signy Island, Antarc-
tica (FOISSNER 1996b). Feeds on bacteria, green algae, and flagellates. Biomass of 106 in-
dividuals about 20 mg (population by KAHL 1932).

Oxytricha otto wi FOISSNER, 1996 (Fig. 73c-h, Table 15)

1996 Oxytricha ottowi FOISSNER, BioI. Fertil. Soils, 23: 289 (two type slides have been deposited in the
OberOsterreichische Landesmuseum in Linz, Upper Austria).

M 0 r p hoI 0 g y and b i 0 log y: In life about 85-110 x 25-35 ~m. Outline

roughly elliptical, both ends broadly rounded, dorsoventrally flattened up to 2:1. Very
flexible. Invariably (n = 19) 8 globular to ellipsoidal macronuclear nodules slightly left of
median, after protargol impregnation 6-9 x 4-7 ~m (mean = 7.9 x 6.0 ~m; n = 14).0-5
(mean = 3.2; n = 19) globular micronuclei, after protargol impregnation 2-3 ~m across.
Contractile vacuole slightly in front of mid-body, without conspicuous collecting canals.
Cortical granules elongate, about 2.0 x 0.5 ~m, colourless, stain deep-blue with methyl
green-pyronin and light-brown with protargol, arranged in many loose rows (Fig. 73d, f).
 .s \ ,1,/ ,/
\ ) /
,.
~
'/~ 8'::::§ ::; \' ) '/
p.,~ s:: /' I ,

l\ ~ '-!./'
~, \ .. ') / ./.\
,/ \ <i.•. r,:. / ./
• ~s §" "" \ • . •,. I • y
'-I.'
~' ~'/
:"),Irt~, .-:\
/' ~.~/ r--
~:~
... /f .. ':.Ii ' ;r\~/ 1
.. , '}
; ~'\~I/
,,
..-,
I \" •... :,.,
)If::~,
__
'" '" \ \'{~:, ~ I ~:::;-
'," '" ~\. ", J ..-- >-" §:
t:l
I I /1
'" ..
'.." 1 ' . \/y/ ~
;- '-;-\' ,. \ 'I~, '. \: I

~j .. ~
.... . . . . ,\. \I) \/;, f
..
. ~~ · \ tj
. ... ' .
.., ./I\'~, (Ii
' "..... ,
..
~~
• •• .
r~
A~.~.~ ••
~ '"
.;~
~.j" • /.""\' . lV'
\.,
e •• g
f
~~~..J 'h
Fig. 73c-h Oxytricha ottowi (from FOISSNER 1996c. c-t: from life; g, h, protargol impregnation). c) Ventral view, 97 J.1m. d) Dorsal view showing rows of cortical gran-
ules. e) Cytoplasmic crystals. 1) Lateral optical section showing that cortical granules (arrowhead) are rod-shaped. g, b) Ventral and dorsal infraciliature and nuclear ap- tv
.....
paratus, 90 J.1m. Arrowhead marks posterior end of dorsal kinety S. The main species character is the macronucleus which invariably consists of eight nodules; p 218. \0
220 SYSTEMATIC SECTION

Cytoplasm colourless, with few to many 3-4 11m sized crystals, some 2-4 11m sized fat
globules, and food vacuoles (Fig. 73c, e). Movement moderately rapid, without peculiari-
ties.
Adoral zone of membranelles about 34 % of body length. Buccal cavity compara-
tively large and deep, right third covered by hyaline lip. Paroral and endoral curved and
tightly spaced, that is, in parallel and/or intersecting optically only in anterior third, both
composed of dikinetids arranged in zigiag. Pharyngeal fibres conspicuous. Frontal and
marginal cirri about 15 !lm, transverse cirri about 20 !lm long. Frontal-ventral-transverse
cirri (except pretransverse ventral cirri) slightly enlarged as compared to marginal cirri.
Invariably (n = 14) 3 frontal cirri, 4 frontoventral cirri, and 1 buccal cirrus; usually 3,
rarely 4, postoral ventral cirri; usually 2, rarely only 1, pretransverse ventral cirri; usually
5, rarely 4, transverse cirri. Marginal rows widely open at posterior end, gap occupied by
caudal cirri right of cell median. Dorsal cilia about 3 !lm long, invariably (n = 14) ar-
ranged in 6 dorsal kineties. Constantly (n = 14) 3 caudal cirri on kineties 1,2, and 4 (Fig.
73h), caudal cirri not conspicuously elongated.

o c cur r e n c e and e colo g y: Locus classicus is the Transvaal Bay, Gough Is-
land, South Atlantic Ocean, where FOlSSNER (l996c) discovered it in fern bush (Histiop-
teris incisa) peat litter, mixed with some soil and moss (PH 4.4; 30 m above sea level).
Feeds on bacteria, heterotrophic flagellates, and coccal cyanobacteria.

Oxytricha procera KAHL, 1932 (Fig. 74)

1932 Opisthotricha procera KAliL, Tierwelt Dtl., 25: 608.

Nom e n c I a t u rea n d t a x 0 nom y: KAHL (1932) classified Opisthotricha as

subgenus of Oxytricha; the correct name in the original description is thus Oxytricha
(Opisthotricha) procera. I do not agree with BORROR (1972a) that this spe-
cies is identical with the limnetic O. simi/is which has only 2 caudal cirri.
However, the validity of O. procera has to be confirmed by a detailed re-
description. Opisthotricha procera sensu NAlDu (1965; Fig. 224k) is insuf-
ficient redescribed.

M 0 r p hoi 0 g y and b i 0 log y: In life about 100-120 !lm long.

Outline slightly spindle-shaped. Contractile vacuole distinctly in front of
mid-body where the body is broadest. Cytoplasm packed with dark gran-
ules, especially in posterior portion. Adoral zone of membrane lies about
28 % of body length. Number and arrangement of frontal, ventral, and
transverse cirri probably genus-specific. Transverse cirri protrude dis-

Fig. 74 Oxytrichaprocera (from KAHL 1932). Ventral view from life, 110 11m.
 Oxytricha 221

tinctly beyond posterior end of cell. Dorsal cilia in life 3-4 Jim long. Three clearly dis-
cernible caudal cirri.

o c cur r e n c e and e colo g y: KAlIL (1932) discovered this little-known species

in terrestrial mosses from Wisconsin and California, USA. Records not substantiated by
illustrations: Canale Naviglio navigabile in Parma, Italy (MAnoN! & GHE1TI 1977); soft
water Escambia River in Florida, USA (PATRICK 1961). Biomass of 106 individuals about
20 mg.

Oxytricha proximata SHIBUYA, 1930 (Fig. 75)

1930 Oxytricha proximata SHIBINA, J. imp. agric. Exp. Stn Nishigahara, 1: 209,213 (Fig. 75).
1935 Oxytricha proximata SHIBINA, 1930 - KAHL, Tierwelt DtI., 30: 841.

T a x 0 nom y: BORROR (1972a) synonymised this terrestrial species with the limnetic O.
fal/ax probably because of the broad body shape. It differs from the limnetic Oxytricha
tenel/a, which has a similar body shape and size, by the lack of cortical granules. Identity
with O. bimembranata cannot be excluded (Fig. 64a). Detailed redescription necessary.

M 0 r p hoi 0 g y and b i 0 log y: In life

70-140 Jim long, about 2.2 times as long as broad. Elon-
gate ovate or elongate elliptical, posterior end obtusely
pointed. Number of micronuclei not mentioned in origi-
nal description, but very probably more than one because
the two macronuclear nodules are widely separated,
which would be rather unusual for a species with a
single, central micronucleus. Contractile vacuole dis-
tinctly in front of mid-body.
Adoral zone of rnembranelles about 40 % of body
length. Frontal and transverse cirri slightly enlarged. Five
transverse cirri, protrude somewhat beyond posterior end
of cell; the figure, however, gives the impression that
only 4 transverse cirri are present (KAHL 1932). Dorsal
cilia 2-4 Jim long. Caudal cirri inconspicuous because
marginal rows appear as being uninterrupted posteriorly.

o c cur r e n c e and e colo g y: Rare species,

probably confined to terrestrial habitats. Locus classicus
is Nishigahara near Tokyo, Japan, where SHIBUYA (1930)
Fig.75 Oxytricha proximata (from
discovered Oxytricha proximata in the soil from a field. SIUBINA 1930). Ventral view from
Records not substantiated by illustrations: soil in Bel- life, 70-140 ~m. TC = right trans-
gium (CHARDEZ 1967); moss on a rock near the village of verse cirrus.
222 SYSTEMATIC SECTION

Boldogkovaralja, Hungary, and 20-30 indo ml- I in soil of a Hungarian deciduous forest
(GELLERT 1956, 1957). Feeds on detritus (GELLERT 1956). Biomass of 106 individuals
about 60 mg.

Oxytricha pseudojusiformis DRAGESCO &

DRAGESCO-KERNEIS, 1986 (Fig. 76)

1986 Oxytricha pseudofusiformis DRAGESCO & DRAGESCo-KERNEIS, Faune

tropica1e, 26: 469.

T a x 0 nom y: This rather small species is somewhat super-

ficially described, detailed redescription thus necessary. Ge-
neric classification uncertain.

M 0 r p hoi 0 g y: After protargol impregnation 35-46 x

Fig. 76 Oxytricha pseudo-
(usiformis (from DRAGESCO
& DRAGESCo-KERNEIS 1986).
Ventral infraciIiature, pro-
targol impregnation, about
65 )lm.
16-19 Jlm. Elliptical, both ends rounded. MacronucIear nod-
ules after protargol impregnation 5--6 Jlm long; central micro-
nucleus only 1.5 Jlm across. Adoral zone of membranelles
about 30 % of body length. Undulating membranes obviously
difficult to discern. 5-7 cirri on frontal area, that is, 1-3 of the
typical 8 cirri (3 frontal, 4 frontoventra1, 1 buccal) are absent.
4-5 postoral and pretransverse ventral cirri. Constantly 5,
about 13 Jim long transverse cirri, which protrude only
slightly beyond posterior end of cell. Marginal cirri about
8 Ilm long. 3-4 inconspicuous caudal cirri. Dorsal cilia pre-
sumably rather short because neither drawn nor mentioned by
DRAGESCO & DRAGESco-KERNErs (1986).
o c c u rr e n c e and e col 0 g y: Locus classicus is the
soil of a savannah in Pendjari, Benin. Not found since. Bio-
mass of 106 individuals about 4 mg.

Oxytricha pseudosimilis HEMBERGER, 1985 (Fig. 77, Tables 3, 15)

1985 Oxytricha pseudosimilis HEMBERGER, Arch. Protistenk., 130: 405 (the holotype slide is deposited in the
Institut filr landwirtschaftliche Zoologie und Bienenkunde at the University of Bonn, Germany).

T a x 0 nom y: Differs from Oxytricha similis in the habitat (terrestrial against limnetic),
in the number of caudal cirri (3 against 2), and in the length of the dorsal cilia (8-10 Ilm
against 2-4 Ilm). The overall morphology is somewhat reminiscent of Tachysoma pe/-
lionel/um sensu Roux (1901), who also drew 2 micronuclei (Fig. 136s); however, the
 Oxytricha 223

Roux population obviously has no caudal cirri, so that a

conspecifity can be excluded. The arrangement of the fron-
toventral cirri and the shape of the undulating membranes
and adoral zone of membranelles of 0. pseudosimilis does
not agree vel)' well with the typical Oxytricha patterns; the
oral apparatus is almost of the Gonostomum type. Rede-
scription, especially live observation, drawing of the dorsal
infraciliature, and morphometric characterisation needed.

M 0 r p hoi 0 g y and b i 0 log y: After protargol im-

pregnation 95-120 x 40-50 ~m. Body margins more or
less parallel, both ends broadly rounded. Rather fragile.
Two micronuclei. Contractile vacuole about in mid-body.
Cytoplasm blackish. Adoral zone of membranelies 114-113
of body length, middle portion laterally positioned and un-
dulating membranes rather short and do not intersect (both
characters almost as in Gonostomum). Frontal and ventral Fig. 77 Oxytricha pseudosimi.
cirri slightly, transverse cirri distinctly enlarged compared lis (from HEMBERGER 1985).
with marginal cirri. 3 about 15-18 ~m long caudal cirri. Ventral infraciliature, protargol
Three dorsal kineties of body length, and usually 1, some- impregnation, 110 11m; p 222.
times 2 kineties of half body length. Dorsal cilia 8-1 0 ~m
long. Morphogenesis commences with proliferation of basal bodies adjacent to left trans-
verse cirrus. Dorsal morphogenesis in Urosomoida pattern (further details, see HEM.
BERGER 1982, P 158-161).

o c cur r e n c e and e col 0 g y: Locus classicus is a woodland soil near Puerto

Maldonado, Peru. Not found since. Biomass of 106 cells 180 mg (ForSSNER 1987a).

Oxytricha quadrici"ata BLATIERER & FOISSNER, 1988 (Fig. 78a, b, Table 15)

1988 Oxytricha granulijera quadricirrala BLAITERER & FOISSNER, Stapfia, 17: 63 (1 slide ofholotype speci·
mens is deposited in the OberOsterreichische Landesmuseum in Lim, Upper Austria).

Nom e n c I at u rea n d t a x 0 nom y: BLATIERER & FOISSNER (1988) established

O. quadricirrata as subspecies of O. granulifera because the live aspect and the infra-
ciliature are vel)' similar. However, there are some minor differences in the infraciliature
which indicate a separation at the species level: (i) in 0. granulifera the right marginal
row begins more anteriorly and terminates more posteriorly than in 0. quadricirrata; (ii)
in O. granulifera the left pretransverse ventral cirrus is located more anteriorly than in O.
quadricirrata; (iii) Oxytricha granulifera has constantly 5 (n = 25, FOISSNER & ADAM
1983b) transverse cirri, whereas 0. quadricirrata usually has 4 (mean = 3.9, n = 15). Ac-
 224 SYSTEMATIC SECTION

Fig. 78a, b Oxytricha quadricirrata (from BLAT-

TERER & FOISSNER 1988. a, b, protargol impregna-
tion). Ventral and dorsal infraciliature, 60 f.lm. Ar-
rowhead marks a single basal body pair left of ki-
nety 5. 1-5 = dorsal kineties 1-5; p 223.

cording to Article 50 c (i) of the IcZN

(1985) a change in rank of a taxon in the
species group does not affect the author-
ship of the name.

M 0 r p hoi 0 g y: In life 70-100 x

, ,
20-30 Jim. Live aspect - including corti-
, , cal granules - as in O. granulifera (see
, , p 197). A single basal body pair left of
, , dorsal kinety 5, possible a further charac-
, ,'. ,, ter which allows a separation from O.

••• - granulifera (Fig. 78b, arrowhead).

o c cur r e n c e: Locus classicus is the

a b
upper soil layer (0-5 cm, pH 6.3) with
roots and litter of a sand-hill in the 99-miles desert north of Lake Alexandrina, near Ade-
laide, Australia. Soil in Brazil (FOISSNER 1997b). Biomass of 106 cells about 13 mg.

Oxytricha rubripuncta BERGER & FOISSNER, 1987 (Fig. 79a-h, Tables 3, 15)

1987 Oxytricha rubripuncta BERGER & FOISSNER, Zoot. ]b. Syst., 114: 222 (Fig. 79!H:; 1 slide of holotype
specimens and 1 slide of paratype specimens are deposited in the OberOsterreichische Landesmuseum
in Linz, Upper Austria).
1994 Oxytricha rubripuncta BERGER & FOISSNER, 1987 - SHIN, Dissertation, p 141 (Fig. 79f-l1).

Nom e n c I a t u rea n d t a x 0 nom y: "Oxytricha rubipuncta" in BLATIERER &

FOISSNER (1988, P 72) is an incorrect subsequent spelling. According to the arrangement
of the frontoventral cirri and the slightly reduced number of transverse cirri this species
could also be classified in Urosomoida. However, since more than 4 dorsal kineties are
present, the generic classification of BERGER & FOISSNER (l987a) is accepted. Oxytricha
rubripuncta should not be confused with O. auripunctata, which has orange-yellow corti-
cal granules. The freshwater species 0. aeruginosa has a rusty granulated cytoplasm and
transverse cirri which do not protrude beyond the posterior end of the cell. In life 0. ru-
bripuncta could be confused with Urosomoida agilis, which sometimes also has a red-
dish granulation, but after protargol impregnation these two species can be easily sepa-
rated by the different number and arrangement of dorsal kineties and transverse cirri. No-
 Oxytricha 225

...
,
,
.-
..
,
,
.. .
.. ....
... ..

Fig. 79a-e Oxytricha rubripuncta (from BERGER & FOISSNER 1987a a-c, from life; d, e, protargol impregna-
tion). a) Ventral view, ISO J.lm. b) Dorsal view, ISS J.lm. The red cortical granules are arranged around the
bases of the cirri and dorsal cilia c) Cytoplasmic crystals. d, e) Ventral and dorsal infraciliature, d, e =
135 J.1m; p 224.

tohymena rubescens and Cyrtohymena muscorum, which are also reddishly granulated,
have a different oral apparatus.

M 0 r p hoi 0 g y and b i 0 log y: In life about 150 x 45 Ilm, according to SHEN

(1994) 120-160 x 40-60 Ilm. Margins slightly converging posteriorly. Somewhat con-
tractile. Macronuclear nodules in life about 21 x 13 Ilm (13-22 x 5-11 Ilm, according to
SInN 1994), after protargol impregnation 15-25 x 6-8 Ilm (mean = 20.1 x 7.2 Ilm; n =
15). 2--4, usually 2, in life about 3 Ilm sized micronuclei. Contractile vacuole about in
mid-body, during diastole with collecting canals. Cortical granules about 1 Ilm across, ar-
ranged around bases of cirri and dorsal cilia (Fig. 79b), so that specimens appear reddish
even at low magnification. Cytoplasm colourless.
Adoral zone of membranelles about 30 % of body length. The SInN (1994) popula-
tion has a distinctly lower number of adoral membranelies and marginal cirri as compared
 226 SYSTEMATIC SECTION

I l i 1

(;~
I
- I l I 1
Q./ ~ 1 l I)
i:?:::;
.:::::::: j
/22
ttl
-
'.

''\.. ::::-
,
,

ty
.~-
~
4

f
() 1

11
J
\ \. J J
,? , l \.
%
; J

:%/
/' IB
l

t
\
l

\
~
~
\.

\.
J

)
I

\ )
\
~ J
\. ) 1
.." . .
\.
I

" \. \ ) }
• \
\. \.
j
•
. h

Fig.79f-b Oxytricha rubripuncta (from SHIN 1994. f, from life; g, b, protargol impregnation). f) Ventral view,
128 Jlm. g, b) Ventral and dorsal infi"aciliature, 145 Jlm; p 224.

to the type material (Table 15). Buccal area flat. Pharyngeal fibres prominent in life.
Frontal cirri about 15 /lm long in life, bases distinctly enlarged. Rarely 1 frontal cirrus
and 1 frontoventral cirrus lacking. Frontoventral cirri arranged as in Urosomoida and not,
as erroneously stated in the original description, as in many Oxytricha species. Usually 3,
rarely up to 5 postoral ventral cirri. Usually 4 (n = IS), sometimes 3 or 5, only slightly
enlarged transverse cirri, situated very close to posterior end of cell and almost between
posterior end of marginal rows. Marginal cirri in life 10012/lID long. Invariably 6 (n =
15) dorsal kineties (Fig. 7ge, h). Dorsal cilia about 3 /lm long. Usually 3, very rarely 4
caudal cirri on kineties 1,2, and 4. The formation of the oral primordium commences ad-
jacent to the postoral ventral cirri.

o c cur r e n c e and e colo g y: Locus classicus is the upper soil layer (0-5 cm) of
an uncultivated grassland dominated by Poa sp. in the Golan Hills, Israel {BERGER &
 Oxytricha 227

FOISSNER 1987a). SHIN (1994) found it in moss-covered soils at Kwanak Campus, Seoul,
Korea. Feeds on bacteria, diatoms (Hantzschia sp.), autotrophic flagellates (Chlamydo-
monas sp.), testate amoebas (Trinema lineare), and ciliates (Sathrophilus sp.). Biomass
of 106 individuals about 200 mg (FOISSNER 1987a).

Oxytricha alfredkahli FOISSNER, 1987 (Fig. 80)

1932 Oxytricha ovalis KAHL, Tierwelt Dtl., 25: 603 (Fig. 80).
1987 Oxytricha alfredkahli FOISSNER, Arch. Protistenk., 133: 223.

Nom e n c I at u rea n d t a x 0 nom y: KAHL (1932) classified Oxytricha as subge-

nus of Oxytricha; the correct name in the original description is thus Oxytricha (Oxytri-
cha) ovalis, which is, however, a junior primary homonym of O. ovalis SCHMARDA, 1854,
and O. ova/is FROMENTEL, 1876 (now O. fromenteli FOISSNER, 1987d). Oxytricha alfred-
kahli is the replacement name for O. ovalis KAHL, 1932. Very little known and perhaps
invalid species from saline waters. Detailed redescription necessary.

M 0 r p hoi 0 g y: In life about 65 J.lm long. Elliptical to oval. Adoral zone of mem-
branelles about 40 % of body length. Sometimes 1 or 2 supernumerary cirri on frontal
area Transverse cirri long, protrude distinctly beyond posterior end of cell. Dorsal cilia
2-4 J.lm long.

o c cur r e n c e and e colo g y: Locus classicus is saline water (2.0-2.5 % salt

content) in Oldesloe near Hamburg, Germany, where it is rather common (KAHL 1932).
All other records not substantiated by illustrations: Parma River near Langhirano, Italy
(MAnoN! & GHETII 1977; further record from Italy: GRISPINI 1938); at 0.6-0.8 % salinity
in the Danzig Bay, Poland (BIERNACKA 1962); reservoirs in Azerbaijan (ALEKPEROV
1982b, 1983); Gulf of Riga, Latvia (BoIKovA 1984a, b); eutrophic lake in China during
summer (GONG 1986); bottom of freshwater ponds from AI-Hassa Oasis, Saudi Arabia
(AL-RAsHEID 1996b); Gulf of Mexico (BORROR 1962). The terrestrial record by LUZZATII
(1938) is unreliable. Biomass of 106 individuals about 13 mg.

Oxytricha discifera KARL, 1932 (Fig. 81)

1932 Oxytricha disci/era KAHL, Tierwelt Otl., 25: 604 (Fig. 81).
1933 Oxytricha disci/era KAHL 1932 - KAHL, TierweltN.- u. Ostsee, 23: 113 (Fig. 81).
1992 Oxytricha disci/era KAHL, 1930-5 - CAREY, Marine interstitial ciliates, p 192.

Nom e n c I at u rea n d t a x 0 nom y: KAHL (1932) divided Oxytricha into several

subgenera; the correct name in the original description is thus Oxytricha (Oxytricha) dis-
228 SYSTEMATIC SECTION

Fig. 80 Oxylricha alfredkahli (from KAHL 1932). Ventral view from

life, 65 11m; p 227.
Fig. 81 Oxylricha disci/era (from KAHL 1932). Ventral view from life,
200 11m; p 227.

cifera. Because of the cephalisation and some uncertain

observations (see next paragraph) the generic classifica-
tion is uncertain. Detailed redescription needed.
80
M 0 r p hoi 0 g y: In life 180-240 J1rn long (240 to
250 J1m according to AL-RAsHEID 1996a), about 4-5 times
as long as broad, that is, slender elliptical; anterior portion
(1/8 of body length) cephalised, posterior end broadly
rounded; undersized specimens with inconspicuous ce-
81 phalisation. Rather fragile. Two macronuclear nodules,
each with a micronucleus. Contractile vacuole absent or
omitted (KAHL 1932). Probably 18 frontal-ventral-
transverse cirri, however, only 5 transverse cirri unequivocally recognised (KAHL 1932);
transverse cirri V11 and VI!l rather strong, protrude distinctly beyond posterior end of
cell. Dorsal cilia long, about 8 J1m. Caudal cirri possibly lacking, that is, classification in
Oxytricha uncertain.

o c cur r e nee and e colo g y: Marine sediments, not very common but sporadi-
cally rather abundant (HARTWIG 1974, KArn.. 1932, PAITERSON et aI. 1989). Locus classi-
cus is the Kiel Bay (Baltic), where KAHL (1932) discovered O. discifera in the sediment.
Almost katharobic (KArn.. 1932). Records not substantiated by illustrations: Venice, Adri-
atic Sea (KIESSELBACH 1936); North Sea (KOSTERS 1974); mesopsammon of Black Sea
(KOVALEVA 1966, KOVAIEVA & GOLEMANSKY 1979); Barents Sea (AzOVSKY 1996, KOVAL-
JEVA 1967); mesopsammon of the Ussuri Gulf, Japan Sea (RAIKov 1963); White Sea
(BURKOVSKY 1968, 1970a, b, 1978, 1984, 1987, BURKOVSKY et aI. 1980, 1994, RAIKov
1962; further record from the USSR: BURKOVSKll & AzOVSKII 1985); in 0.1-0.4 mm sized
sand of westem Caspian Sea (AGAMALIEV 1970, 1983, AGAMALlYEV 1974); in 26-75 % of
sediment samples of Loch Eil, Scotland, a marine habitat enriched with pulp mill eftluent
(WYATI & PEARSON 1982); Kandalakshsky Bay, White Sea (AzoVSKY et aI. 1996); marine
interstitial of Saudi Arabian Gulf Islands of AI-Batinah and Abu Ali (AL-RAsHEID 1996a);
sediment of eutrophic Mullet Bay, St. George's Harbour, Bermuda (HARTWIG 1980).
Grazes on detritus and bacteria (WYATI & PEARSON 1982).

Oxytricha durhamiensis nov. spec. (Fig. 186d)

1972 Oxytricha ha/ophila (KAHL, 1932) - BORROR, Acta Protozool., 10: 66 (misidentification).
 Oxytricha 229

T a x 0 nom y: This population differs from 0. halophila in that it has cortical granules
and two micronuclei. Thus, identification with KARL's (1932) species as supposed by
BORROR (1972b) is not justified. Reinvestigation recommended. Oxytricha durhamiensis
differs from the other marine Oxytricha species by the cortical granulation.

D i a g nos i s: In life 60-100 x 33-45 !lm sized, marine Oxytricha(?) with about
0.5 flID sized spherical cortical granules. 5 or 6 dorsal kineties.

T Ypel 0 cat ion: Tidal marsh ditches and pools near Adam's Point, Durham, New
Hampshire, USA.

M 0 r p hoi 0 g y and b i 0 log y: Outline elliptical; flattened about 2: 1 dorso-

ventrally. Cell obviously not rigid because it can bend laterally, indicating that classifica-
tion in Oxytricha is correct. Two macronuclear nodules, each with one micronucleus.
Contractile vacuole present. Cortical granules in linear oblique groups of 2-5 granules
parallel to dorsal kineties; granulation rows similarly on ventral side, between most mas-
sive frontal cirri each row of granules approximately 10 in number; colour of cortical
granules not mentioned. Posterior portion of cell with irregular oblong granules about 4 x
2 !lm. Free-swimming individuals move in a slow, narrow counter-clockwise helix; most
specimens creep rapidly and irregularly forward, showing a strong tendency to veer to the
right.
Adoral zone of membranelles ("buccal cavity") 26-33 !lm long, about 25-30 adoral
membranelIes, each with 3 (likely 4, as in other hypotrichs) closely set rows of cilia. Cilia
on mediad edge of each membrane lIe form a narrow field and beat independently of
membranelIe. Paroral probably composed of basal body pairs; parora! and endoral sepa-
rated by sharp ridge. 18 frontal-ventral-transverse cirri. Frontal cirri distinctly enlarged.
Transverse cirri protrude distinctly beyond posterior end of cell. 24 right marginal cirri,
8 !lm apart in single row, marginal cirri about 20 !lm long. Dorsal cilia 4-5 !lm long and
4 Ilm apart. 3-4 caudal cirri (''terminal marginal cirri bristle-like"), about 16 !lm long;
may beat when animal is manoeuvring in a small place.

o c cur r e n c e and e colo g y: The population became abundant on or near the

surface scum of cultures derived from a collection made on 18 December, 1968 from
Adam's Point tidal marsh. Food vacuoles contained chloroplasts and diatoms (BORROR
1972b).

Oxytricha enigmatica DRAGESCO & DRAGESCO-KERNEIS, 1986 (Fig. 82a, b,

246a-j, Tables 15,53, Addendum 5)
1986 Oxytricha enigmatica DRAGESCO & DRAGESCo-KERNEIS, Faune tropicale, 26: 463.
1997 Hemigastrostyla enigmatica (DRAGESCO & DRAGESCO-KERNEIS, 1986) - SONG & WILBERT, Arch. Protis-
tenk., 148: 421 (Fig. 246a-j; detailed redescription and new combination, see Addendum 5).
230 SYSTEMATIC SECTION

T a x 0 nom y: I am rather cer-

Fig. 82a, b Oxytricha en-
igmatica (from DRAGESCO
and DRAGESCo-KERNEIs
1986. a, b, protargo\ im-
pregnation). a) Ventral in-
fraciliature, 120 ~m. b)
Ventral infraciliature of
posterior part of cell. Arrow
marks 2 additional cirri to
right of transverse cirri;
p229.
tain that this species is not an
Oxytricha. The straight undu-
lating membranes are reminis-
cent of Stylonychia, and the
longitudinal arrangement of the
frontoventral cirri is similar to
in Urosoma. However, since O.
enigmatica obviously has a
flexible body and, furthermore,
some characters are still un-
known, the originally proposed
classification is preliminarily
maintained. The slightly twisted
body and the right marginal
row which terminates immedi-
ately in front of the right trans-
verse cirrus, is reminiscent of
O. geleii. However, in this
somewhat smaller species, the
pretransverse ventral cirri are
conspicuously displaced ante-
riad and no cirri are present
right of the transverse cirri
(compare Fig. 82b with Fig.
83b). Both species occur in sa-
line waters! Redescribed and
transferred to Hemigastrosty/a
by SONG & WILBERT (l997a; see
Addendum 5).

M 0 r ph 0 log y and b i 0 log y: After protargol impregnation 75-128 x 32-56 J.lm.

Macronuclear nodules after protargol impregnation 11-19 J.lm (mean = 15 J.lm; n = 15)
long. Number of micronuclei unknown. Adoral zone of membrane lIes about 45 % of body
length, wide overlapping on right margin. Undulating membranes rather straight, arranged
side by side as in Sty/onychia pattern. Arrangement of frontal, ventral, and transverse cirri
somewhat deviating from typical Oxytricha pattern (Fig. 82a). Five (!) frontoventral cirri
and 4-5 postoral and pretransverse ventral cirri. Five enlarged transverse cirri protrude
slightly beyond posterior end of cell. Right marginal row terminates in front of right trans-
verse cirrus. Two curious cirri right of posterior end of right marginal row (Fig. 82b,
arrow). 5-7 fme caudal cirri. Number of dorsal kineties not known. According to the text
and Figure 82b the dorsal cilia are long, in Figure 82a, however, they look rather short.
 Oxytricha 231

",
I
,, , '~~~~l~ ~
...., ::, ~!mop ~
•,, ,, ...
••
,, , ,, ,
..,
, ,
,
. ,, ,
, ,,
,,
,
~
,
••
.
.-.
..
~ .• ..
••
•
,
,,

•
c d

\.

.. '
.••' e

Fig. 83a-g Oxytricha geleii (from WILBERT 1986b. a, from life and after protargol impregnation; b-g, protar-
gol impregnation). a) Ventral view, 70 Jim. b, c) Ventral and dorsal infraciliature, b, c = 45 Jim. Arrows mark
anterior and posterior postoral ventral cirrus; immediately behind are the two pretransverse ventral cirri. d)
Ventral infraciliature ofa very early morphogenetic stage, 55 11m. e) Ventral infraciliature ofa middle morpho-
genetic stage, 65 11m. C, g) Ventral and dorsal infraciliature oflate morphogenetic stages, f, g = 70 11m; p 232.

o c cur r e n c e and e colo g y: Perhaps restricted to saline waters (PATIERSON et

al. 1989). Locus classicus are saline pools near the shore of Lake Nokoue, Benin
(DRAGESCO & DRAGESCO-KERNEIS 1986). Feeds on diatoms and dinoflagellates. Biomass
of 106 individuals (after protargol impregnation) about 50 mg.
232 SYSTEMATIC SECTION

Oxytricha ge/eii (WILBERT, 1986) comb. nov. (Fig. 83a-g, Tables 3, 15)
1986 H%sticha ge/eii WILBERT, Symposia Biologica Hungarica, 33: 251.

T a x 0 nom y: Although the arrangement of the ventral cirri is reminiscent of H%sti-

cha, a detailed inspection of Figure 83b and the morphogenesis indicate that it is more
probably an oxytrichid. The unusual appearance of the infraciliature is mainly caused by
the conspicuously anteriorly displaced pretransverse ventral cirri. The slight torsion, the
right marginal row which terminates immediately in front of the right transverse cirrus,
and the saline habitat indicate a close relationship to Oxytricha enigmatica. The undulat-
ing membranes do not intersect and the number of dorsal kineties is rather low, hence the
classification in Oxytricha cannot be definite.

M 0 r p hoi 0 g y and b i 0 log y: In life 4~0 J.lm long, about 2.3 times as long as
broad. Body slightly twisted. Flexibility of body not known. Macronuclear nodules after
protargol impregnation 5-10 J.lm long (mean = 7.4 J.lm; n = 11). Contractile vacuole
slightly in front of mid-body. Adoral zone of membranelies about 1/3 of body length. Un-
dulating membranes slightly bent, but not intersecting (similar to in o. enigmatica). Pre-
transverse ventral cirri arranged immediately behind postoral ventral cirri, giving the im-
pression of a mid-ventral row (Fig. 83b). Usually 9, sometimes only 7 frontoventral and
postoral and pretransverse ventral cirri. 4-{i, usually 5 transverse cirri. Invariably 3 dorsal
kineties of body length, each with a caudal cirrus (Fig. 83c). Dorsal cilia 2-4 J.lm long.
Some morphogenetic stages are shown in Figures 83d-g. The formation of only 6
longitudinally arranged streaks (Fig. 83e) producing 18 frontal-ventral-transverse cirri
(Fig. 83b) is typical for the Oxytrichidae. The morphogenesis of the dorsal infraciliature
proceeds in Gonostomum pattern.

o c cur r e n c e and e colo g y: Probably restricted to saline habitats. Locus classi-

cus of this somewhat unusual species is the Little Manitou Lake (salinity 9.6 %) in Sas-
katchewan, Canada. Feeds on diatoms. Biomass of 106 individuals about 6 mg.

Oxytricha ha/ophi/a KARL, 1932 (Fig. 84a, b)

1932 Opisthotricha ha/ophi/a KAHL, Tierwelt DtI., 25: 611 (Fig. 84a, b).
1933 Opisthotricha halophila KAHL 1932 - KAHr., Tierwelt N.-u. Ostsee., Lieferung 23 (feillI. C3): 114.

Nom e n c I a t u rea n d t a x 0 nom y: Incorrect subsequent spellings: Opisthotri-

cha ha/ophy/a (TUCULESCU 1965, p 201); Oxytricha (Opis/otricha) ha/ophi/a (TUCULESCU
1965, P 160). KAHL (1932) classified Opisthotricha as subgenus of Oxytricha; the correct
name in the original description is thus Oxytricha (Opisthotricha) ha/ophi/a. Oxytricha
ha/ophi/a is possibly the senior synonym of the limnetic 0. parahalophila (Fig. 84c).
 Oxytricha 233

Oxytricha halophila sensu AGAMALIEV

(1978, 1983) is rather certainly not identi~
cal to the population described by KAHL
(1932) because 2-3 micronuclei are re-
ported; it is classified as insufficient rede-
scription (Fig. 233~ m). The same is true
for O. halophila sensu CZAPIK & JORDAN
(1976a) because the single micronucleus is
not described (Fig. 232m). Oxytricha halo-
phila sensu BORROR (1972b) differs signifi-
cantly from the type population in that it
has two micronuclei and cortical granules,
strongly indicating that is a different spe-
cies (see O. durhamiensis; Fig. 186d).

M 0 r p hoi 0 g y and b i 0 log y: In

life about II 0 Ilm long, 4 times as long as
broad. Long elliptical (Fig. 84a). Contrac- C

tile vacuole at about level of buccal vertex. Fig. 84a, b Oxytricha ha/ophila (from KAHL 1932).
Adoral zone of membranelles 114 of body Ventral views from life, a = 110 11m, b = '1 11m. Ar-
length. About 113 of length of transverse row marks single micronucleus; p 232.
Fig.84e Oxytricha paraha/ophi/a (from WANG &
cirri protrudes beyond posterior end of cell. NIE 1935). Ventral view, 80 11m; p 170.
Marginal cirri inserted very close to body
margin. Three fme, setiform caudal cirri,
conspicuously elongated. Dorsal cilia 2--4 Ilm long. KAHL (1932) mentioned another very
similar population (Fig. 84h) under this species name with an oblique right anterior end
and transverse cirri protruding somewhat further beyond the posterior end.

o c cur r e n c e and e colo g y: Probably confmed to saline waters (PATIERSON et

al. 1989). Locus classicus is the detritus in a marine ditch of the Sylt Island, Germany
(KAHL 1932). The second population described by KAHL (1932, Fig. 84h) occurred in sa-
line inland waters of Oldesloe, near Hamburg, Germany. Records not substantiated by il-
lustrations: Bay of Danzig, Poland (BIERNACKA 1962); west coast of Caspian Sea (AGA-
MALIEV 1971); saline lake near Black Sea, Romania (TuCOLESCO 1962b); Barents Sea
(AzOVSKY 1996). Biomass of 106 individuals about 23 mg.

Oxytricha oxymarina nom. nov. (Fig. 85a-e)

1932 Oxytricha marina KAHL, Tierwelt Dtl., 25: 603 (Fig. 85b).
1933 Oxytricha marina KAHL 1932 - KAlIL, Tierwelt N.- u. Ostsee, Lieferung 23 (Teil II.el): 113.
1972 Oxytricha marina KAHL? - DRAGESCO, Annis Fac. Sci. Univ. fed. Cameroun, 11: 88 (Fig. 85a).
1974 Oxytricha marina KAlIL, 1932 -JONES, Univ. South Alabama Monogr., 1: 41 (Fig. 8Se).
 234 SYSTEMATIC SECTION

Fig. 85a-d Oxytricha oxymarina (a, from DRAGESCO 1972b; b, from l<AHL 1932; e, d, from AGAMALIEV 1978.
a, protargol impregnation; b, from life; e, d, wet silver impregnation). a, c) Ventral infraeiliature, a = 90 11m, e
= 85 11m. b) Ventral view, 110 11m. d) Dorsal infraciliature, very likely not correct, 85 J1ffi; P 233.

1978 Oxytricha marina KAHL, 1932 - AGAMALIEV, Aeta Protozool., 17: 437 (Fig. 85e, d).
1983 Oxytricha marina KAHL, 1932 - AGAMALIEV, Ciliates of Caspian Sea, p 108 (Fig. 85e, d).
1986 Oxytricha marina KAHL, 1932 - DRAGESCO & DRAGESCo-KERNEIS, Faune tropicale, 26: 472.

Nom e n c I at u rea n d t a x 0 nom y: KAm.. (1932) divided Oxytricha into several

subgenera (Table 7); thus, Oxytricha (Oxytricha) marina kARL, 1932, is a primary homo-
nym of Oxytricha (Steinia) marina KAm.., 1932 (Tierwelt Dtl., 25: 614). Since I found no
literature where either of these two species was given a new name as demanded by the
IcZN (1985), I hereby rename the first species: Oxytricha oxymarina nom. nov., for Oxy-
tricha (Oxytricha) marina. The other species is now considered to be a Cyrtohymena, C.
marina (KAm.., 1932) FOISSNER, 1989.
The descriptions listed above do not agree very well in all characters, so that an iden-
tification should be based mainly on the original description. Oxytricha marina sensu
 Oxytricha 235

BIERNACKA (1967; Fig. 226d) and sensu GANAPATI & RAo (1958;
Fig. 228a) are insufficient redescriptions. Detailed reinvestiga-
tion needed.

M 0 r p hoI 0 g y and b i 0 log y: In life 80-120 /.lm long.

Body margins parallel or slightly converging anteriorly. Both
ends broadly rounded. Slightly contractile. Macronuclear· nod-
ules, according to DRAGESCO (1972b), after protargol impregna-
tion 14-18/.lm; according to JONES (1974), in life only
10-12/.lm long. 2-3 about 3/.lm sized spherical micronuclei.
Contractile vacuole distinctly in front of mid-body. Cytoplasm
colourless, posterior cell portion usually with dark granules (Fig.
85b).
Adoral zone of membranelles about 113 of body length, af-
ter DRAGESCO (1972b) 30-32, after AGAMALIEV (1978) 30-35
membranelles. Buccal area obviously very small. According to
KAHL (1932) and DRAGESCO (1972b) transverse cirri protrude
distinctly beyond posterior end of cell, after JONES (1974) they
do not. Number of marginal cirri: 33-39 right and 31-33 left Fig. 8Se Oxytricha OX)'-
(DRAGESCO 1972b), 24-26 right and 18-20 left (AGAMALIEV marina (from JONES
1974). Ventral view from
1978). Dorsal infraciliature shown in Figure 85d rather certainly
life, 110 11m; p 233.
incorrect. Dorsal cilia short, caudal cirri obviously inconspicu-
ous.

o c cur r e n c e and e colo g y: Common in brackish and marine habitats and in

inland salt waters. Locus classicus is a marine ditch on Sylt Island, Germany (KAHL
1932). DRAGESCO (1972b) found 0. oxymarina in a saline pond at Lake Tchad, Tchad.
JONES (1974) observed it in Mobile Bay (Alabama, USA), AGAMALIEV (1978) in the Cas-
pian Sea.
Records from marine habitats not substantiated by illustrations (see also PATIERSON et
al. 1989): Bay of Naples, Adriatic (NOBill 1957); Kiel Bay and supra-littoral zone of the
Bay of Eckemforder, Baltic Sea (BOCK 1952, HARTWIG 1974); Danzig Bay and a harbour
in Poland (BIERNACKA 1962, 1968); Gulf of Riga, Baltic Sea (BoIKovA 1984b); Caspian
Sea (AGAMALIEV 1971, 1983, 1990); saline lake in Romania near Black Sea (TUCOl.ESCO
1962b); Barents Sea (AzOVSKY 1996); Logy Bay in Newfoundland, Canada, Atlantic
Ocean (LACKEY & LACKEY 1970); marine interstitial of Saudi Arabian Gulf Islands of AI-
Batinah, Abu Ali, and Tarot (AL-RAsHEID 1996a, 1997); Bay of Bengal, Indian Ocean
(RAo & GANAPATI 1968).
Feeds on diatoms, small flagellates, and small ciliates. Biomass of 106 individuals
about 20 mg (population ofKAHL 1932).
236 SYSTEMATIC SECTION

Oxytricha saltans (COHN, 1866) REES, 1881 (Fig. 86a-k, 87a-l, 88a-g,
8ge, 245a-e, Table 15, Addendum 4)

1866 Actinotricha sa/tans COHN, Z. wiss. Zool., 16: 283 (Fig. 87!H:).
1867 Actinotricha sa/tans COHN - QUENNERSTEDT, Acta Univ. lund., 4: 40 (Fig. 87t).
1881 Oxytricha sa/tans - REES, Zur Kenntniss der Bewimperung der hypotrichen Infusorien, p 16.
1882 Actinotricha sa/tans, COHN - KENT, Manual infusoria IT, p 790.
1883 Actinotricha sa/tans (COHN) - MAuPAs, Archs Zool. expo gen., 1: 544 (Fig. 87i,j).
1884 Oxytricha sa/tans, COHN sp. - REES, Tijdschr. ned. dierk. Vereen., I: 646,647 (Fig. 87e, g).
1884 Oxytricha sa/tans COHN sp. - ENlZ, Mitt. zoo!. Stn Neapel, 5: 367 (Fig. 88a-g).
1886 Actinotricha hyalina PEREYASLAWZEWA, Zap. novoross. Obshch. Estest, 10: 88 (Fig. 8ge).
1926 Actinotricha sa/tans CoHN - LEpSI, Infusorien des SOSswassers und Meeres, p 86 (Fig. 87h).
1932 Actinotricha sa/tans COHN, 1866 - KAHL, Tierwelt Dtl., 25: 604 (Fig. 87d).
1933 Tachysoma (Actinotricha) sa/tans COHN 1866 - KAHL, Tierwelt N.- u. Ostsee, 23: 113.
1934 Actinotricha sa/tans COHN 1866 - WANG, Rep. mar. bioI. Ass. China, 3: 63 (Fig. 871).
1965 Actinotricha sa/tans COHN - UPSI, Protozoologie, p 975 (Fig. 87k; poor drawing).
1972 Tachysoma sa/tans (COHN, 1866) - BoRROR, J. Protozool., 19: 15.
1991 Oxytricha sa/tans (COHN, 1866) KARL, 1932 - SONG, SHIN & KIM, Korean J. syst Zool., 7: 235 (Fig.
86a-k; authoritative redescription).
1997 Oxytricha sa/tans (COHN, 1866) - SONG & WILBERT, Arch. Protistenk., 148: 420 (Fig. 245lH:).

Nom e n cia t u rea n d t a x 0 nom y: Actinotricha hyalin in PEREYASLAWZEWA

(1886, P 97) is an incorrect original spelling. KAHL (1932) classified Actinotricha as sub-
genus of Oxytricha; the correct name in his revision is thus Oxytricha (Actinotricha) sal-
tans (COHN, 1866). See Addendum 4 for an important nomenclatural note. The combining
author is REEs (1881), who correctly included Actinotricha in Oxytricha. Caudal cirri are
present, proving that the classification in Tachysoma, as suggested by BORROR (1972a), is
not justified. Synonymy of Actinotricha saltans and Actinotricha hyalina was already
suggested by KAHL (1932). The description is mainly based on the authoritative rede-
scription by SONG et al. (1991). The ALzAMORA (1929; Fig. 89a) and LEPSI (1927; Fig.
89b, c) populations are insufficiently redescribed, although the distinctly shortened right
marginal row in Figures 89b, c indicates that LEPSI'S identification was correct. The iden-
tification by DRAGESCO (1963; Fig. 89d) is rather uncertain and was thus not included in
the list of synonyms (see insufficient redescriptions).

M 0 r ph 0 log y and b i 0 log y: In life 50-80 x 15-25 !lm (SONG et al. 1991),
60!lm (CoHN 1866; erroneously "0,6 Mm" [= 600 !lm] are given), 80-1OO!lm (MAUPAS
1883), 50-82!lm (REES 1884), 50-100 x 3~0!lm (ENTZ 1884), 50-100 x 35!lm

Fig. 86a-f Oxytricha sa/tans (from SONG et al. 1991. a-d, from life; e, f, protargol impregnation). a) Ventral --)
view, 80 Ilm. b) Lateral view. Nuclear apparatus after Feulgen staining. c, d) Well (with ingested ciliates) and
poorly fed (with greasily shining globules) specimens, c = 781lm, d = 70 Ilm. e, f) Ventral and dorsal infra-
ciliature, 75 Ilm. Long arrow in (e) marks anterior-most postoral ventral cirrus, short arrow denotes anterior
end of right marginal cirral row. Anterior arrowhead marks buccal cirrus, posterior one denotes caudal cirri;
p236.
 Oxytricha 237

# #
I
, '"

'
'"
f # • .'
...
~
, .. ~
e
238 SYSTEMATIC SECTION

9 h

' .. k

..

Fig. 86g-k Oxytricha saltans (from SONG et aI. 1991. g-k, protargol impregnation). g-j) Schematic illustra-
tions of variability of dorsal infraciliature. k) Detail of ventral ciliature showing fibres anchoring cirri in the
cell. Explanation of original labels: AL = anterior longitudinal fibre, C = cirrus, LMC = left marginal cirral
row, PF = peripheral fibre, PL = posterior longitudinal fibre, RMC = right marginal cirral row, VC = postoral
ventral cirrus; p 236.

(WANG 1934); according to COHN (1866), about twice as long as broad. Ovoid to
elongate-Ianceolate, usually elliptical and anterior portion slightly converging. Flexible
but not distinctly contractile. Macronuclear nodules distinctly separate, after protargol
impregnation 11-21 x 8-13 !lm (mean = 16 x 10 Jlm; n = 16).2-3 spherical micronuclei.
Contractile vacuole lacking (CoHN 1866, SONG et al. 1991); according to the detailed re-
description by ENTZ (1884), contractile vacuole in mid-body but period between systoles
usually rather long, so that cells have no conspicuous vacuole for much of the time. Corti-
cal granules lacking. Cytoplasm colourless, posterior cell portion often packed with
1-5 !lm sized, greasily shining globules (Fig. 86d); in anterior and posterior end often
one conspicuous shining globule each (like the "FettkOmchen" in Tachysoma pellionel-
lum). Movement characteristic, that is, often immobile for several minutes with radiating
distal adoral membranelies, then jumping suddenly; also evenly creeping among algae
(COHN 1866). Notes on movement, see BULLINGTON (1925).
Adoral zone of membranelIes about 25 % of body length, that is, buccal cavity rather
short and narrow. Distal (about 5) membranelles about 20 !lm long, spine-shaped and
 Oxytricha 239

g
a·
e

Fig. 87a-1 Oxytricha saltans (a-c, from COlIN 1866; d, after ENlZ and REEs from KAHL 1932; e, g, from REEs
1884; f, after QUENNERSTEDT 1867; h, from LEPSI 1926b; i,j, from MAUPAS 1883; k, from UPSI 1965; I, from
WANG 1934. a-I, from life). a, d, e, g-i, k, I) Ventral views, a = 60 J.lm, d = 70 J.lm, e, g = 50-82 J.lDl, h =
100 J.lm, i = 90 J.lm, k = size not indicated, 1= 95 J.lm. b, c) Small specimens. 1) Ventral ciliature seen from
dorsal, 83 J.lm. j) Left lateral view, 90 J.lm; p 236.

thus radiating (thus the name Actinotricha!). Undulating membranes rather short, parallel,
that is, not in the typical Oxytricha pattern. Proximal 2-3 membranelles covered by lip
and usually separate from anterior (main) portion (for example, Fig. 86a, e, 87a, f, i, I,
88a). All cirri, except transverse cirri, rather fme. Arrangement of cirri rather invariable
(Fig. 86e). Constantly (n = 26) 3 frontal cirri, only 3 frontoventral cirri, 1 buccal cirrus
 240 SYSTEMATIC SECTION

88a b

\'i;\~\"\\~. fk.
1/ .

~"'~~.'
~ :. ~·.~i
:.\.~
. . •. :. . . .
'~ri
'.• •. • . <.:
. . .••.. •. .} . . :. .•. .••.. .

~. :.:. •. . . . '...•........ ·~~J·'.···:tt·

1[.<: . . .' .. \{.~.......
. ~.
-A0'.. ,,;1

l..\~j"f?(
\<~~i<'!
~~~+r
nf d

Fig. 88a-g Oxytricha saltans (from ENTZ 1884. From life). a-d) Dorsal (showing mainly ventral side),
ventral, and lateral view, 50-100 Jlm. e) Dorsal view of a late morphogenetic stage. r, g) Prominent broom-
shaped transverse cirrus and radiating adoral membranelles; p 236. Fig. 89a-d Insufficient redescriptions of
Oxytricha saltans (a, from AizAMORA 1929; b, c, from LEpSI 1927; d, from DRAGESCO 1963.8-<), from life; d,
protargol impregnation). a) Ventral side seen from dorsal, 60 Jlm. b, c) Ventral views, 100 Jlm. Shortened right
marginal cirra1 row indicates that the LEPSI identification is correct. d) Ventral infraciliature, 35-75 Jlm; p 236.
 Oxytricha 241

(near buccal vertex), 3 postoral ventral cirri (about in mid-

body), 2 pretransverse ventral cirri. Invariably (n = 26) 5
very strong, broom-shaped transverse cirri (Fig. 88t) which
protrude distinctly beyond posterior end of cell; COHN
(1866) counted 6-8 transverse cirri which is likely a misob-
servation. Right marginal cirral row begins in mid-body.
Usually 5, rarely 6 (mean = 5.3; n = 11) dorsal kineties (in-
cluding dorsomarginal rowls]), cilia 9-13 ~m long (Fig.
86a, f-j). Constantly (n = 7) 3 fine caudal cirri which con-
sist of2 basal body pairs only (Fig. 86e, t).

o c cur r e n c e and e colo g y: Common in marine

biotopes. Locus classicus is Heligoland, Germany, North
Sea (COHN 1866). Locus classicus of the synonym, Actina-
tricha hyalina, is the Black Sea (PEREYASLAWZEWA 1886).
SONG et al. (1991) collected O. saltans from shrimp culture
ponds (containing a lot of organic matter) around the Gulf
of Jiaozhou, Qingdao, China, where it occurred from May
to July. SONG & WANG (1993) found it in marine cultiva-
tions along the Bohai Bay and in the Yellow Sea, China.
Fig. 8ge Actinotricha sa/tans
Records largely substantiated by illustrations: Kattegat (from PEREYASLAWZEWA 1886).
(Sweden), Baltic (QUENNERSTEDT 1867, 1869); Bay of Ventral view from life, size not
Naples and coast of Algiers, Mediterranean (ENTZ 1884, indicated; p 236.
MAUPAS 1883, REES 1881); the Netherlands, North Sea
(REES 1884); highly abundant from a standing culture of
sea-water from Bay of Amoy, China, in late July (WANG 1934). Records not substantiated
by illustrations: North Sea (KUSTERS 1974); sediment of Kiel Bay, Baltic (BOCK 1952,
HARTWIG 1974); Genoa and Venice, Mediterranean and Adriatic (GRUBER 1884, KIEssEL-
BACH 1936); saline lake sometimes supplied with sea water near Sebastopol, Ukraine (DA-
GAJEVA 1930); Black Sea (JELIASKOWA-PASPALEWA 1933, LEPSI 1965); harbour of Ostend
(Belgium), Atlantic Ocean (PERSOONE 1968); coast of the Netherlands, Atlantic Ocean
(REES 1881); Caspian Sea (AGAMALIEV 1971, 1983); rock pools in New Hampshire, At-
lantic Ocean (MARTINEZ 1980); Rio de Janeiro Bay, Atlantic Ocean (FARIA et aI. 1922);
Barents Sea (KOVAIJEVA 1967). Feeds on bacteria and small flagellates and ciliates (SoNG
et al. 1991, Fig. 86c). SONG et al. (1991) found O. saltans at 19-21 °C, pH 7.9-8.0, about
100 % oxygen saturation, and 3.0-3.2 % salinity. Temperature optimum for growth 34
°C, lethal (LDso) temperature 37 °C (MARTINEZ 1980).

Oxytricha tricorn is MILNE, 1886 (Fig. 90)

1886 Oxytricha tricornis MILNE, Proc. Manchr Fld Nat. Archaeol. Soc., 18: 52 (Fig. 90).
242 SYSTEMATIC SECTION

1932 Oxytricha tricornis MILNE, 1886 - KARL, Tierwelt Dtl., 25: 603.

Nom e n c I at u rea n d t a x 0 nom y: KAHL (1932) divided Oxytricha into several

subgenera; the correct name in his revision is thus O. (Oxytricha) tricornis. Little known
species (very likely because of small size) urgently in need of de-
tailed redescription. Classification in Oxytricha uncertain. BORROR
(1972a) synonymised it with Gonostomum strenuum, which is,
however, broader and lives in freshwater.
MADRAZO-GARlBAY & LoPEZ-OCHOTERENA (1973) recorded and
figured a "Stylonichia tricornis (TAGLIANI, 1922)" from freshwater
in Mexico (Fig. 240j). This is not a ''new combination" of the
MILNE species, but of TAGLIANI'S (1922) Dipleurostyla tristyla,
which is very likely a fragment of a (oxytrichid?) hypotrich. Fur-
thermore, Figure 240j is only a redrawing ofTAGLIANI's illustration.

M 0 r p hoi 0 g y: In life about 70 11m long and five times as long

Fig. 90 Oxytricha
tricornis (from MIL- as broad, that is, very slender, widest in mid-body, tapering gradu-
NE 1886). Ventral ally toward ends. Flexible and highly contractile. Two macronu-
view from life, about clear nodules. Three very conspicuous adoral membranelles, one
70 JUIl. Note slender fourth to one third of body length (other, small adoral membranelles
outline and three
possibly overlooked). Eight or ten scattered "ventral cirri" and 6
conspicuous adoral
membranelles; p 242. long transverse cirri. Two rows of marginal cirri. Dorsal cilia likely
short (''minute'').

o c cur r e nee: Locus classicus is very likely the Irish Sea near
Glasgow, England (MILNE 1886). AGAMALIEV (1971, 1973a) recorded Oxytricha tricornis
from the Caspian Sea.

Oxytricha species at present classified in taxa not revised in this book

Opisthotricha elongata GRANDORI & GRANDORI, 1934, Boll. Lab. Zoo1. agr. Bachic. R. 1st.
sup. agr. Milano, 5: 290, Tavola XIII, fig. 278. Remarks: GRANDORI & GRANDORI (1934)
classified Opisthotricha as subgenus of Oxytricha so that the correct name is Oxytricha
(Opisthotricha) elongata, and it is thus not a primary homonym of Opisthotricha elon-
gata SMITH, 1897. According to the shape of the adoral zone of membrane lies, GRANDORI
& GRANDORI'S (1934) species very likely belongs to the TrachelostylalTrachelochaeta
group. Opistotricha elongata on page 338 is an incorrect subsequent spelling of Opistho-
tricha. In life about 120 11m long. Soil in Italy. Detailed redescription necessary.

Oxytricha alba FROMENTEL, 1876, Etudes microzoaires, p 268, Planche XIII, fig. 16. Re-
marks: Junior synonym of Holosticha pullaster (MOLLER, 1773) ForSSNER, BLATIERER,
BERGER & KOHMANN, 1991 (for details, see FOISSNER et al. 1991, p 240).
 Oxytricha 243

Oxytricha auricularis CLAPAREDE & LACHMANN, 1858, Mem. Inst. natn. genev., 5: 148,
Planche V, fig. 5,6. Remarks: According to CAREY & TATCHELL (1983) the junior syno-
nym of EpiclintesJelis (MOLLER, 1786) CAREY & TATCHELL, 1983.

Oxytricha capitata PEREYASLAWZEWA, 1886, Zap. novoross. Obshch. Estest., 10: 91, fig.
15. Remarks: KAHL (1932) transferred it to Amphisiella GOURRET & ROESER, 1888.

Oxytricha crassa CLAPAREDE & LACHMANN, 1858, Mem. Inst. natn. genev., 5: 147,
Planche VI, fig. 7. Remarks: FROMENfEL (1875) fixed it as type species of Oxytricha,
which was, however, neglected by later authors. KAHL (1932, p 588) transferred it (as
"Trichotaxis (Oxytricha) crassa") to the subgenus "Holosticha Trichotaxis" ("Holosti-
cha (I'richotaxis) crassel'; incorrect spelling of Trichototaxis).

Oxytrichaflava COHN, 1866, Z. wiss. Zool., 16: 288, Tafel XV, Fig. 27-29. Remarks:
Now Pseudokeronopsisflava (COHN, 1866) WIRNSBERGER, LARSEN & UHLIG, 1987.

Oxytricha flava var. carnea CoHN, 1866, Z. wiss. Zool., 16: 288. Remarks: Now Pseu-
dokeronopsis carnea (COHN, 1866) WIRNSBERGER, LARSEN & UHLIG, 1987.

Oxytricha fusca PERTY, 1852, Zur Kenntniss kleinster Lebensfonnen, p 154, Tafel VI,
Fig. 19A, B. Remarks: Junior synonym of Urostyla grandis EHRENBERG, 1830.

Oxytricha gibba CLAPAREDE & LACHMANN, 1858, Mem. Inst. natn. genev., 5: 144, Planche
V, fig. 8. Remarks: According to CLAPAREDE & LACHMANN (1858) possible identical with
Oxytricha gibba (MOLLER, 1786) sensu EHRENBERG (1938), after KAHL (1932, p 547) an
Uroleptus.

Oxytricha kessleri WRZESNIOWSKI, 1877, Z. wiss. Zool., 29: 275, Tafel XIX, Fig. 12-15.
Remarks: Now Holosticha kessleri (WRZESNIOWSKI, 1877) WRZESNIOWSKI, 1877 (see
FOIsSNER et al. 1991, p 228).

Oxytricha longi-caudata WRIGIff, 1862, Q. Jl microsc. Sci., 2: 220, Planche IX, Fig. 7, 8.
Remarks: Junior synonym of Psammomitra retractilis (CLAPAREDE & LACHMANN, 1858)
BORROR, 1972, a species recently transferred to Uroleptus (SONG & WARREN 1996).

Oxytricha micans ENGELMANN, 1862, Z. wiss. Zool., 1: 387. Remarks: Junior synonym of
Holosticha pullaster (MOLLER, 1773) FOISSNER, BLATIERER, BERGER & KOHMANN, 1991
(see FOISSNER et al. 1991, p 240).

Oxytricha oculata MERESCHKOWSKY, 1877, Trudy imp. S-peterb. Obshch. Estest., 8: 232,
Tafel I, fig. 9, 10. Remarks: Uncertain species! According to KAHL (1932, P 582) a
Holosticha.
244 SYSTEMATIC SECTION

Oxytricha pernix WRZESNIOWSKI, 1877, Z. wiss. ZooI., 29: 273, Tafel XIX, Fig. 10, 11.
Remarks: Now Pseudokeronopsis pernix (WRZESNIOWSKI, 1877) BORROR & WICKLOW,
1983.

Oxytricha retractiUs CLAPAREDE & LACHMANN, 1858, Mem. Inst. natn. genev., 5: 148,
Planche V, fig. 3, 4. Remarks: Now Uroleptus retraetiUs (CLAPAREDE & LACHMANN, 1858)
SONG & WARREN, 1996. Previously classified in Psammomitra (BORROR 1972).

Oxytricha rubra EHRENBERG, 1835, Abh. preuss. Akad. Wiss., year 1835: 164 and Mit-
theilungen der Gesellschaft naturforschender Freunde zu Berlin, year 1835: 5. Remarks:
Now Pseudokeronopsis rubra (EHRENBERG, 1835) BORROR & WICKLOW, 1983.

Oxytricha scuttelum COHN, 1866, Z. wiss. Zoo!., 16: 287, Tafel XV, Fig. 43-46.
Remarks: According to KAHL (1932, P 579) a Holostieha.

Oxytricha tubieola GRUBER, 1880, Z. wiss. ZooI., 33: 450, Tafel XXVI, Fig. 11, 12. Re-
marks: Now Stichotrieha tubicola (GRUBER, 1880) BORROR, 1972.

Oxytrieha urostyla CLAPAREDE & LACHMANN, 1858, Mem. Inst. natn. genev., 5: 141,
Planche V, fig. 2. Remarks: Now Pseudourostyla urostyla (CLAPAREDE & LACHMANN,
1858) BORROR, 1972. According to FOISSNER et al. (1991, p 260) a supposed synonym of
Paraurostyla weissei (STEIN, 1859) BORROR, 1972 (see also Fig. 2271, m).

Oxytricha velox QUENNERSTEDT, 1869, Acta Univ. lund., 6: 20, Fig. 20, 21. Remarks:
KAHL (1932, P 589) transferred it to "Trichotaxis" (incorrect spelling of Trichototaxis).

Oxytricha viridis PEREYASLAWZEWA, 1886, Zap. novoross. Obshch. Estest., 10: 91, Fig.
16. Remarks: A junior primary homonym of 0. viridis FROMENTEL, 1876, a species inde-
terminata. According to BORROR & WICKLOW (1983), O. viridis PEREYASLAWZEWA is syn-
onymous with Pseudokeronopsis rubra; thus, I do not replace its name.

Oxytricha wrzesniowskii MERESCHKOWSKY, 1877, Trudy imp. S-peterb. obshch. Estest., 8:

231, Tafel II, fig. 6. Remarks: According to KAHL (1932, P 583) a Holosticha.

Species indeterminata

DUMAS (1929, 1930, 1937), a French amateur protistologist, described about 550 new
species, mainly ciliates from various biotopes of the French Massif Central. Probably
most of the about 80 hypotrichous species (mainly assigned to Kerona, Oxytricha, and
Stylonychia) are species indeterminata. A list of all DUMAS' names is provided by FOISS-
NER (1 995a).
 Gxytricha 245

New species of Oxytricha - SESHACHAR & KAsTURI BAI, 1963, Arch. Protistenk., 106: 456
(Fig. 222c, d). Remarks: In life 200-450 x 100-150 flm. Anterior end narrow rounded,
posterior pointed. 3 (!) about 10 x 7 flm sized macronuclear nodules, sometimes with a
thin, threadlike connection. 7-14 micronuclei. Contractile vacuole large, situated at pos-
terior end (?). Adoral zone of membrane lIes about 1/2 of body length. Marginal rows
converging posteriorly. 5 transverse cirri, 3 of them are elongated and protrude beyond
posterior end of cell. Isolated from a freshwater fish tank in Bangalore, India. The ani-
mals were cultured in 0.01 % Knops solution, Erd-Schreiber, and hay infusion and were
fed with Quaker oats, lettuce infusion, and Horlick's malted milk. SESHACHAR & KAsTURI
BAI (1963) did not mention a species name. The position of the contractile vacuole in the
posterior portion is rather unreliable. Sterkiella thompsoni (Fig. 182f-k) also has 3 mac-
ronuclear nodules; however, this species is much smaller (90-139 flm long) and possibly
endemic to Antarctica.

Gpisthotricha sordis LEPSI, 1965, Protozoologie, p 22 (Fig. 224h). Remarks: In life (?)
130 flm long. Polysaprobic. Oxytricha sordis in LEPSI (1965, P 975) is very likely only
another combination of the same species.

Opistotricha terricola GEU. ERT, 1957, AnnIs Inst. bioI. Tihany, 24: 21 (Fig. 222b). Re-
marks: Incorrect subsequent spelling of Gpisthotricha. "G. tericola" in LEPSI (1965, P
975) is also an incorrect subsequent spelling. About 80 flm long. 2 spherical macronu-
clear nodules, 2 micronuclei. About 20 adoral membranelles, adoral zone of mem-
branelles 25 % of body length. 3 frontal, 1 buccal, and only 2 frontoventral cirri. Postoral
ventral cirri lacking, 2 pretransverse ventral cirri, 4 transverse cirri, only 1 long caudal
cirrus. Feeds on bacteria. Soil ofa deciduous forest in Hungary. The arrangement of the
cirri is rather unusual; somewhat reminiscent of Urosoma similis.

Oxitricha joblolii BORY DE SAINT-VINCENT in LAMOUROUX et aI., 1824, Encyclopedie

methodique, p 596. Remarks: According to EHRENBERG (1838, P 367) possible synony-
mous with Stylonychia pustulata or Tachysoma pellionellum.

Oxitricha nfun. 1 - IzQUIERDO, Protozoos, P 191 (Fig. 229g). Remarks: Incorrect subse-
quent spelling of Oxytricha. In life 130 flm long. Freshwater in Chile.

Oxitricha nWn. 2 - IzQUIERDO, Protozoos, P 192 (Fig. 229t). Remarks: Incorrect subse-
quent spelling of Oxytricha. In life 140 flm long. Freshwater in Chile.

Oxitricha nfun. 3 - IzQUIERDO, Protozoos, p 191 (Fig. 22ge). Remarks: Incorrect subse-
quent spelling of Gxytricha. In life 92 flm long. Freshwater in Chile.
246 SYSTEMATIC SECTION

Oxitricha pullicina BORY DE SAINT-VINCENT in LAMOUROUX et aI., 1824, Encyclopedie

methodique, p 595. Remarks: Established for a broad form of Trichoda pulex MOLLER,
1773.

Oxitricha variabilis BORY DE SAINT-VINCENT in LAMOUROUX et aI., 1824, Encyclopedie

methodique, p 597. Remarks: Possible only a (unjustified?) replacement name for Tri-
choda praeceps MOLLER, 1786. See also Oxytricha variabilis GROLIERE, 1975.

Oxytricha acuminata DUMAS, 1930, Microzoaires, p 57 (Fig. 239t). Remarks: Oxytricha

acuminata DUMAS is a junior primary homonym of O. acuminata STOKES, 1887. Because
it is a species indeterminata, I do not replace the name. In the legend to the figure
(planche XXVIII, fig. 24) designated as "Oxytrichafastigiata (labiata}". Infusion ofter-
restrial moss, France.

Oxytricha acuminata MAIwAN, 1977, Rec. Surv. India, 72: 223 (Fig. 234d). Remarks:
Oxytricha acuminata MAIwAN is the junior primary homonym of O. acuminata STOKES,
1887, O. acuminata DUMAS, 1930, and O. acuminata VUXANOVICI, 1963 (see previous
and next entry). Because it is a species indeterminata, I do not replace the name. Al-
though I did not check the type slides (Z. S. I. Reg. No. Pt 694-701), I am rather sure that
this taxon is indeterminable. 100-115 x 50-60 Jim. Nine frontoventral, 5 ventral, and 5
transverse cirri; marginal rows interrupted posteriorly. Freshwater in Fatch Sagar, Udai-
pur, India.

Oxytricha acuminata VUXANOVICI, 1963, Studii Cerc. BioI., 15: 215 (Fig. 223k).
Remarks: Oxytricha acuminata VUXANOVICI, 1963, is the junior primary homonym of O.
acuminata STOKES, 1887 (now in Urosoma) and O. acuminata DUMAS, 1930. Because it
is a species indeterminata, I do not replace the name. A special feature of this form is a
conspicuous single cirrus just behind the contractile vacuole, which is displaced posteri-
orly. Stagnant waters in Romania.

Oxytricha ambigua DUJARDIN, 1841, Zoophytes, p 419, Planche XI, fig. 15. Remarks: A
nominal species and not a redescription of Trichoda ambigua MOLLER, 1786 (now Spi-
rostomum ambiguum) because, unlike in other redescriptions by DUJARDIN, no synonyms
are listed in a footnote,

Oxytricha anca DUMAS, 1930, Microzoaires, Planche XXIX, fig. 3 (Fig. 2390). Remarks:
Without description. France.

Oxytricha arcuata DUMAS, 1930, Microzoaires, p 54 (Fig. 239d, e). Remarks: Bog in
France,
 Oxytricha 247

Oxytricha barbula DUMAS, 1930, Microzoaires, p 55 (Fig. 239k). Remarks: Ditch with
grass in France.

Oxytricha becciformis DUMAS, 1929, Microzoaires, p 72 (Fig. 2361). Remarks: In the leg-
end to the figure (planche XXVII, fig. 15) designated as Oxytricha labiata. Balaine,
France.

Oxytricha bilobata FROMENTEL, 1876, Etudes microzoaires, p 264, Planche XII, fig. 6.
Remarks: Very likely a fragment of a hypotrichous ciliate. Never(?) mentioned in the lit-
erature since.

Oxytricha cornipes DUMAS, 1929, Microzoaires, p 70 (Fig. 236d). Remarks: Rather

surely a small fragment of a hypotrich. Balaine, France.

Oxytricha corn uta DUMAS, 1929, Microzoaires, p 73 (Fig. 236n). Remarks: Oxytricha
cornuata in the legend to the figures (Planche XXVII, fig. 18) is an incorrect original
spelling. Balaine, France.

Oxytricha curta DUMAS, 1929, Microzoaires, p 72 (Fig. 236k). Remarks: Pond with cress,
Balaine, France.

Oxytricha cypris DUMAS, 1929, Microzoaires, p 74 (Fig. 243t). Remarks: Certainly a

fragment. France.

Oxytricha decumana PERTY, 1852, Mitt. naturf. Ges. Bern, 1852: 64. Remarks: No illus-
tration available. I assume that PERTY (1852a) was published before PERTY (1852b),
where O. decumana is described on page 154. Freshwater, Switzerland.

Oxytricha deformis FROMENTEL, 1876, Etudes microzoaires, p 267, Planche XIII, fig. 8.
Remarks: FROMENTEL (1876) selected an appropriate species name: very likely a de-
formed specimen.

Oxytricha dubia O. F. M. - KOVALEVA & GOLEMANSKY, 1979, Acta Protozool., 18: 275.
Remarks: Mentioned in a list of psammobiotic ciliates of the Black Sea. I could not find
the basionym.

Oxytricha dujardiniana DIESING, 1866, Sber. Akad. Wiss. Wien, 53: 95. Remarks: Estab-
lished for 0. gibba sensu DUJARDIN (1841, p 418, Planche XI, fig. 12), which is, however,
rather certainly a species indeterminata.

Oxytricha ehrenbergiana DIESING, 1866, Sber. Akad. Wiss. Wien, 53: 95. Remarks: It is
very difficult to fmd out for which population this species was established. From the list
of synonyms in DIESING (1850, P 157) and DIESING (1866b), I conclude that the limnetic
248 SYSTEMATIC SECTION

Oxytricha gibba sensu EHRENBERG (1838, Infusionsthierchen, p 365, Tafel XLI, II) was
meant. However, this form is rather superficially described, so that an identification will
never be possible.

Oxytricha ephippioides DUMAS, 1929, Microzoaires, p 74 (Fig. 243r). Remarks: Certainly

a fragment. France.

Oxytricha exociformis DUMAS, 1930, Microzoaires, p 56 (Fig. 2391). Remarks: Possibly

an Uroleptus-like species because ventral rows are present. Marshy ditch in France.

Oxytrichafimbriata DUMAS, 1930, Microzoaires, Planche XIV, fig. 7 (Fig. 239b, c). Re-
marks: In the text (p 53) designated as "Oxytricha jimbriata, DE FROMENTEL". This
author, however, redescribed Trichoda fimbriata MOLLER, 1786, which is the anterior
portion of Stylonychia mytilus (EHRENBERG 1838, P 371).

Oxytrichaformosa ALEJ<PEROV, 1984, Zool. Zh., 63: 1461 (Fig. 223m, n). Remarks: Infra-
ciliature unreliable; very likely some cirri were not recognised with the wet silver impreg-
nation method. Freshwater from Azerbaijan.

Oxytricha fromenteli FOISSNER, 1987, Arch. Protistenk., 133: 223. Remarks: Oxytricha
fromenteli is the replacement name for O. ovalis FRoMENTEL, 1876, Etudes microzoaires,
p 264, Planche XII, fig. 2, which is a junior primary homonym of 0. ovalis SCHMARDA,
1854. Oxytrichafromenteli is presumably a Steinia or Cyrtohymena species because the
buccal area is very large. Oxytricha ovalis, DE FROMENTEL sensu DUMAS (1929, p 70) is a
redrawing from the original description (Fig. 236c).

Oxytrichafurcatus SMITH, 1897, Am. mono microsc. J., 18: 147 (Fig. 220e). Remarks:
"Orytrichafurcatas" in the legend to the figure of the original description is an incorrect
original spelling. KARL (1932, P 602) - who considered it as a valid species - made a jus-
tified emendation: Oxytricha furcata. Thus, his own species Oxytricha (I'achysoma) fur-
cata KARL, 1932, is a junior primary homonym which has to be replaced. The very
prominent adoral zone of membranelles and the obviously straight undulating membranes
would require a classification in Stylonychia. On the other hand, the very soft and flexible
body is typical for members of Oxytricha. In life 127-170 Ilm long, elliptical, both ends
rounded. Two ellipsoidal macronuclear nodules. Contractile vacuole about in mid-body.
Arrangement of frontal and ventral cirri as in species of the Stylonychia mytilus complex,
but all cirri, except the frontal cirri, furcated. Five transverse cirri, funbriated at their
ends, only the left transverse cirrus does not protrude beyond posterior end of cell. Mar-
ginal rows "continuous, but heavier and longer posteriorly", indicating that caudal cirri
present. SMITH (1897b) found this, in my opinion, doubtful form in an old infusion of rose
fission petals (USA). PATRICK (1961) and PATRICK et al. (1967) recorded it from rivers in
the USA and DINGFELDER (1962; Fig. 224c) from a puddle in Bavaria.
 Oxytricha 249

Oxytricha galeata DUMAS, 1929, Microzoaires, p 75 (Fig. 243s). Remarks: Certainly a

fragment. France.

Oxytricha gyrinioides DUMAS, 1929, Microzoaires, p 74 (Fig. 243q). Remarks: France.

Oxytricha hengshanensis Lru et aI., 1992 in SHEN, Lru, SONG & Gu, 1992, Subtropical
soil animals of China, p 155 (Fig. 2401, m). Remarks: I have some doubt that the ventral
infraciliature is correctly drawn. A micrograph of a protargol-impregnated, slightly dam-
aged specimen, kindly supplied by SHEN YUNFEN, shows that it is possibly a Cyrtohymena
species because the undulating membranes are distinctly bent. The anteriorly displaced
''postoral ventral cirri" are perhaps an artefact. In spite of these uncertainties, a short de-
scription (based on a translation by Mr Wu DONGLI): in life (?) 124-153 x 53-76 f.1m.
Body ovoid, both ends rounded, flexible. Two macronuclear nodules. Contractile vacuole
about in mid-body. Cytoplasm colourless, cortical granules lacking. Many food vacuoles
with algae and amoebas. Rapid movement. Adoral zone of membranelies 113 of body
length, 23 adoral membranelles. 3 enlarged frontal cirri, 1 buccal cirrus, 3 longitudinally
arranged cirri, I cirrus behind right frontal cirrus and at level of anterior end of right mar-
ginal row. 3 ''postoral'' ventral cirri longitudinally arranged right of the proximal portion
of adoral zone of membranelles (Fig. 2401, arrow). 2 pretransverse ventral cirri and 5 al-
most horizontally arranged transverse cirri. Left marginal row longer than right one (pos-
sibly including caudal cirri). 3 dorsal kineties. Locus classicus is a subtropical soil on the
Hengshan Mountain, Hunan, China.

Oxytricha immemorata ALEKPEROV, 1984, Zool. Zh., 63: 1460 (Fig. 224a, b). Remarks:
Infraciliature unreliable; possibly some cirri were not recognised with the wet silver im-
pregnation method. Freshwater from Azerbaijan.

Oxytricha incrassata DUJARDIN, 1841, Zoophytes, p 418, Planche XI, fig. 14. Remarks:
Probably indeterminable.

Oxytricha labiata DUMAS, 1930, Microzoaires, p 55 (Fig. 239j). Remarks: A junior pri-
mary homonym of the next species; because it is a species indeterminata, I do not replace
the name. Bog in France.

Oxytricha labiata FROMENfEL, 1876, Etudes microzoaires, p 264, Planche XII, fig. 1. Re-
marks: Likely a deformed specimen. A senior primary homonym of O. labiata DUMAS.

Oxytricha lacerata DUMAS, 1929, Microzoaires, p 71 (Fig. 236g). Remarks: Freshwater

in France.

Oxytricha lacrimula DUMAS, 1929, Microzoaires, p 71 (Fig. 236f). Remarks: Freshwater

in France.
250 SYSTEMATIC SECTION

Oxytricha lata STERK!, 1878, Z. wiss. ZooL, 31: 44, 55 (Fig. 224d). Remarks: STERK!
(1878) mentioned and drew only the fan-shaped adoral zone of membranelIes.

Oxytricha lingua DUJARDIN, 1841, Zoophytes, p 418, Planche XI, fig. 11. Remarks: In life
about 125 J.lm long. Presumably cortical granules present. Stagnant freshwater in Paris,
France. Very likely not an Oxytricha, perhaps a Holosticha or Urostyla.

Oxytricha longipes DUMAS, 1930, Microzoaires, p 55 (Fig. 239i). Remarks: Roadside

ditch on a meadow in France.

Oxytricha luteolucens DUMAS, 1937, Microzoaires, p 20 (Fig. 239q). Remarks: Only 1

macronuclear nodule. France.

Oxytricha merula FROMENTEL, 1876, Etudes microzoaires, p 269, Planche XX, fig. 8. Re-
marks: It is uncertain if the specimen observed is a hypotnchous ciliate at all.

Oxytricha monstrosa VUXANOVICI, 1963, Studii Cerc. BioI., 15: 214 (Fig. 2231). Remarks:
In life 75-85 J.lm long. Body rigid, not contractile, hence very likely no Oxytricha; the
overall view is reminiscent of a Histriculus. Frontal cirri about 13 J.lm long. 9 left and 11
right marginal cirri. Stagnant freshwater in Bucharest, Romania. STILLER (1974a, P 133)
transferred this superficially described form to Tachysoma (T. monstrosa) because the
marginal rows are widely separated posteriorly and no caudal cirri are drawn.

Oxytricha nova KLOBUTCHER, SWANTON, DONINI & PRESCOTT, 1981, Proc. natn. Acad. Sci.
U.S.A., 78: 3015 (Fig. 235A, B). Remarks: This name is a nomen nudum (lCZN 1985, Ar-
ticle 13) because the publication is accompanied neither by a description or a definition
that states characters that are purported to differentiate the taxon, nor by a bibliographic
reference to such a published statement. Only very few morphological data of this organ-
ism are known (2 macronuclear nodules, 4 micronuclei, Fig. 235A). The scanning elec-
tron micrographs available (Fig. 235B, Figure 1 in AMMERMANN 1985, Figure 1b in STEIN-
BROCK 1986 and SCHLEGEL & STEINBROCK 1986) do not allow an identification to species
or genus level. The straight to slightly curved paroral and the long adoral zone of mem-
branelles (almost 50 % of body length) indicate that O. nova belongs to the Stylonychi-
nae (and not to the Oxytrichinae), a classification supported by biochemical investiga-
tions (SCHLEGEL 1985, SCHLEGEL & STEINBROCK 1986, SCHLEGEL et al. 1991). The bio-
chemical data also indicate that 0. nova is closely related to the "Oxytricha hi/arid' of
the Italian protozoologists, a form which I (preliminary) classifY as Stylonychia pustulata.
Recently, we recognised that 0. nova is a species of the Sterkiella histriomuscorum
group (FOISSNER & BERGER 1999; see this paper for further details). Oxytricha nova is the
subject of many molecular-biological studies: ALLEN & OLINS (1984), BAROIN ToURAN-
CHEAU et aL (1995), BERCHTOLD et aL (1995), BERNHARD et aL (1995), BOSWELL et aL
(1982, 1983), BUTLER et al. (1984), DELGADO et al. (1988, 1991), DUBOIS & PRESCOTT
 Oxytricha 251

(1995), ELWOOD et al. (1985), FANG & CECH (1993b, 1995), FANG et al. (1993), GAJADHAR
et al. (1991), GOTISCHLING & CECH (1984), GOTISCHLING & ZAKIAN (1986), GRAY et al.
(1991), GREENWOOD et al. (1991a, b), GRESLIN et al. (1988, 1989), GUNDERSON et al.
(1995), HARPER & JAHN (1989), HERRICK (1992), HICKE et al. (1990, 1995), HOFFMAN &
PRESCOTI (1966, 1997a, b), JAHN (1988), JAHN et al. (1988), KLoBurCHER (1987),
KLoBurcHER & JAHN (1991), KLOBurCHER & PREsCOTI (1986), KLoBurcHER et al. (1984,
1986, 1988), KRAur et al. (1986), KRIsHNAN et al. (1990), LAPORTE & THOMAS (1997,
1998), LEE & KUGRENS (1992), LINGNER et al. (1994), LIPPS et al. (1982), MARTIN &
GRIMES (1987), MELEK et al. (1994), MITCHAM et al. (1992, 1994), OKA & THOMAS (1987),
OLINS et al. (1993), PRESCOTI (1984, 1992a, b, 1993, 1994), PRESCOTI & DuBoIS (1996),
PRESCOTI & GRESLIN (1992), PRESCOTI et al. (1982), PRICE & CECH (1987, 1989), RAGAN
et al. (1996), RAGHURAMAN & CECH (1989, 1990), RAGHURAMAN et al. (1989), RAIKov
(1989), RIBAS-APARICIO et al. (1987), SAPRA et al. (1985), SCHLEGEL (1991), SHIPPEN et al.
(1994), SOOIN et al. (1986), STEINBROCK (1990), SWAGER et al. (1990), SWANTON et al.
(1980a, b, 1982), WILLIAMSON et al. (1989), WILLIS et al. (1993), ZAHLER & PREScOTI
(1988, 1989), ZAHLER et al. (1991). Complete small subunit rRNA sequence 1771 nucleo-
tides long; G * C content 45 % (LEIPE et al. 1994). LAUGHLIN et al. (1983) described a
method for the large-scale cultivation of Oxytricha nova. Before 1978, PRESCOTI and co-
workers studied a rather similar organism which was originally referred to as Stylonychia
mytilus prior to being identified as Oxytricha sp. in LAUTH et al. (1976; further details on
the confusing "nomenclature" of the organisms used by the American workers, see
KLOBurCHER & PRESCOTI 1986, P 113).

Oxytricha oblonga DUMAS, 1929, Microzoaires, p 72 (Fig. 236j). Remarks: Oxytricha ob-
longua in the legend to Planche XXVII is an incorrect original spelling. Posterior end
with 4 cirri. Balaine, France.

Oxytricha oblongatus - SHARMA, SOBTI & KA1HURIA, 1986, Res. Bull. Panjab Univ. Sci.,
37: 119. Remarks: I do not know the original description of this "species". Possibly it is a
numen nudum. In life 80-100 J.1m x 40-50 J.1m. 2 macronuclear nodules. India.

Oxytricha obtusa DUMAS, 1930, Microzoaires, p 53 (Fig. 239a). Remarks: Ventral sur-
face covered with small cirri; thus surely not an Oxytricha. France.

Oxytricha oltenica LEPSI, 1965, Protozoologie, p 972, 975 (Fig. 224e). Remarks: Perhaps
a Stylonychia? Romania?

Oxytricha ovata DUMAS, 1929, Microzoaires, Planche XXVII, fig. 6 (Fig. 236p) and Du-
MAS (1930; as "Oxytrica ovatd') Planche XII, fig. 6 not fig. 11 (Fig. 236q). Remarks: A
senior primary homonym of Oxytricha (Opisthotricha) ovata KAHL, 1932. Without de-
scription. Only 1 macronuclear nodule. France.
252 SYSTEMATIC SECTION

Oxytricha ovalis SCHMARDA, 1854, Denkschr. Akad. Wiss., Wien, 7: 17,25 (Fig. 222h).
Remarks: In life 87-104 J.lm long. Ovoid. Ephemeral pond near Luxor, Egypt. I agree
with FOISSNER (1987d) that this species is so insufficiently described that an identification
is impossible.

Oxytricha (?) parvula VUXANOVICI, 1963, Studii Cerc. BioI., 15: 214 (Fig. 223i, j). Re-
marks: About 40 J.lm long, oval. Adoral zone of membranelIes 112 of body length. Stag-
nant freshwater in Bucharest, Romania.

Oxytricha pisciunculiformis DUMAS, 1929, Microzoaires, p 75 (Fig. 243u). Remarks:

Likely an Uroleptus. France.

Oxytricha pistilloiaes DUMAS, 1929, Microzoaires, p 71 (Fig. 236h). Remarks: Very

poorly described. Balaine, France.

Oxytricha pleuronectes DUMAS, 1929, Microzoaires, p 73 (Fig. 2360). Remarks: Possibly

a fragment of a hypotrich. France.

Oxytricha plicata EICHWALD, 1852, Bull. Soc. imper. nat. Moscou, 25: 518, Tafel VI, fig.
14. Remarks: Indeterminable, perhaps a euplotid. Baltic Sea

Oxytricha praeceps DUMAS, 1930, Microzoaires, p 54 (Fig. 239f, n, u). Remarks: The
body shape is reminiscent of that of Holosticha pullaster, which has, however, the con-
tractile vacuole distinctly behind mid-body. Bog and river in France. In DUMAS (1937, p
20) again designated as new species. "Oxytricha maeceps" in the legend to the figure
(Planche V, fig. 2) is an incorrect subsequent spelling.

Oxytricha proboscis DUMAS, 1930, Microzoaires, p 56 (Fig. 239r, s). Remarks: In wet
mosses (Hypnum) in France.

Oxytricha protensa PERTY, 1852, Mitt. naturf. Ges Bern, 1852: 64. Remarks: PERTY
(1852a) mentioned only the name. An insufficient description is given by PERTY (1852b,
Kenntnis kleinster Lebensformen, p 153, Tafel VI, Fig. 20A-E). Very likely not an Oxy-
tricha because of the conspicuously elongated body. Freshwater in Switzerland.

Oxytricha quadrinucleata CULBERSON, 1986, Diss. Abstr. Int., 46B: 2589. Remarks: Very
likely, CULBERSON (1986) is the author of this nomen nudum, that is, the name is pub-
lished without description (possibly available in his dissertation).

Oxytricha quercineti LEPSI, 1948, Notat. bioI., Buc., 6: 154. Remarks: LEPSI (1948a) pro-
vided a long description but no illustration of this soil species. In life 100 x 40 J.lm. Body
shape similar to that of Tachysoma pellionel/um, elliptical, left margin not concave. Very
flexible. Frontal cirri enlarged. 2-3 frontoventral and 3 postoral ventral cirri. Caudal cirri
 Oxytricha 253

lacking. Marginal rows not interrupted posteriorly. 5 transverse cirri, distinctly protruding
beyond posterior end. Dorsal cilia short. Adoral zone of membranelIes narrow, 30-35 /lm
long. Peristomial lip anteriorly not conspicuously bent. Undulating membranes short. 2
macronuclear nodules. Cytoplasm rather colourless, without symbiotic algae. Feeds on
bacteria. Lively movement. Polysaprobic. Litter of an oak forest near Harbovat,
Romania. I do not believe that this species can be identified. However, it is not impossi-
ble that this particular biotope contains some characteristic species.

Oxytricha radians DUJARDIN, 1841, Zoophytes, p 420, Planche XI, fig. 16. Remarks: In
life 50 /lm long. Possibly an oligotrichid or a euplotid ciliate.

Oxytricha rostrata DUMAS, 1930, Microzoaires, p 56 (Fig. 239m). Remarks: In the legend
to the figure (planche XXVI, fig. 20) designated as 0. cultriformis. Bog and ditch with
plants in France.

Oxytricha saltans VUXANOVICI, 1963, Studii Cerc. BioI., 15: 214 (Fig. 2320). Remarks:
STILLER (1974a, P 133) transferred it to Tachysoma (T. saltans) because the marginal
rows are widely separated posteriorly. BORROR (1972a) synonymised it with Actinotricha
saltans COHN, 1866, which he also transferred to Tachysoma. Oxytricha saltans VUXANO-
VICI is somewhat reminiscent of Tachysoma humicola, which is, however, restricted to
terrestrial habitats. In life 40-55 /lm long, not contractile but flexible. Adoral zone of
membranelles 113 of body length, consists of 9 or 10 membranelles. 5 transverse cirri, in
life 20 /lm long, protrude distinctly beyond posterior end of cell. 8-9 left and right mar-
ginal cirri, respectively, in life 10 /lm long. Lake "Floreasca" in Bucharest, Romania.
VUXANOVICI (1963) found it also in a freshwater habitat in the botanical garden ofBucha-
rest.

Oxytricha stratiformis DUMAS, 1930, Microzoaires, p 54 (Fig. 239h). Remarks: Roadside

ditch on a meadow in France.

Oxytricha striata SCHMARDA, 1854, Denkschr. Akad. Wiss., Wien, 7: 17,24 (Fig. 222i).
Remarks: In life 208 /lm long, both ends rounded. Ephemeral stagnant freshwater near
Luxor, Egypt. Reminiscent of Tachysoma pellionellum, which is, however, distinctly
smaller.

Oxytricha subcylindrica DUMAS, 1929, Microzoaires, p 71 (Fig. 236i). Remarks: Anterior

end with 6-8 enlarged cirri, posterior end with 5 cirri. Balaine, France.

Oxytricha tetranucleata NEMETH in GELEI, 1950, Acta bioI. hung., 1: 80 (Fig. 240g). Re-
marks: The reference ''NEMETH 1950" in GELEI is given without source; possibly this is an
unpublished paper. Only the figure is available. Generic classification uncertain. Hun-
gary. The same is true for the second NEMETH species described in GELEI (1950), namely,
254 SYSTEMATIC SECTION

Holosticha oxytrichoidea (= Holostischa oxytrichoidea in the legend; Fig. 222a), which

is probably an oxytrichid because it has only 18 frontal, ventral, and transverse cirri.

Oxytricha trifallax GRESLIN, PRESCOTI', OKA, LoUKIN & CHAPPELL, 1989, Proc. natu. Acad.
Sci. U. S. A., 86: 6264 and Oxytricha trifallax HUNTER, WILLIAMS, CARTINHOUR &
lIERR.ICK, 1989, Genes & Development, 3: 2110. Remarks: I am not sure if these are the
"original descriptions"; both do not contain morphological data, thus a nomen nudum
(lcZN 1985). The author's got it from R lIAMMERSMTIlI, Ball State University. Further lit-
erature: DOAK et aI. (1997), DUBOIS & PRESCOTI' (1995), HOFFMAN & PRESCOTT (1997a, b),
KLOBUTCHER & lIERR.ICK (1997), LINGNER et aI. (1994), PRESCOTI' & DUBOIS (1996), SEEG-
MILLER & HERRICK (1998), SEEGMILLER et aI. (1997), WILLIAMS et aI. (1993), WITIIERSPOON
et al. (1997). SEEGMILLER et al. (1996) collected wild 0. trifallax cells from diverse lim-
netic sites in Indiana, USA. Recently we found that O. trifallax belongs to the Sterkiella
histriomuscorum group (details see ForSSNER & BERGER 1999).

Oxytricha truncata VUXANOVICI, 1963, Studii Cerc. BioI., 15: 212 (Fig. 222o-t).
Remarks: According to Article 45 (g) of the IcZN (1985) the forms Oxytricha truncata f.
piriforme VUXANOVICI, 1963 (p 213; Fig. 222s, t) and Oxytricha truncata f. dUatata Vux-
ANOVICI, 1963 (p 213; Fig. 222q, r) are of infrasubspecific rank. BORROR (1972a) syn-
onymised this rather superficially described species with O. fallax because the body
shape and the size are very similar. I do not accept this synonymisation because VUXANO-
VICI (1963) mentioned a granulation and a brown-reddish colour, perhaps similar to in O.
aeruginosa. However, in this species the body is not as broad and the transverse cirri do
not protrude beyond the posterior end of the cell. In life 100-150 !lm, "formapiriforme"
100-120 !lm, "forma dUatatd' 80-100 !lm long; only about 1.6 times as long as broad.
Oval or distinctly pyriform (Fig. 222o-q, s). Two macronuclear nodules. Micronuclei not
mentioned. Contractile vacuole about in mid-body. Cells packed with granules and small
globules giving them a rusty colour, "forma piriforme" appears reddish. Adoral zone of
membranelies 40-45 % of body length. Arrangement and number of frontal and ventral
cirri certainly not exactly observed. 5 transverse cirri. Dorsal cilia 2-4 !lm long. Locus
classicus is the Lacul Fundeni, a pond in Bucharest, Romania, where VUXANOVICI (1963)
discovered it between decaying plant material. Not found since.

Oxytricha viridis FROMENTEL, 1876, Etudes microzoaires, p 266, Planche XIII, fig. 5. Re-
marks: LONGHI (1895) found 0. viridis FROMENTEL in a lake in Italy.

EHRENBERG (1831, P 119; 1838, p 366) transferred Trichoda cicada MOLLER, 1786 (now
Aspidisca cicada) to Oxytricha.
 Oxytricha 255

Insufficient redescriptions

Actinotricha saltans COHN - ALzAMORA, Notas Resfun. Inst. esp. Oceanogr., 2: l3 (Fig.
89a). Remarks: Poor drawing. Bay of Palma de Mallorca, Mediterranean.

Actinotricha saltans COHN - DRAGEsco, 1963, Cab. BioI. mar., 4: 265 (Fig. 89d).
Remarks: Some characters (length of right marginal cirral row, number of transverse
cirri) of this redescription, which is obviously exclusively based on silver impregnated
material, do not agree very well with the authoritative redescription. Saprobic sediment
inlnear Roscoff, France, Atlantic Ocean.

Actinotricha saltans COHN - LEPSI, 1927, Studii Cerc. Acad. RPR, 12: 124 (Fig. 89b, c).
Remarks: Poor drawing, although the distinctly shortened right marginal cirral row and
the radiating distal adoral membranelles indicate that the LEPSI identification is correct. In
life 102 x 34 J.lm. Black Sea? LEPSI (1926a; without illustration) population in life
162 x 54 J.lm, that is, too large; Black Sea.

Opisthotricha parallela ENGELMANN - SCHUSTER, 1899, Sci. Gossip, 5: 203 (Fig. 240i).
Remarks: Freshwater in England.

Opisthotricha parallelis ENG. - KAlIL, 1928, Arch. Hydrobiol., 19: 211 (Fig. 52d). Re-
marks: KAlIL (1928b) gives only an illustration and no description of this form. It has 2
buccal cirri, indicating that it is not 0. parallela. In life about 100 J.lm long and three
times as long as broad. Both ends rounded. Contractile vacuole slightly in front of mid-
body. Adoral zone of membranelIes about 113 of body length. Frontal cirri distinctly en-
larged. Two (!) buccal cirri near anterior end of undulating membranes. 5 transverse cirri,
protrude slightly beyond posterior end of cell. Caudal cirri obviously present, however,
number not discernible. Dorsal cilia short (2-4 J.lm). KARL (1928a, b) found this ciliate in
the Oldesloer Salzstellen in a small trench (salinity 0.29 %, containing numerous Rhodo-
bacteriaceae) near Alt-Fresenburg near Hamburg, Germany.

Opisthotricha procera KAlIL - NAlDu, 1965, Hydrobiologia, 25: 560 (Fig. 224k). Re-
marks: Arrangement of ventral cirri unreliable. Among algae in an old well in Nakash,
Cuddapah, India.

Opisthotricha similis ENGELMANN, 1862 - CURDS, 1969, Wat. Poll. Res., No. 12: 68 (Fig.
22lh). Remarks: The drawing is rather schematic and does not agree with the authorita-
tive redescription by KAlIL (1932; Fig. 57a).

Opistotricha ovata KAlIL 1930 - BIERNACKA, 1967, Wiss. Z. Ernst Moritz Amdt-Univ.
Greifswald, 16: 244 (Fig. 226e). Remarks: Incorrect subsequent spelling of Opistho-
256 SYSTEMATIC SECTION

tricha. In life 100-120 /lm long, 3 caudal cirri. Feeds on flagellates. Polysaprobic; among
decaying algae in polluted littoral areas ofthe Hiddensee Island, Baltic Sea.

Oxitricha fallax - BATTISH, 1992, Freshwater zooplankton of India, p 57, Fig. 54. Re-
marks: Incorrect spelling of Oxytricha. Very superficially illustrated. Planktonic in India.

Oxytricha - DUMAS, 1929, Microzoaires, Planche XXXVI, fig. 38 (Fig. 236m). Remarks:
Without description. France.

Oxytricha - MAUPAS, 1888, Archs Zool. expo gen., 6: 222, 276 (Fig. 230a, b). Remarks:
In life 100-190 /lm long, very flexible but only slightly contractile. Arrangement of cirri,
see Figure 230a. Freshwater. Cyst spherical, wall with more or less regularly arranged
thickenings (Fig. 230b). Although this population is well described, I could not identify it
with any species. The overall morphology is reminiscent of o. fallax, whereas the con-
spicuous cyst indicates that it is perhaps a species of its own.

Oxytricha aeruginosa WRZESNIOWSKI - PATSCH, 1974, Arb. Inst.landw. Zool. Bienenk., 1:

58 (Fig. 220m). Remarks: 100-200 x 40-70 /lm. Slender. 2 macronuclear nodules, 2 mi-
cronuclei. Contractile vacuole at about level of buccal vertex. Marginal rows confluent
posteriorly, marginal cirri about 10 /lm long. Transverse cirri do not protrude beyond
posterior end of cell. Adoral zone of membranelles 113 of body length, undulating mem-
branes slightly bent. 4-6 dorsal kineties of body length. Morphogenesis commences with
the proliferation of the oral primordium very close to the left transverse cirrus. Freshwa-
ter near Bonn, Germany. PATSCH (1974) described no red granulation of the cytoplasm;
thus, the identification with 0. aeruginosa is unjustified. Perhaps identical with Histricu-
Ius histrio because the ventral infraciliature is rather similar and no caudal cirri are de-
scribed. However, since no information is available on the flexibility of the cell, I avoid
synonymisation with H. histrio.

Oxytricha aeruginosa WRZESNIOWSKI, 1870 - AGAMALIEv, 1974, Acta Protozool., 13: 71

and AGAMALIEv, 1983, Ciliates of Caspian Sea, p 108 (Fig. 221a, b). Remarks: About
120-150 /lm long; 30-35 adoral membranelles; 40 right and 38 left marginal cirri; mar-
ginal rows widely open posteriorly; only 3 dorsal kineties and probably no caudal cirri.
Solid surface overgrowth of the eastern coast of the Caspian Sea. The identification is
very doubtfully, since no red colouring is mentioned. Furthermore, AGAMALIEV (1974) de-
scribed only 3 dorsal kineties. Very likely he overlooked some kineties in the wet silver
impregnated material because he (AGAMALIEV 1983, P 108ft) described 3 kineties also in
O. marina, Opisthotricha halophila, and Stylonychia mytilus which is, of course, a mis-
observation.

Oxytricha chlorigella KAHL - HORVATH, 1939, Allatt. Kozl., 36: 83 (Fig. 22li). Remarks:
Incorrect subsequent spelling of Oxytricha chlorelligera. HORVATH (1939b) did not men-
 Oxytricha 257

tion the symbiotic algae, which are characteristic for the species. Furthermore, there are
some differences in the arrangement of the postoral ventral and transverse cirri. Reminis-
cent of 0. hymenostoma.

Oxytricha crassistilata KAHL, 1932 - ALEKPEROV, 1993, Zoosystematica Rossica, 2: 26

(Fig. 241a, b). Remarks: In life 25-30 Ilm long. Cytoplasm transparent. 2 bean-shaped
macronuclear nodules, each with 1 micronucleus. 25-30 adoral membranelles. 10 ventral
cirri (buccal, frontoventral, postoral ventral, pretransverse ventral), 5 transverse cirri, 12
right and 10 left marginal cirri. 4 dorsal kineties of body length, 1 shortened kinety, 3
caudal cirri. Soils of st. Petersburg and its suburbs, Russia. Does not match the original
description in many characters (size, nuclear apparatus ... ).

Oxytrichafallax - BARKER, 1949, J. Proc. Inst. Sew. Purif., year 1949: 11 (Fig. 232i). Re-
marks: Only 86 Ilm long! Sewage treatment plant.

Oxytrichafallax - BISHOP, 1943, J. Morph., 72: 443 (Fig. 221t). Remarks: I do not be-
lieve that BISHOP (1943) worked with o. fallax because the outline does not show the
typical pyriform shape.

Oxytrichafallax - GREGORY, 1923, J. Morph., 37: 555 (Fig. 240n). Remarks: Neithersize
(80-130 Ilm) nor body shape allow an unequivocally identification with O. fallax. Fur-
thermore, the arrangement of the ventral cirri is insufficiently observed.

Oxytricha fallax STEIN - HEBERER, 1928, Treubia, 10: 27 (Fig. 229h). Remarks: HEBERER
wrote that his form is probably identical with O. fallax. He observed two cells bursting
and the release of many spheres of different size; he interpreted this process as mUltiple
division without encystment and designated the spheres as anisogametes. Freshwater in
West-Lombok, Indonesia.

Oxytricha fallax - QUENNERSTEDT, 1865, Acta Univ. lund., 2: 59 (Fig. 38k). Remarks:
Sweden.

Oxytrichafallax - SMITH, 1978, Br. Antarct. Surv. Sci. Rep., No. 95: 75 (Fig. 221e). Re-
marks: Soil of various Antarctic islands. Oxytricha fallax is not reliably recorded from
terrestrial habitats (FOiSSNER 1987a). Perhaps Sterkiella histriomuscorum.

Oxytricha fallax, ST. - KOFFMAN, 1926, Acta zoo1., Stockh., 7: 3 10 (Fig. 231 d). Remarks:
Garden soil near Stockholm, Sweden. Oxytrich fallax rather certainly does not inhabit
terrestrial habitats (FOISSNER 1987a).

Oxytricha fallax STEIN - CONN, 1905, Bull. Conn. St. geol. nat. Hist. Surv., 2: 60 (Fig.
221 c). Remarks: Number and arrangement of cirri rather doubtful. Furthermore, the body
258 SYSTEMATIC SECTION

shape does not match the characteristic fallax type. Freshwater habitats in Connecticut,
USA.

Oxytricha fal/ax STEIN - WEST & LUNDIN, 1963, Pap. Mich. Acad. Sci., 48: 108 (Fig.
220k). Remarks: See also LUNDIN & WEST (1963) and WEST (1953). Freshwater habitats
of the Upper Peninsula, Michigan, USA.

Oxytrichafal/ax STEIN - CHARDEZ, 1967, Revue Ecol. BioI. Sol, 4: 294 (Fig. 223p). Re-
marks: Possibly identical with O. proximata (a species also mentioned in his faunal list!),
which is originally described from terrestrial habitats and has a similar body shape and
size as O. fal/ax. Soil in Belgium.

Oxytrichafallax STEIN - FORMISANO, 1957, AnnaIi Fac. Sci. agr. Univ. Napoli, 22: 26
(Fig. 234b). Remarks: Rhizosphere of cultivated plants in the Campania soils, Italy.

Oxytrichafallax STEIN, 1859 - CELA, 1972, Physis, 31: 576 (Fig. 232n). Remarks: In life
97-160 x 3~8 J.l.m (mean = 129 x 52J.1.m; n = 5). In the Yalca Lagoon, Argentina, on
Azolla filiculoides.

Oxytricha furcata SMI1H 1897 - DINGFELDER, 1962, Arch. Protistenk., 105: 621 (Fig.
224c). Remarks: In life 91-116 J.l.m long. Elliptical. About 30 right and 20 left marginal
cirri; marginal rows converging posteriorly. About 35 adoral membranelles. 4 transverse
cirri, protrude distinctly beyond posterior end of cell. Arrangement of ventral cirri unreli-
able; especially the longitudinal row behind the buccal vertex is probably a misobserva-
tion. Infusion of mud from a puddle, Bavaria, Germany.

Oxytricha halophila KAHL, 1932 - AGAMALlEv, 1978, Acta Protozool., 17: 439 and AGA-
MALIEV, 1983, Ciliates of Caspian Sea, pliO as Opisthotricha halophila KAHL, 1932 (Fig.
2331, m). Remarks: In life 160-180 x 40-50 J.l.m. Two macronuclear nodules, 2 or 3 mi-
cronuclei. Contractile vacuole about at level of buccal vertex. Live specimens yellowish-
white. Adoral zone of membranelles 1/3 of body length, about 34-38 adoral mem-
branelles. Distal end of adoral zone conspicuously overlapping to right side of cell.
About 28-35 cirri in each marginal row, rows distinctly separate posteriorly. Three dorsal
kineties of body length (misobservation). Caudal cirri? Fine sand, algae detritus, and peri-
phyton of the Caspian Sea (see also AGAMALIEV 1971). The identification is certainly in-
correct because Oxytricha halophila sensu KAHL (1932) has only a single micronucleus
and prominent caudal cirri (Fig. 84a, b).

Oxytricha halophila KAHL, 1932 - CZAPIK & JORDAN, 1976, Acta Protozool., 15: 441, 442
(Fig. 232m). Remarks: The characteristic single micronucleus is not described; thus, the
identification cannot be accepted. Brackish water in the Gdansk Bay (Baltic Sea) at a sa-
linity of 0.3-0.7 %.
 Oxytricha 259

Oxytricha henegui F.-F., 1924 - MATIs, 1977, Acta Fac. Rerum nat. Univ. comen.
BratisI., 22: 25. Remarks: Possibly erroneously written for Opisthonecta hennegui
FAURE-FREMIET, 1924, a peritrichous ciliate (see ForssNER et aI. 1992c).

Oxytricha hymenostoma STOKES - CONN, 1905, Bull. Conn. st. geoI. nat. Hist. Surv., 2:
60 (Fig. 221 d). Remarks: Freshwater habitats in Connecticut, USA.

Oxytricha /udibunda STOKES, 1891 - CURDS, 1969, Wat. Poll. Res., No. 12: 67 (Fig.
221g). Remarks: The drawing is rather schematic and does not allow unequivocal identi-
fication. CURDS (1969) assumed that the posteriorly displaced macronuclear nodules can
be used to separate this species from congeners like O. fallax.

Oxytricha ludibunda STOKES - NAIDu, 1965, Hydrobiologia, 25: 559 (Fig. 224m). Re-
marks: In life 90--103 x 42-46 ~m. Numerous in dirty water in Cuddapah, South India.

Oxytricha marina KAHL 1930 - BIERNACKA, 1967, Wiss. Z. Ernst Moritz Arndt-Univ.
Greifswald, 16: 244 (Fig. 226d). Remarks: In life about 120 ~m long, flexible, rapid
movement. Feeds on diatoms. Polluted areas of the littoral of the Hiddensee Island, Baltic
Sea.

Oxytricha marina KAHL, 1932 - GANAPATI & RAo, 1958, Andhra Univ. Mem. Oceanogr.,
2: 86 (Fig. 228a). Remarks: In life 150--170 ~m long. Caudal cirri lacking, indicating that
they observed a Tachysoma. Sea near Visakhapatnam, India.

Oxytricha minor KAHL, 1932 - CHORIK, 1968, Free-living ciliates, p 133 (Fig. 231h). Re-
marks: CHORIK (1968) did not draw the single micronucleus and the somewhat elongated
marginal cirri at the posterior end of the rows. Small water bodies in Moldova

Oxytricha parvisty/a STEIN - EDMONDSON, 1920, Trans. Am. microsc. Soc., 39: 185. Re-
marks: STEIN (1859b) did not describe an O. parvistyia. I assume that it should read O.
platystoma. EDMONDSON (1920) also recorded Tachysoma parvisty/a STOKES; thus, a con-
fusion with this species must be excluded. Devil's Lake Complex, North Dakota, USA.

Oxytricha pustulata EHRBG. - JAWOROWSKI, 1893, Spraw. Kom. fizyogr., 28: 41.
Remarks: JAWOROWSKI (1893) mentioned this binomen in a faunal list of Krakow, Poland.
Probably he meant Stylonychia pus/ufa/a, which he, however, also listed.

Oxytricha saprobia (KAHL, 1935) - VUXANOVICI, 1963, Studii Cerc. BioI., 15: 215 (Fig.
2201). Remarks: This population has a distinctly longer adoral zone of membrane lies
(46 % of body length) than the population described by KAHL (1932; about 37 %).
Moreover, KAHL (1932) drew prominent transverse cirri and a single micronucleus be-
tween the macronuclear nodules, which is not the case in Figure 2201. Romania.
260 SYSTEMATIC SECTION

Oxytricha sardida ENTz - BULLINGTON, 1925, Arch. Protistenk., 50: 271. Remarks:
120-140 x 40 Ilm. I do not know this species. Perhaps an incorrect spelling of O.
sordida, which is, however, from WRZESNIOWSKIEGO (1866).

Oxytricha similis ENGELMANN - PATSCH, 1974, Arb. Inst. landw. Zool. Bienenk., 1: 61
(Fig. 230h). Remarks: In life 100-150 x 40-50 !lm. Flexible. Adoral zone of membrane1-
les 30-40 Ilm long. Only 3 frontoventral cirri, 3 caudal cirri, 5 dorsal kineties. Brook
near Bonn, Germany. Very likely not identical with 0. similis, which has only 2 caudal
cirri.

Oxytricha sp. - CHAUDHURI, 1929, AnnIs Protist., 2: 54, 57 (Fig. 242a, b). Remarks: Soil
in India.

Oxytricha sp. - CONN, 1905, Bull. Conn. St. geol. nat. Hist. Surv., 2: 60 (Fig. 224i, j). Re-
marks: CONN (1905) found two Oxytricha sp. in freshwater habitats of Connecticut, USA.
Both forms have an unreliable arrangement of ventral cirri.

Oxytricha sp. - CZAPIK, 1959, Acta Hydrobiol., 1: 191 (Fig. 228i). Remarks: Perhaps a
Sty/onychia. Benthic area of fishponds in Poland.

Oxytricha sp. - GRANDORI & GRANDORI, Boll. Lab. Zoo!. agr. Bachic. R 1st. sup. agr. Mi-
lano, 5: 289, Tavola XIII, fig. 269. Remarks: Soil in Italy.

Oxytricha sp. - JACOBSON, 1931, Arch. Protistenk., 75: 76 (Fig. 226i). Remarks: An iden-
tification is impossible because no morphological details (for example, flexibility) are de-
scribed.

Oxytricha sp. - MOTE, 1954, Proc. Iowa Acad. Sci., 61: 588 (Fig. 222e). Remarks: The
supernumerary cirri are reminiscent of Gastrosty/a. Soil of a virgin prairie in Iowa, USA.

Oxytricha sp. - NIKOUUK & GELTZER, 1972, Pocvennye prostejsie SSSR, p 128, Plate XII,
Fig. 228. Remarks: Perhaps Gonostomum sp. Soil from the USSR.
 Allotricha 261

Allotricha STERK!, 1878

1878 Allotricha STERK!, Z. wiss. Zoo!., 31: 37,48,57 - Type (by monotypy): Allotricha mollis STERK!, 1878.
1882 Allotricha, STERK!- KENT, Manual Infusoria II, p 782.
1889 Pleurotricha (Allotricha) STERK! 1878 - BOTsCHLI, Protozoa, p 1747.
1979 Allotricha STERK!, 1878 - CORLISS, Ciliated proto:roa, p 312.
1987 Allotricha STERK!, 1878 - TUFFRAU, AnnIs Sci. nat. (Zoo!.), 8: 116.
1994 Allotricha STERK!, 1878 - TUFFRAU & FLEURY, Traite de Zoologie, 2: 144.
1996 Onychodromopsis STOKES, 1887 - PEn: & FOISSNER, Acta Proto:roo!., 35: 264.
1997 Onychodromopsis STOKES, 1887 - BERGER & FOISSNER, Arch. Protistenk., 148: 146.

C h a r act e r i sat ion: Adoral zone of membranelles formed like a question mark.
Undulating membranes in Oxytricha pattern or rather straight and arranged almost side
by side. Frontoventral cirri in V-shaped pattern. Postoral ventral cirri in dense cluster be-
hind buccal vertex. Two pretransverse ventral and 5 transverse cirri. Two or more right
and I or more left rows of marginal cirri. Caudal cirri present. Primordia V and VI of the
proter originate from cirrus V/4 and V/3 respectively. No parental marginal rows retained
after division. Dorsal morphogenesis in Oxytricha pattern.

T a x 0 nom y: The characterisation is that of Onychodromopsis in BERGER & FOISSNER

(1997), where we followed the PETZ & FOISSNER (1996) definition of Onychodromopsis.
For a detailed description of patterns, see chapter 2 in the general section.
Allotricha was established for a metabolic (that is flexible) hypotrich with typical
Oxytricha cirral pattern but more than 2 (marginal) cirral rows (see below). Thus, BOT-
SCHLI (1889) classified Allotricha as subgenus of P/eurotricha, whereas KAHL (1932,
P 593) mentioned the type species (as A. mollis) in Pleurotricha. BORROR (1972a, P 15)
synonymised Allotricha mollis with Pleurotricha lanceo/ata, which has, however, a rigid
body. CORLISS (1979), TUFFRAu (1987), and TUFFRAu & FLEURY (1994) classified Allotri-
cha in uncertain position in the Sporadotrichina and Oxytrichidae, respectively. Very re-
cently, PETZ & FOISSNER (1996) considered A. mollis as genus and species indeterminata
because the original description lacks details and a figure.
STERKI (1878) made rather exact observations (including morphogenetic ones) and
wrote on p 48 "Der einfacheren Bezeichnung wegen benenne ich die 8 charakteristischen
Stimwimpern, die bei Sty/onychia, Oxytricha, Histrio, P/eurotricha und Allotricha offen-
bar vollkommen homolog sind, mit Zahlen und zwar...". Furthermore, he gave a diagno-
sis on p 59: "Allotricha n. gen. Die Stim-, Bauch- und Afterwimpern der Oxytrichen,
ausserdem rechts zwei Reihen borstenf6rmiger Wimpern (ahnlich den Pleurotrichen).
Klirper metabolisch. Peristom, Consistenz, Aussehen, Far-bung und Bewegungsweise der
Urostylen. 1st also in keinem der bisherigen Genera unterzubringen. A. mollis. Sehr viel
beobachtet; haufig". Unfortunately, STERKI (1878) did not figure the type species. Never-
theless, I reactivate Allotricha because the diagnosis exactly matches the descriptions of
Pleurotricha variabilis REuTER, Onychodromopsisjlexilis sensu PETZ & FOISSNER (1996),
and my nomen nudum species Parurosoma granulifera in FOISSNER et al. (1991, p 262).
262 SYSTEMATIC SECTION

The "Reihen borstenf6rmiger Wimpern" mentioned in the original description are cer-
tainly not misobserved dorsal kineties because STERK! (1878, p 48) was the first to de-
scribe these inconspicuous structures minutely. Further details, see Allotricha mollis.

Key to species

1 Length in life usually less than 150 Jim; transverse cirri conspicuously displaced an-
teriad, protrude only slightly beyond posterior end of cell, hook-shaped arrangement
(Fig. 92a, h); cirrus III12 arranged left of posterior migratory cirrus (cirrus VII3; Fig.
92h, j, arrow); prefers (exclusively in?) terrestrial habitats .... A. antarctica (p 268)
Length in life usually more than 150 Jim; transverse cirri arranged more or less termi-
nally, protrude distinctly beyond posterior end of cell, arranged in oblique row (Fig.
91a, c); cirrus III12 arranged left between level of cirri IV/3 and VIl3 (Fig. 91c, ar-
row); prefers (exclusively in?) freshwater ................................. 2
2 Cortical granules (0.5-1.0 Jim) present (Fig. 91b) ............... A. mollis (p 262)
Cortical granules absent (Fig. 91 s) ............................................ .
. . . . . . . . . . . . . . . . Pleurotricha variabilis, a supposed synonym of A. mollis (p 268)

Allotricha mollis STERK!, 1878 (Fig. 91a-g, t, Tables 3, 19)

1878 Allotricha mollis STERK!, Z. wiss. Zoo!., 31: 48, 58 (no illustration).
1932 Allotricha mollis STERK! - KAHL, Tierwelt DtI., 25: 593 (mentioned at Pleurotricha; no own obser-
vations).
1982 Pleurotricha tihanyiensis (GELLERT & TAMAS, 1958) - HEMBERGER, Dissertation, p 223 (Fig. 91g; un-
published thesis; see taxonomy).
1986 Pleurotricha tihanyiensis (GEL. & TAM). HEM. - SCHMITZ, Dissertation, p 94, Abb. 40.
1988 Pleurotricha spec. - KRAMER, Diplomarbeit, p 29 (Fig. 9lf; unpublished thesis).
1991 Parurosoma granulifera BERGER - FolSsNER, BLATTERER, BERGER & KOHMANN, Informationsberichte
Bayer. Landesamtes fUr Wasserwirtschaft, 1191: 261, 262 (nomen nudum species; Fig. 91c).

T a x 0 nom y: STERK! (1878) provided a short defmition of A. mollis but unfortunately

no illustration. It was transferred to Pleurolricha (KAHL 1932), synonymised with P. lan-
ceolala (BORROR 1972a), or classified as species indeterminata (PETZ & FOISSNER 1996).
Conversely, I consider the characterisation (see relevant chapter in genus section) as suf-
ficient and assume that STERK! observed the same species that I previously called (in
FOISSNER et al. 1991) Parurosoma granulifera because (i) STERK! (1878, P 48, 58) de-
scribed a ventral cirral pattern (only including frontal-ventral-transverse cirri) identical to
in other I8-cirri oxytrichids; my "P. granuliferd' has a typical Oxytricha cirral pattern
(Fig. 9Ic, f). (ii) STERK! mentioned 2 right cirral rows (similar to in pleurotrichs); "P.
granulifera" usually has 2 right marginal rows (Fig. 91c, f). (iii) The body of Allotricha
is metabolic (STERK! 1878), that is, flexible as in "P. granuliferd'; thus, synonymisation
with the rigid Pleurotricha lanceolala, as suggested by BORROR (1972a), is unjustified.
 Allotricha 263

(iv) Peristome, consistence, appearance, colouring, and movement is as in urostylids

(STERKI 1878); all of which matches "Parurosoma granulifera", which was mentioned in
FOISSNER et al. (1991) in the chapter similar species of Paraurostyla weissei, which was
considered as typical urostylid in STERKI'S time!
The HEMBERGER (1982; Fig. 91g) popUlation has distinctly larger frontal-ventral-
transverse cirri than "Parurosoma granulifera" (Fig. 91c) or Pleurotricha spec. sensu
KRAMER (1988; Fig. 91t). Furthermore, in the HEMBERGER population the right transverse
cirrus (VIII) is somewhat displaced anteriorly, indicating that it is possibly a different
species. Unfortunately, neither HEMBERGER (1982) nor KRAMER (1988) mentioned whether
cortical granules are present or not. Pleurotricha variabilis REUTER is classified as sup-
posed synonym of A. mollis (see below for details).
In life, Allotricha mollis can be easily confused with the equally sized Paraurostyla
weissei, also having cortical granules, 2 macronuclear nodules, and additional cirral rows;
however, P. weissei has more transverse cirri and cirral rows and a different arrangement
and number of frontal and ventral cirri.
Most data in the following chapter are own observations (Fig. 91a-e; slides are de-
posited in the Ober5sterreichische Landesmuseum in Linz, Upper Austria). Data from
KRAMER (1988) are indicated, and data from HEMBERGER (1982) are mentioned separately
because they presumably relate to a different species.

M 0 r ph 0 log y and b i 0 log y: In life about 160-220 x 60-80 ~m (original data),

according to SCHMITZ (1986) 180-290 x 75-110 ~m. Flattened about 2-2.5:1 dorso-
ventrally. Long elliptical, usually both ends rounded, posterior one sometimes incon-
spicuously tapered. Very flexible. Two macronuclear nodules in life about 30 x 15 ~m,
nucleoli globular, about 1.5 ~m across. Usually 2 (sometimes 3), 4 ~m sized, ellipsoidal
or spherical micronuclei (however, see Table 19). Contractile vacuole somewhat in front
of mid-body, during diastole with long canals (Fig. 91a-<:). Cortical granules yellowish,
in older cultures almost yellow-red, about 1 ~m across, usually arranged in loose groups
and short rows, especially around cirri and dorsal cilia (Fig. 91b). Pellicle and cytoplasm
colourless. Posterior cell portion packed with cytoplasmic crystals and 2-4 ~m sized
globules. Movement moderately fast gliding.
Adoral zone of membranelles about 35 % of body length, bases of largest mem-
branelles in life 12-14 ~m broad. Buccal area broad and deep, almost as in Cyrtohymena.
Undulating membranes distinctly curved and intersecting, that is, more or less in Oxytri-
cha pattern (Fig. 91c, f, t). Frontal cirri enlarged, in life about 20 ~m long. Frontoventral
and ventral cirri slightly enlarged, rather invariably arranged as in Figure 91c. Transverse
cirri arranged in an oblique row, slightly enlarged, in life 25-30 ~m long, protrude dis-
tinctly beyond posterior end of cell (Fig. 91a, c, f, t). Usually 2 right marginal rows of
body length, a third row is distinctly shorter and confined to mid-body (Fig. 91e), some-
times a fourth right marginal row with 7-19 cirri (SCHMITZ 1986). Invariably 1 left mar-
ginal row. Marginal cirri in life 15-20 ~m long. Six dorsal kineties, including 2 dorso-
marginal rows. Kineties 4--6 shortened anteriorly, kineties 3 and 6 distinctly shortened
264 SYSTEMATIC SECTION

r ,
r,
; (

,
I f
(

r
(

(
{

\ i r
.....
(
(

(}
--.
a
~

...
~

, .. .~

",
I

,
I
"
,
I

,
,"
,

.. •. ••••.
I
I
#• •
, 9

e
. -.. ,,' f
 Allotricha 265

h k m

n o p q r
Fig. 91b-s Pleurotricha variabilis, a supposed synonym of Allotricha mollis (after RwrER 1961. h-s, from
life). b-r) Ventral views of 11 specimens from 4 different clones (h-j, culture 1; k, culture 2; H>, culture 3;
p-r, culture 4). s) Ventral view of a typical specimen, 200-220 ~m. Arrow marks frontoventral cirrus III12;
p268.

posteriorly. Dorsal cilia in life about 3 ~m, according to SCHMITZ (1986) 6-8 ~m long, as
in the HEMBERGER population. Caudal cirri about 15 ~m long, at posterior end of kineties
1, 2, and 4. Very likely usually more than 3 caudal cirri (Fig. 91d), the KRAMER (1988)
and SCHMITZ (1986) populations with 3~ cirri (4.4 on average; n = 9).

-(- Fig.91a-g Allotricha mollis (a-e, originals; f, from KRAMER 1988; g, from HEMBERGER 1982. a, b, from life;
c-g, protargol impregnation). a, b) Ventral views, c = 180 ~m, d = 200 ~m. The yellowish cortical granules are
shown in the right posterior quarter of (b). c, f, g) Ventral infraciliature, c = 190 ~m, f = 150 ~m. g = 270 ~m.
Arrow in (c) marks frontoventral cirrus IIII2, arrow in (g) denotes right transverse cirrus which is displaced an-
teriorly in the HEMBERGER population, indicating that it is probably another species. d) Dorsal infraciliature,
215 ~m. Arrows mark posterior end ofkinety 3 and anterior end ofkinety 4. This specimen has 2 caudal cirri
each on kineties 1 and 2 and 1 cirrus on kinety 4. Very likely this is the typical pattern in this species. e) Mid-
dle stage of morphogenesis in ventral view, 170 ~m. The cirri of a frontal-ventral-transverse primordium are
connected by a broken line. Parental cirri white, new cirri black. Arrow marks new dorsomarginal kineties of
opisthe; p 262.
266 SYSTEMATIC SECTION

Fig. 91t Allotricha mol/is (original scan-

ning electron micrograph kindly supplied
by W. FOISSNER). Ventral view, 12511m.
The arrows mark the 3 postoral ventral cirri
which are slightly larger than the cirri of
the innermost (third) right marginal row.
The transverse cirri are arranged in an
oblique row as in the neotype material (cf.
Fig. 91c) and not hook-shaped as in the
HEMBERGER population (Fig. 91g). Note
curved paroral; p 262.

Cyst spherical, in life 80 J1m

in diameter on average (75 to
90 J1m; n = 14). Cyst wall about
3-5 J1m thick, moderately wrin-
kled but without spines, which is a
further difference to A. antarctica.
Formation of the frontal-
ventral-transverse CIm streaks
typically oxytrichid (Fig. 9le). All
new right marginal rows are
formed from streaks occurring in
the outer parental right marginal
row; no parental right marginal
cirri are retained after division.
The HEMBERGER (1982; Fig.
9lg) population in life (?) about
270 x 90 J1m, ratio of length to
width about 3: 1. Elliptical, both
ends broadly rounded. Two mac-
ronuclear nodules, 2-4 micronu-
clei (Fig. 9lg). Adoral zone of
membrane lies about 113 of body
length, about 50 adoral mem-
branelles. Typical Oxytricha cirral
pattern, frontal-ventral-transverse
cirri distinctly enlarged, transverse
cirri arranged in hook shape.
Three right marginal rows with
31-37 (outer), 29-35 (middle),
and 20-22 (inner) cirri. 37-40 left
 Allotricha 267

Table 19 Morphometric data of Allotricha antarctica (ant, from PETZ & FOISSNER 1996) and A. mollis (mol,
original data; m02, from KRAMER 1988; m03, from SCHMnz 1986). All data are based on protargol-impregnated
specimens. All measurements in micrometres. ? = sample size not indicated. CV = coefficient of variation (in
%), Max = maximum value, mean = arithmetic mean, Min = minimum value, n = sample size, SD = standard
deviation

Character Species mean SD CV Min Max n

Body, length ant 139.6 16.2 11.6 105 194 31
mol 206.2 19.5 9.5 158 231 12
m02 167.7 105 205 58
Body, width ant 63.1 9.2 14.6 43 83 31
mol 64.4 8.5 13.2 52 76 12
m02 47.8 30 72 38
Anterior macronuc\ear ant 28.2 4.8 17.0 22 38 31
nodule, length mol 34.3 6.7 19.5 23 44 10
Anterior macronuclear ant 15.1 1.4 9.5 12 18 31
nodule, width mol 19.5 3.9 21.0 11 24 10
Posterior macronuclear ant 26.8 4.5 16.7 22 39 31
nodule, length mol 32.9 7.7 23.5 23 45 10
Posterior macronuclear ant 16.6 2.0 11.9 13 22 31
nodule, width mol 18.6 3.5 18.8 10 23 10
Macronuclear nodules, ant 15.5 5.1 33.1 4 27 31
distance between mol 28.0 7.4 26.7 18 40 10
Micronuclei, number ant 2.3 0.6 25.3 4 31
mol 2.4 0.5 21.8 2 3 8
m02 11.8 6 17 25
Micronucleus, diameterl ant 4.4 0.4 9.5 4 5 31
mol 3.0 0.5 16.7 2 4 9
Adoral membranelles, number ant 33.0 1.7 5.0 30 37 31
mol 56.4 3.4 5.9 50 61 11
m02 49.0 38 57 31
m03 48 64 ?
Adoral zone of membranelIes, ant 54.0 3.3 6.2 47 61 31
length moll 71.2 8.1 11.3 60 82 12
m02 68.7 60 75 20
Right marginal row I, ant 28.8 2.2 7.6 24 33 31
number of cirri mol 34.0 3.5 10.4 28 40 11
m02 29.9 24 36 27
m03 30 48 ?
Right marginal row 2, ant 17.1 3.5 20.3 10 25 31
number of cirri mol 33.1 3.5 10.6 26 38 11
m02 21.6 10 31 27
m03 22 44 ?
Right marginal row 3, mol 10.9 5.3 48.5 3 21 9
number of cirri m02 8.9 4 20 15
m03 14 26 ?
Left marginal row, ant 25.6 2.6 10.1 20 31 31
number of cirri mol 36.8 3.1 8.4 33 42 12
m02 29.1 24 38 27
m03 36 47 ?

I The micronuclei of both species are almost spherical.

268 SYSTEMATIC SECTION

marginal cirri. At least 4 dorsal kineties, cilia about 6-7 Jim long. Six caudal cirri, diffi-
cult to recognise.

o c cur r e n c e and e colo g y: Probably common in freshwater and thus presuma-

bly sometimes confused with Paraurostyla weissei. Locus classicus not mentioned by
S1ERKI (1878), but possibly Schleitheim in Canton Schafthausen, Switzerland, where he
lived and worked. According to S1ERKI, A. mollis is a frequent and abundant species. I
found it in the aufwuchs of an oligosaprobic brook in Aigen, a suburb of Salzburg, Aus-
tria (Fig. 91a-e), and in a xenosaprobic alpine brook in Kaprun, Austria. W. FOISSNER
(pers. comm.) sampled A. mollis in a mesosaprobic river in Bavaria immediately after
flooding (Fig. 91t). These records indicate that A. mollis prefers clean or only slightly
polluted running waters. HEMBERGER (1982) gives no locality. SCHMITZ (1986) found it in
the aufwuchs of lentic areas in the mesosaprobic Rhine River, Germany. The KRAMER
(1988) population is from Ithaca, USA. Feeds on green algae, diatoms, and ciliates. Bio-
mass of 106 individuals about 180 mg.

Sup p 0 sed s y non y m: Pleurotricha variabiJis REuTER, 1961, Acta zool. fenn., 99:
17 (Fig. 91h-s). Remarks: REUTER (1961) did not mention cortical granules. I presume
that he overlooked them; however, further populations have to be checked because it can-
not be excluded that a species without cortical granules exists. PErZ & FOISSNER (1996)
transferred it to Onychodromopsis (0. variabilis) and considered it as synonym of O.
jlexilis in the legend to the figures. In life 200-220 Jim long. Elliptical, broadest at about
level of contractile vacuole. Two macronuclear nodules. Adoral zone of membranelles
about 113 of body length. Occasionally 1 or 2 additional ventral cirri at level of posterior
macronuclear nodule (Fig. 9li-I). Transverse cirri protrude slightly beyond posterior end
of cell. Often a surplus, more or less long cirral row between or left of the 2 right mar-
ginal rows (Fig. 91h, n-r). Locus classicus is a pool with melted snow on grass about
60 m from the seashore near Helsingfors, Finland. Not found since.

Allotricha antarctica nov. spec. (Fig. 92a-y, 93a, b, Tables 3, 4, 19)

1972 Pleurotricha sp. - THOMPSON, Antarctic Res. Ser., 20: 285 (Fig. 92j; a slide of wet silver impregnated
specimens is deposited in the U. S. National Museum, Washington, D. C.; see THOMPSON 1972, P 287).
1978 P/eurotricha lanceolata - SMITH, Scient. Rep. FalIdd lsI. Depend. Surv., No. 95: 78 (Fig. 92k; misiden-
tification).
1996 Orrychodromopsisj/exilis STOKES, 1887 - PErz & FOISSNER, Acta Protozool., 35: 264 (Fig. 92a-i, I-y,
93a, b; authoritative description; type slides are deposited in the OberOsterreichische Landesmuseum in
Linz, Upper Austria).

Tax 0 nom y: PErZ & FOissNER (1996, 1997) identified this species with the type of
Onychodromopsis STOKES, 1887, which has, however, a different cirral pattern (Fig.
143a-d). Allotricha antarctica differs from A. mollis in body size and correlated charac-
 Allotricha 269

..
, '
e

Fig. 92a-g Allotricha antarctica (from PETZ & FOISSNER 1996. a--g, from life). a) Ventral view of a typical
specimen, 125 Jlm. b) Broad specimen. c) Lateral view. d) Cortical granulation consisting of small and large
granules. e) Cytoplasmic crystals. 1) Optical section of resting cyst, 45 Jlm. g) Detail of cyst wall; p 268.

ters (for example, number of membranelIes), position and arrangement of transverse cirri,
position of frontoventral cirrus 11112, shape and arrangement of undulating membranes
(cp. Fig. 91c with 92h), and geographic distribution so that synonymy can be excluded.
The following chapter is based mainly on the very detailed description by PIITz & FOISS-
NER (1996); data from THOMPSON (l972) and SMITH (1978) are indicated.

D i a g nos i s: Field populations in life about 90-125 x 40-70 ~m, cultured stocks
100-200 x 40-80 ~m. Cortical granules colourless. On average 33 adoral membranelles.
 270 SYSTEMATIC SECTION

1/
/ ,/
1/ ,I )
1/ ,/: ;.-6
j
;/
./ / /
: I
: \'/
/ , ,1 '\;
,/ /
I
j ,

~i :J A

T
~

."~~
j /
.~ 1-
<T ¥

.-

./

'---,

Fig. 92b, i Allotricha antarctica (from PEIZ & FOISSNER 1996. h, i, protargol impregnation). Ventral and dor-
sal infraciliature of same specimen, 155 11m. Arrow in (h) marks frontoventral cirrus III12 which is differently
arranged to in A. mollis (Fig. 91c, f). Arrowhead marks surplus dorsal kinety seen only in this specimen, arrow
in (i) denotes interkineta1 dorsal dikinetid. Explanation of original labels: Cc = caudal cirri, I, 4, 5, 6 = dorsal
kineties; p 268.

Undulating membranes almost straight and arranged side by side. Frontoventral cirrus
11112 immediately left of migratory cirrus VII3. Transverse cirri arranged in hook shape,
distinctly displaced anteriad. 2-3 right and I or 2 left marginal cirral rows.

T Y p e l 0 cat ion: Terrestrial moss from the coast of Core Bay, Prince Edward
Island, Prince Edward Islands, Subantarctic.
 Allotricha 271

M 0 r p hoi 0 g y and b i 0 log y: Field populations in life on average 113 x 55 ~m

(n = 30), specimens of cultured stocks on average 24 % larger than in raw culture.
THOMPSON (1972) population, after wet silver impregnation, 100-125 x 40-55 ~m (mean
= III x 48 ~m; n = 25). Body very flexible, broadly elliptical and dorso-ventrally flat-
tened up to 2: 1. Macronuclear nodules ellipsoidal, nucleoli globular to ellipsoidal,
1-4 ~m across after protargol impregnation. Micronuclei almost globular, usually one in
variable position in indentation of each macronuclear nodule. Contractile vacuole in mid-
body, with 2 collecting canals. Cortical granules 0.8-1.0 ~m across, arranged in rather
narrowly spaced longitudinal rows underneath entire cell surface as in Oxytricha granu-
Ii/era, rather compact, stain blue but are not extruded when methyl green-pyronin is
added, impregnate faintly to intensely with protargol depending on method and bleaching
time. A second type of granules, in great numbers between larger granules, stains red with
methyl green-pyronin (Fig. 92d); when the stain is added, many of them are extruded and
adhere to cirri and adoral membranelles. Cytoplasm colourless, in field populations with
many about 1-4 J.1m sized lipid droplets and 3-5 ~m long, colourless crystals mainly in
posterior portion (Fig. 92e). Food vacuoles 15-20 ~m across. Glides moderately fast.
Adoral zone of membranelles about 40 % of body length (PETZ & FOISSNER 1996,
THOMPSON 1972), distal portion ventrally covered by frontal scutum, cilia gradually short-
ened from distal to proximal end of zone, widest bases about 10 ~m in life. 35 adoral
membranelles on average (THOMPSON 1972). Buccal cavity occupying about one third of
cell width, anteriorly slightly curved and merging into frontal scutum, posteriorly gradu-
ally deepened and covered by buccal lip. Buccal lip hyaline, bent upright to and continu-
ous with frontal cell surface, commences at level of buccal cirrus, gradually widened pos-
teriorly to form wedge-shaped structure obliquely merging into posterior buccal vertex;
longitudinally bipartite by deep cleft containing proximal 2/3 of paroral, right sheet nar-
rower than left. Paroral after protargol impregnation 25-32 ~m long (mean = 28 ~m, n =
31), anteriorly slightly curved, cilia about 8 ~m long. Endoral after protargol impregna-
tion 25-35 ~m long (mean = 29 ~m, n = 31), straight, inserts on right wall of buccal cav-
ity, not recognisable in scanning electron micrographs because underneath buccal lip. Pa-
roral and endoral usually side by side in silvered specimens, posterior portions rarely
slightly intersecting in surface view of cell, both very likely composed of dikinetids (Fig.
92h). Pharyngeal fibres 30-55 ~m long in protargol slides, originate from adoral mem-
branelles and undulating membranes. All cirri of field populations in life about 13 ~m
long, insert in shallow cortical pits. Cirral composition as shown in Figure 92h, only
slightly variable, that is, one basal body row ± may occur in most frontal-ventral-
transverse cirri; first cirrus of inner left marginal row usually composed of three kineties.
Constantly (n = 31) 3 frontal cirri, 1 buccal cirrus, 4 frontoventral, 5 postoral and pre-
transverse ventral cirri, and 5 transverse cirri. Arrangement of frontal-ventral-transverse
cirri invariably as shown in Figure 92h, that is, almost identical to in Oxytricha granulif-
era! Transverse cirri only slightly protruding beyond posterior end of cell. Marginal rows
open at posterior end, gap occupied by caudal cirri. Usually 2 left (81 %, n = 31) and 2
right (86 %, n = 80) marginal rows; left outer row, however, frequently consisting on av-
 272 SYSTEMATIC SECTION

Or-, Q-Ir
",
"Q

Q
Q
Q
/)
q
I
,
o

CC-o 00 ..

,
17

,
Q

,
IV/2-0
V/4-0 .,
..
..
,
V/3-Q , ,
,,·-iL
,
,
Or- ~I m

Fig. 92j-o Allotricha antarctica (j, from THOMPSON 1972; k, from SMITH 1978; 1-0, from PEn & FOISSNER
1996. j, wet silver impregnation; k, NOLAND'S stain fixation; l-n, protargol impregnation). j, k) Ventral infra·
ciliature of non·dividing specimens, j = 100-125 ~m, k = 140 ~m. Arrow in (j) marks frontoventral cirrus
III12, located left of cirrus VIl3; this is an important difference to A. mollis (cf. Fig. 91c). The drawing by
SMITH (1978) is somewhat superficial, however, it very likely shows the same species. I) Very early divider
with oral primordium close to the transverse cirri. m) Early divider showing basal body patch close to postoral
ventral cirrus V/3. Arrowhead denotes outer left marginal row. n) Anlage 2 (arrow) extending anteriorly from
oral primordium. Explanation of original labels: Azm = adoral zone of membraneIles, Cc = caudal cirri, Em =
endoral, Fc = right frontal cirrus, iL = inner left marginal row, Ir = inner right marginal row, oL = outer left
 Allotricha 273

erage of 5 cirri only, in 19 % of specimens even lacking, one cell with short third row
(Fig. 92a, h). Rarely a very short third right marginal row is present, namely 1-7 cirri left
of inner row (6 %) or 2-3 cirri between inner and outer row (8 %); very likely, these are
remnants from last generation and/or young postdividers with parental cirri still in resorp-
tion (Fig. 92y, Table 19). Dorsal cilia 3-5 ~m long, insert in shallow cortical pits. Kine-
ties 1-3 almost as long as body, kinety 4 curved and commencing behind mid-body, ki-
nety 5 extends from anterior to mid-body, kinety 6 in anterior third of cell, consists of
4--5 bristles only; frequently, some irregularly arranged cilia between kineties 3 and 4,
very likely remnants from last generation. Rarely 7 kineties. According to THOMPSON
(1972) likely 5 dorsal kineties (presumably he overlooked the short dorsomarginal row!).
Three (4--5 in six of34 specimens) caudal cirri on dorsal kineties 1,2, and 4, inconspicu-
ous because not elongated and composed of only 4, rarely 6 cilia (Fig. 92h).
Resting cysts globular to slightly ellipsoidal (Fig. 92f), in life about 40 ~m
(3~8 ~m; n = 31) in diameter. Cyst wall colourless, about 2 ~m thick, compact; surface
studded with about 2 ~m long, hyaline spines and sometimes separated by narrow, trans-
parent layer from compact part of wall (Fig. 92g). Cyst contents (cell) usually conspicu-
ously lobed, does not occupy entire cyst volume (Fig. 92f); however, when cyst is slightly
pressed, its contents expand, completely filling interior. Cortex distinctly striated. Cyto-
plasm contains many 1-3 ~m sized, colourless globules. Macronuclear nodules and mi-
cronuclei do not fuse (Fig. 92f).

M 0 r p hog e n e sis (Fig. 921-y, 93a, b): This part of life cycle is described in great
detail by PETZ & FOISSNER (1996; see this paper for a detailed documentation with micro-
graphs).
Oral primordium and cirral streaks (Fig. 921-y): Stomatogenesis commences with
the apokinetal (?) proliferation of basal bodies close to the left transverse cirri (Fig. 921).
The anarchic field then elongates anteriorly in a narrow streak. Simultaneously, a small
field of basal bodies appears adjacent to postoral ventral cirrus V/3, which disintegrates
slightly later; both the basal body field and the dissolved cirrus produce the opisthe's an-
lage 6 (Fig. 92m). Very likely, this anlage does not contribute to the oral primordium be-
cause it is slightly but distinctly separate from the oral anlage in 81 % of the cases ob-
served (n = 16) and cirrus V/3 is inactive during physiological regeneration. Slightly be-
fore or after disintegration of cirrus V/3, some basal bodies separate from the anterior end
of the oral primordium, migrate anteriad and become the opisthe's anlage 2, seemingly
touching the parental undulating membranes (Fig. 92n, 0). However, scanning electron
micrographs reveal that this streak is on the cell surface and does not contact the parental
membranes, which are still intact and within the buccal cavity and the longitudinal cleft
of the buccal lip; very likely, this streak does not contact the proter's anlage 2 (dissolved
buccal cirrus) either, although sometimes being rather close to it.

E-- marginal row, Op = oral primordium, Or = outer right marginal row, Pf= frontoventral cirrus VI/4, Pm = par-
oral, Tc = transverse cirri, Vc = pretransverse ventral cirrus VII2; p 268.
 274 SYSTEMATIC SECTION

, t7

,
d

Q
Q

o p Cl

(; --
D o ~-Pm
i
'" i
E

o
:
2\~-Em
\"

'.
,
Q

.-
'"
Q

d
p
Q

q
,
Q

,
Q t7
Q

1 U
\. .. \. " Q
t7

s
Fig. 920-s Allotricha antarctica (from PETZ & FOISSNER 1996. Morphogenetic stages after protargol impreg-
nation). o-q) Early dividers showing development of the opisthe anlagen 2-6. r, s) Early dividers. Arrows
mark disaggregating cirrus IV/3 (r) and IllI2 (s). Arrowhead denotes dedifferentiating buccal cirrus. Explana-
tion of originallabeis: Em = endoral, Pm = paroral, 2-6 = frontal-ventral-transverse cirri primordia; p 268.
 Allotricha 275

~f
I

.. '
Fig. 92t, u Allotricha antarctica (from PETZ & FOISSNER 1996. t, U, protargol impregnation). Middle dividers
showing separation of cirral streaks 5 and 6 and marginal anlagen formation (arrowheads). 1-{; = frontal-
ventral-transverse cirri primordia; p 268.

Almost simultaneously with the dedifferentiation of cirrus V/3, the middle postoral
ventral cirrus Vl4 disaggregates, fonning opisthe's anlage 5; in 28 % of cases (n = 18) it
even dissolves immediately before cirrus V/3. Membranelles begin to differentiate in the
oral anlage from right to left in a posteriad direction (Fig. 920). Subsequently, the ante-
rior postoral ventral cirrus N 12 breaks up into files of basal bodies aligning to anlage 4
of the opisthe (Fig. 92p). Simultaneously, the opisthe's anlage 3 develops near, and possi-
bly from, the posterior ends of anlagen 5 and 6 or from remnants of the oral primordium
(Fig. 92q). At this stage, the opisthe's cirral streaks often appear to be connected with the
oral primordium (Fig. 92r), which has, however, usually commenced to fonn adoral
membranelles (Fig. 92p, q, s).
Posterior frontoventral cirrus IV/3 disintegrates next and produces anlage 4 of the
proter (Fig. 92r). Then, frontoventral cirrus III12 and buccal cirrus 1112 disaggregate al-
most simultaneously, fonning proter's anlagen 3 and 2, respectively (Fig. 92s). Slightly
later, the opisthe's primordia 5 and 6 separate in midregion. The anterior portions move
276 SYSTEMATIC SECTION

, 0

!
/ ~

f
!
i
J •
1 l'
i!~ (J
• I
i Q

\; ",
q "

o 0
o
"'0
o
• I ,
.
,
w

:.: ~,
f' ~

o ,. ~

,
~0 ,
,~
; ~
, ...... "~,
II'
,
, ,, ~
,,
,
, • ,,
..
,
,
,,
•.
,,
••• •
#

•• .
• ,f,
.. o
..
#
I 0

0
0
0
x y
 Allotricha 277

.
~.

....
t't, •

,. ,

.
"
.

,,

,,
....
,
, " ,

\. =-
I
;
.I

a b

Fig. 93a, b Allotricha antarctica (from PElZ & FOISSNER 1996; a, b, protargol impregnation). Morphogenesis
of dorsal infraciliature proceeds in Oxytricha pattern, that is, dorsal kinety 3 fragments (arrows) and 2 dorsa-
marginal kineties (kineties 5, 6) originate near right marginal row anlagen (see Fig. 92w); p 268.

forward, as indicated by the decreased distance to frontoventral cirrus VII3, becoming

proter's anlagen 5 and 6 (Fig. 92s--u). The origin of the opisthe's anlage 1 (undulating
membranes) could not be clarified. It develops rather late from anarchic basal bodies lo-
cated between the oral primordium and the posterior ends of the cirral streaks. Finally, all
primordia of the opisthe distinctly separate from those of the proter due to cell elongation
and/or anlagen movement (Fig. 92t, u). In the broadened middle dividers, 6 cirral streaks
each are recognisable in proter and opisthe (Fig. 92v-x). Cirri differentiate from anterior
to posterior in the classic oxytrichid pattern (Fig. 92w, x).

~ Fig. 92v-y Allotricha antarctica (from PElZ & FOISSNER 1996. v-y, protargol impregnation). v) Middle di-
vider showing marginal anlagen formation. Arrows mark anlage for inner right marginal row. w, x) Late divid-
ers showing migration of cirri, development of dorsomarginal kineties (= dorsal kineties 5, 6; arrowheads), and
origin of outer left marginal row (large arrows). y) Postdivider (opisthe; 90 11m) still having some parental cirri.
Parental cirri white, new cirri black; p 268.
278 SYSTEMATIC SECTION

Concomitant with cirral streak separation, distinct changes occur in the daughter's
oral structures. The opisthe's oral primordium invaginates, while the parental adoral zone
and buccal cavity evaginate. The evagination is accompanied by a dedifferentiation of the
parental paroral, which proliferates basal bodies at least at its anterior end (Fig. 92u, v).
This anlage forms the leftmost frontal cirrus. The endoral, which is always separate from
the paroral anlage, is very likely also partially or completely reorganised as indicated by
its slightly loosened kinetids; a special primordium is, however, not formed. When the
buccal cavity evaginates, the undulating membranes become more narrowly spaced and
optically intersect in their posterior third. When the buccal field invaginates in early post-
dividers, the membranes move apart and thus do not intersect any longer (Fig. 92w-y).
While the buccal cavity is restored, the pharyngeal fibres are resorbed and rebuilt.
Marginal anlagen (Fig. 92t-y): Differentiation commences in middle dividers. The
second (47 % of cases), third (35 %) or, rarely, first (18 %) cirrus of the outer right row
and, slightly later, invariably the fIrst cirrus of the inner left row commence anlagen for-
mation in the proter; the primordia for the opisthe originate from parental cirri at about
mid-body (Fig. 92t, u). A few cirri each disintegrate into files of basal bodies which align
longitudinally to form a row. These primordia subsequently become double-rowed and
elongate by basal body proliferation and incorporation of additional cirri at their posterior
ends (Fig. 92u, v). The inner right and outer left marginal row are morphogenetically in-
active, but originate from primordia formed at the anterior ends of the remaining parental
outer right and inner left marginal rows respectively (Fig. 92v). The primordia in the left
marginal row are distinctly smaller than those in the right row and thus produce fewer
cirri. The parental inner right marginal row is completely resorbed only after cytokinesis.
Therefore, specimens having three right marginal kineties are very likely postdividers
(Fig. 92y).
Dorsal morphogenesis is in Oxytricha pattern. Usually 1, rarely 2, caudal cirri each
differentiate at the posterior ends of kineties I, 2, and 4 (Fig. 93a, b). Nuclear division
proceeds as usual, that is, the macronuclear nodules frrst fuse and then split twice, the last
macronuclear division being completed in the postdividers. Physiological regeneration
(reorganisation) largely resembles the development of the proter. For details, see PETZ &
FOISSNER (1996).

o c cur r e n c e and e colo g y: Common in terrestrial habitats of the Antarctic re-

gion. PETZ & FOISSNER (1996) found their populations at 2 sites of the Prince Edwards Is-
lands (lat 46°35'S, long 37°56'E; Subantarctic), namely in a slightly saline and acidic (PH
6.1) grass sward on rock at the seaward limit of vegetation on Marion Island and in a
slightly saline and acidic (PH 6.3) moss sample collected between Poa and Callitricha
vegetation near a penguin rookery on Prince Edward Island (collected by Dr J. COOPER,
University of Cape Town). At these sites, A. antarctica occurred together with Acineria
uncinata, Colpoda aspera, C. cucullus, C. injlata, C. stein ii, Cyrtolophosis mucicola,
Haplocaulus terrenus, Leptopharynx costatus, Nivaliella plana, Platyophrya vorax, and
Pseudoplatyophrya nana. SMITH (1978) observed Allotricha antarctica in moss peat,
 Cyrtohymena 279

grass soil, and Aceana soil in South Georgia, the South Orkney Islands, the Elephant Is-
land, and many other sites of the Antarctic region (SMITH 1972, 1973a, 1974a, 1981,
1982, 1984). Conversely, THOMPSON (1972) found his population in a rock tidal pool with
freshwater dilution on one of the Joubin Islands, near the Antarctic peninsula. FOISSNER
(1996c) recorded A. antarctica from soil of Marion Island, South Atlantic Ocean.
Feeds on bacteria (up to 45 11m long), starch, flagellates (Polytoma sp.), and ciliates
like Colpoda steinii and Cyclidium glaucoma (PETZ & FOISSNER 1996). Biomass of 106 in-
dividuals about 80 mg (PETZ & FOISSNER 1996).

Cyrtohymena FOISSNER, 1989

1989 Cyrtohymena FOISSNER, Sber. 05t. Akad. Wiss., 196: 238 - Type (original designation): Oxytricha
(Steinia) muscorum KAlu., 1932.
1997 Cyrtohymena FOISSNER, 1989 - BERGER & FOISSNER, Arch. Protistenk., 148: 145.

C h a r act e r i sat ion: Adoral zone of membranelies formed like a question mark.
Undulating membranes in Cyrtohymena pattern. Frontoventral cirri form V-shaped pat-
tern. Postoral ventral cirri in dense cluster behind buccal vertex. Two pretransverse and 5
(rarely 4) transverse cirri. One right and 1 left row of marginal cirri. Usually 6 dorsal ki-
neties. Caudal cirri present. Primordia V and VI of proter originate de novo. Dorsal mor-
phogenesis in Oxytricha pattern.

Add i t ion a I c h a r act e r s: Outline long elliptical, rarely distinctly converging

posteriorly. Usually 2:1, rarely more flattened. Body rather flexible, some species, how-
ever, almost rigid (see taxonomy). Two or 4 (c. quadrinucleata) macronuclear nodules.
One, two, or more micronuclei. Contractile vacuole in or near mid-body; collecting ca-
nals usually present. About half of species with cortical granules; Cyrtohymena mUSCD-
rum additionally with reddish granules in cytoplasm. Pellicle and cytoplasm colourless
(not homogeneously coloured), an important difference to Rubrioxytricha. Movement
usually rapid, sometimes slowly gliding. Buccal cavity broad and very deep, thus bright
at low magnification. Frontal cirri slightly to distinctly enlarged as compared with fronto-
ventral cirri, often arranged in a steeply rising row (from right to left). Presumable all
species with 6 dorsal kineties, inclusively 2 dorsomarginal rows (see, however, the very
large Cyrtohymena australis). Three inconspicuous to distinctly elongated caudal cirri on
dorsal kineties 1,2, and 4. Dorsal cilia 3-5 11m long.

T a x 0 nom y: The characterisation is according to BERGER & FOISSNER (1997). For a de-
tailed description of the patterns mentioned in the characterisation, see chapter 2 in the
general section. Cyrtohymena contains the bulk of the species originally described in
Steinia. BORROR (1972a) and HEMBERGER (1982) synonymised Steinia with Oxytricha,
considering the differences in the oral apparatus as insufficient to distinguish genera. This
simplification was not accepted by GROLIERE (1975), and FOISSNER (1982), who re-
280 SYSTEMATIC SECTION

established Steinia (now Cyrtohymena) because of the apomorphic pattern of undulating

membranes (Fig. 199, 20e, 2Ic). Somewhat later, FOISSNER (1989) described Steinia
sphagnieola, a species very closely related to the type of Steinia, S. platystoma. Both
species have a fragmented sinusoidal endoral which does not intersect with the curved pa-
roral (Fig. 20t). Thus, the other Steinia species were assigned to a new taxon, Cyrtohy-
mena (FOISSNER 1989).
Cyrtohymena is well defined and usually easily identified by the sole autapomorphy,
namely the distinctive arrangement of paroral and endoral and the broad, deep and thus
very transparent buccal cavity (Fig. 981). However, some species (c. inquieta, C. quadri-
nucieata, C. tetracirrata) deviate from the majority in that the body is rather inflexible,
the relative length of the adoral zone is 40-50 % of body length, and the postoral ventral
cirrus V/3 is distinctly displaced posteriad and thus possibly not involved in primordia
formation. All three character states indicate that these three species belong to the Stylo-
nychinae, that is, Cyrtohymena is possibly not monophyletic and the distinctive pattern of
undulating membranes evolved twice. However, detailed morphogenetic and molecular
biological data are needed to prove this hypothesis.
Cyrtohymena should not be confused with Steinia, which has another pattern of undu-
lating membranes, a peculiar pit in the buccal cavity, and a very rigid body (Fig. 22b).
Possibly some of the little known Cyrtohymena species (c. gracilis, C. sapropeliea, C.
aestuarii, C. granuiata, C. torrentieola) are synonyms of other species or belong to
closely related taxa, such as Notohymena.
Very likely, Cyrtohymena has a world-wide distribution; however, there is some evi-
dence that certain species do not occur in Australia. BUTIERER & FOISSNER (1988), for in-
stance, studied 21 soil and moss samples from this continent, but did not find C. museD-
rum, which is rather common in European terrestrial habitats. Cyrtohymena comprises
terrestrial, freshwater, and marine species.

Key to species

Species identification in Cyrtohymena unconditionally needs live observation because

several species have cortical granules, that is, identification after only protargol impreg-
nation is insufficient.

1 2 macronuclear nodules ............................................... 2

4 macronuclear nodules (Fig. 103a, b) ............... C. quadrinucieata (p 320)
2 1 micronucleus, usually arranged between the 2 macronuclear nodules (for example,
Fig. 96f, h, 10 Ii) .................................................... 3
2 or more micronuclei ................................................ 5
3 Cortical granules brownish, arranged in longitudinal rows (Fig. 96f) ............ .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . C. gracilis (p 305)
Cortical granules absent .............................................. 4
 Cyrtohymena 281

4 Transverse cirri conspicuously displaced anteriad, do not protrude beyond posterior

end of cell; in life about 100 Ilm long; sapropelic form (Fig. 96h) ............. .
· . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. . C. sapropelica (p 306)
Transverse cirri protrude distinctly beyond posterior end of cell; in life about 150 to
180 Ilm long; terrestrial habitats, especially mosses (Fig. 10 1h, i) ................ .
· ........................................... " . Cyrtohymena sp. (p 307)
5 (2) Cortical granules present ........................................... 6
Cortical granules absent .............................................. 8
6 Cortical granules pink to ruby-red, granules occur also in cytoplasm (Fig. 94a, b, d,
g, h) .............................................. C. muscorum (p 281)
Cortical granules not pink to ruby-red, do not occur in cytoplasm ........... 7
7 Marine habitats ....................................... C. marina (p 323)
Freshwater or terrestrial habitats ....................................... 13
8 (5) 4 transverse cirri .................................................. 9
5 transverse cirri ................................................... 10
9 Terrestrial (Fig. 102a, c, f, h) ......................... C. tetracirrata (p 317)
Freshwater or brackish water (Fig. 101 e, t) ................ C. aestuarii (p 308)
10 (8) Especially central portion of cell with dark (sometimes greenish) granules (no
cortical granules; Fig. 96g) ............................ C. granulata (p 306)
Cell without conspicuous dark granules ................................. 11
11 Outline rather slender, posterior end of cell narrowly rounded (Fig. 96i) ......... .
· . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . C. torrenticola (p 307)
Outline otherwise ................................................... 12
12 Length in life less than 150 Ilm (Fig. 101a~) ............... C. inquieta(p 313)
- Length in life about 150-250 Ilm (Fig. 99a-m) .............. C. candens (p 308)
13 (7) Length in life 250-400 Ilm (Fig. 97a) .................. C. australis (p 296)
Length in life 90-170 J.1m ............................................ 14
14 Transverse cirri displaced markedly anteriad (Fig. 98a, t) .. C. primicirrata (p 300)
Transverse cirri displaced not markedly anteriad (Fig. 96a) ..... C. citrina (p 293)

Cyrtohymena muscorum (KARL, 1932) FOISSNER, 1989 (Fig. 94a-z,

95a-s, Tables 3, 4, 20)

1932 Steinia muscorum KAHL, Tierwelt Dtl., 25: 613 (Fig. 94a).
1956 Steinia dubia GElLERT, Acta bioI. hung., 6: 349 (Fig. 94n; new synonym).
1963 Steinia inquieta (STOKES, 1887) - REllrER, Sarsia, 10: 7 (Fig. 94m; misidentification).
1970 Steinia muscorum, KAHL, 1932 - DRAGESCO, Annis Fac. Sci. Univ. fed. Cameroun (Numero hors-serie):
118 (Fig. 94k).
1972 Oxytricha muscorum KAHL, 1932 - BORROR, J. Protozool., 19: 14.
1972 Tachysoma dubia (GEU.ERT, 1956)-BoRROR, J. Protozool., 19: 15.
1977 Oxytricha rubra BUITKAMP, Decheniana, 130: 122 (Fig. 941).
282 SYSTEMATIC SECTION

1982 Steinia muscorum KAHL, 1932 - FOISSNER, Arch. Protistenk., 126: 105 (Fig. 94b-f; authoritative
redescription; neotype slides are deposited in the Ober()sterreichische Landesmuseum in Linz, Upper
Austria).
1986 Steinia muscorum KAHL, 1932 - DRAGESCO & DRAGESco-KERNEIS, Faune tropicale, 26: 482.
1987 Oxytricha germanica FOISSNER, Arch. Protistenk., 133: 224.
1989 Steinia muscorum KAHL, 1932 - BERGER & FOISSNER, Bull. Br. Mus. nat. Hist. (Zool.), 55: 41 (Fig.
94g-j; a protargol-impregnated slide [1988:2:1:23] is deposited in the British Museum [Natural
History] in London).
1989 Cyrtohymena muscorum (KAHL, 1932) - FOISSNER, Sber. ()st. Akad. Wiss., 196: 239.
1991 Cyrtohymena muscorum (KAHL, 1932) FOISSNER, 1989 - Voss, Arch. Protistenk., 140: 67 (Fig. 94o--z,
95a-s).
1996 Cyrtohymena muscorum - BONKOWSKI, Berichte des Forschungszentrums Wald()kosysteme, 134: 80,
Abb. 6.2.2a.

Nom e n c I a t u rea n d t a x 0 nom y: KAHL (1932) classified Steinia as subgenus

of Oxytricha; the correct name in the original description is thus Oxytricha (Steinia) mus-
corum. Since the presence of a subgeneric name does not affect homonymy between spe-
cies group names (ICZN 1985, Article 57 d), this name is a primary homonym of3 other
muscorum species described by KAHL (1932) in other subgenera (Opisthotricha, Histrio,
Stylonychia) of Oxytricha. Hence, 3 of these 4 species required replacement names
(FOISSNER et al. 1991). "Steinia muscorum" was fixed as type of Cyrtohymena (FOISSNER
1989); thus, we did not replace its species name. The synonym Oxytricha rubra Burr-
KAMP, 1977a is preoccupied by Oxytricha rubra EHRENBERG, 1835 (Abh. preuss. Akad.
Wiss., year 1835: 164). EHRENBERG'S species is the well-known marine Pseudokeronopsis
rubra, whereas the BUITKAMP (1977a) species was renamed as Oxytricha germanica and
concomitantly synonymised with Cyrtohymena muscorum (FOISSNER 1987d).
Steinia dubia, also having a pale pink cytoplasm and a semicircularly curved paroral,
is very likely a further synonym of Cyrtohymena muscorum. In Steinia dubia the
posterior-most marginal cirri are longer than the other marginal cirri and the marginal
rows are not confluent posteriorly (Fig. 94n), which is, according to GElLERT (1956),
typical for Tachysoma; very likely for the reason BORROR (1 972a) transferred it to Tachy-
soma, which is, however, unreasonable because the type species of Tachysoma, T. pel-
lionel/um, has a rather different arrangement of undulating membranes. I assume that
GELLERT (1956) overlooked the caudal cirri.
REUTER (1963a) stated that his Steinia inquieta is not identical with Cyrtohymena
muscorum. However, the habitat, the brownish cortical granules (which are irregularly
distributed in the cytoplasm), and the habitus do not support his statement.
DRAGESCO (1970) mentioned neither cortical granules nor any colour of the cyto-
plasm, very likely because he did not make live observations. In spite of this uncertainty I
accept his identification, because all other data agree rather well with the original descrip-
tion. However, DRAGESCO (1970) provided differing sizes, namely 96-116 ~m in the text
and about 260 ~m for Figure 94k.
 Cyrtohymena 283

r
(' ,
('

a
c d

Fig. 94a-f Cyrtohymena muscorum (a, from KAHL 1932; 1>--1; from FOISSNER 1982. b, e, alpine population
from Hochmais [neotype material]; d, 1; population from mosses in Morzg, Salzburg city. a--d, from life; e, f,
protargol impregnation). a-d) Ventral, lateral, and dorsal view, a = 200 11m, b = 150 11m, d = 185 11m. e, 1)
Ventral and dorsal infraciliature, e = 120 11m, f= 105 11m. CC = caudal cirri, K = large vacuole with crystals,
sG = ruby to pale red, about 0.5-1.5 11m sized cortical granules occurring also in the cytoplasm; p 281.

M 0 r p hoI 0 g y and b i 0 log y: In life 130-200 x 30--60 J.1m (BERGER & FOISSNER
1989a, FOISSNER 1982), after KAHL (1932) up to 250 J.1m, according to REUTER (1963a)
130 J.1m long. Length of synonyms 140-150 J.1m (GELLERT 1956) and, respectively,
180-200 J.1m (BUITKAMP 1977a). Elliptical, both ends more or less broadly rounded. Body
rather flexible. Macronuclear nodules in vivo about 20 x 12 J.1Ill, after protargol im-
pregnation 14-25 x 6-14 J.1m, with many small nucleoli. Usually 2, rarely up to 5, in life
about 6 x 4 J.1m sized micronuclei. Contractile vacuole with long collecting canals, ante-
rior one often with a small dilatation. Pellicle colourless, close underneath and in cyto-
plasm few to very many 0.5 J.1m (sometimes up to 1.5 J.1m) sized, ruby to pale red gran-
ules so that cells are slightly to distinctly reddish; granules usually irregularly arranged,
mainly around dorsal bristle rows and cirri, sometimes forming short rows (Fig. 94a, d,
h). After death granules loose colour very rapidly. Cells usually packed with 2-5 J.1m
sized, yellowish, spindle- or drumstick-shaped crystals. Movement moderately rapid.
Adoral zone of membranelles about 35 % of body length. Bases of largest mem-
branelles in life about 8 J.1m wide. Undulating membranes consist of2 rows of basal bod-
ies. Buccal area partially covered by a hyaline cytoplasmic fold (Fig. 94g). Frontal cirri
strongly enlarged. Rather invariably 2 pretransverse ventral cirri, according to FOISSNER
 284 SYSTEMATIC SECTION

,
, ~

" ,
" ,
,"
.
: .
~

. .- ,,.• . \ '\
.. \.
\
\ ;
,--
Fig.94g-j Cyrlohymena muscorum (from BERGER & FOISSNER 1989a g, h, from life; i,j, protargol impregna-
tion). g, h) Ventral and lateral view, 160 ~m. i, j) Ventral and dorsal infraciliature, 145 ~m. Note that all pos-
toral ventral cirri are arranged rather closely to buccal vertex; p 281.

(1982), however, rarely up to 6 such cirri present. Transverse cirri in life 20-30 f!m long,
protrude distinctly beyond posterior end of cell. Left marginal row J-shaped (Fig. 94e, i),
after BUITKAMP (1977a), however, tenninates about at level of posterior-most transverse
cirrus. Marginal cirri in life 20 f!m, dorsal cilia 3 f!m, after BUITKAMP (1977a) 5 f!m long.
Dorsal kineties 3 and 5 slightly shortened posteriorly, kinety 4 begins and kinety 6 tenni-
nates somewhat in front of mid-body. Very rarely a seventh kinety with 2-4 basal body
pairs; DRAGESCO (1970) counted only 5 kineties (very likely he overlooked the short ki-
nety 6). Caudal cirri in life 15-25 f!m long.
Cyst spherical, 40-55 f!m across, according to REUTER (1963a) only about 35 f!m.
Outer membrane about 0.7 f!m thick, smooth, without a mucous layer. Going inwards, 3

Fig. 94k-n Cyrtohymena muscorum (k, from DRAGESCO 1970; I, from BUITKAMP 1977a; m, after RE1.rrER ~
1963a; n, from GEIl.ERT 1956. k, I, protargol impregnation; m, from life; n, sublimate fixation and opalblue
staining). k, I, n) Ventral infraciliature and nuclear apparatus, k = 260 11m, I = 185 11m, n = 135 11m. m) Ven-
tral view, size not indicated; p 281.
Cyrtohymena 285

o
o·

n
 286 SYSTEMATIC SECTION

5~~
,,
,
\
(

,I (, I
I I
II I
~(
pvc- :
#
,
I(
RMR- -;; - ·LMR
't
.. /

... "
;
;
"-
-., "
,
"
"
PRVC- _#-. ; /'
I
I
TC---';--: \
• •
If·
,~
- I
\

0°

:"\\;
0

\~
"
" !~: o .'
~op
" ~ :
"
" "
... c
c
"
""
""

Fig. 940-t Cyrtohymena muscorum (from Voss 1991b. 0, from life; p-t, protargo( impregnation). o-q) Ven-
tral view and ventral and dorsal infraciliature of non-dividing specimens, 0 = 135 J.lm, p, q = 160 J.lm. r-t)
Ventral infraciliature of very early and early morphogenetic stages, r = 160 J.lm, s = 165 J.lffi, t = 185 J.lm. AZM
= adoral zone of membranelles, BC = buccal cirrus, CC = caudal cirri, e = endoral, FC = frontal cirri, FVC =
frontoventral cirri, LMR = left marginal row, Ma = macronuclear nodule, Mi = micronucleus, OP = oral pri-
mordium, p = paroral, PRVC = anterior pretransverse ventral cirrus, PVC = postoral ventral cirri, RMR = right
marginal row, TC = transverse cirri, 1-6 = dorsal kineties 1-6; p 281.
 Cyrtohymena 287

AO- D

aM- D

" a
"

"
o o
o
~ 0

0:
"',\ , 0
"~ O~
0
• ' •• 0 W

o
• 0
Q/
oa 0 0 Z

Fig. 94u-z Cyrtohymena muscorum (from Voss 1991b. u-z, ventral infraciliature of morphogenetic stages af-
ter protargol impregnation). u-z) Early and middle stages, u = 175 11m, v, x = 145 11m, w = 150 11m, y =
140 11m, z = 125 11m. Some parental cirri are modified to primordia, namely the postoral ventral cirri (v,
arrows), the posterior frontoventral cirri (w, arrows), and the buccal cirrus (x, arrow). AO = anlage of frontal
cirri of opisthe, aM = new adoral membranelles; p 281.
 288 SYSTEMATIC SECTION

(( I

"
~~.,
"

',~.
\
,

OP-r~!~:
, '. ",'
# ••

I ,':t,
OP- -j ..;J
I

":G':
.
.... .
.'
"
.
• t .:. I

,
• • • I

,.• e. •
"

Fig. 95a-g Cyrtohymena muscorum (from Voss 1991b; a-g, ventral and dorsal infraciliature of morphoge-
netic stages after protargol impregnation), a-g) Middle stages, a, b = 135 Jlm, c, g = 130 Jlm, d-f= 120 Jlm,
Arrows in (a) mark anterior-most cirri of marginal rows which are modified to primordia; arrowhead denotes
anlage of undulating membranes of opisthe, The anlagen 5 and 6 of the proter originate de novo, the autapo-
morphy of the Cyrtohymena-Notohymena-Urosomoida clade. Parental cirri white, new cirri black, OP = dorsal
primordia, 1-6 = frontal-ventral-transverse cirri streaks I-VI; p 281.
 Cyrtohymena 289

Fig.9Sh-m Cyrtohymena muscorum (from Voss 1991b. 1Hn, ventral and dorsal infraciliature and nuclear
apparatus of morphogenetic stages after protargol impregnation). h-m) Late stages showing migration of new
cirri, formation of marginal rows, and early stages of dorsal morphogenesis which is in Oxytricha pattern. Bars
= 50 11m. Arrows mark anlagen of dorsomarginal kineties. Note that the undulating membranes are not curved
and arranged side by side at this stage of division. Parental structures white, new black; p 281.
 290 SYSTEMATIC SECTION

. ,.
.:
,,
,,,
;
~f
'fff f
,
,
," '
. .J
.. ,,
,, •••
t, •
~,
~
't
"o
If ,

•.....
••
,
• :
,,,• , .'; ~~

, · '.
,
••,
, ••
••

•
I
I :f4
· ..
· ~:.',
• • ,,
f -I ,,,
•t
f
I'

•,.t
t
t j "

to q r s

Fig. 950-s Cyrtohymena muscorum (from Voss 1991b. n-s, ventral and dorsal infraciliature of morphoge-
netic stages after protargol impregnation). o-p) Very late stages, bar = 50 Ilm. q-s) Postdividers, bar = 50 Ilm.
Note, that the genus-specific shape of the undulating membranes is only attained just after cell fission. Parental
cirri white, new cirri black; p 281.

zones are discernible: (i) a colourless, vitreous, 2 Jlm thick, shining layer, which remains
after squeezing the cyst, (ii) a dense layer of red granules, and (iii) the centrally arranged,
colourless cytoplasm with the nuclear apparatus (ForSSNER 1982).
 Cyrtohymena 291

Table 20 Morphometric data of Cyrtohymena australis (from FOISSNER 1995b. aul, population from Peru;
au2, population from Costa Rica), C citrina (cit, from BERGER & FOISSNER 1987a), C. muscorum (mul, from
BUITl(AMP 1977a; mu2, from FOISSNER 1982; mw, from BERGER & FOISSNER 1989a; mu4, from Voss 1991b), C.
primicirrata (pri, from FOISSNER 1984), C. quadrinucleata (qui, from ORAGESCO & NJlNE 1971; qu2, from
FOISSNER 1984), C. tetracirrata (tet, from BERGER & FOlSSNER I 987a). All data are based on protargol-
impregnated specimens. All measurements in micrometres. '/ = sample size not indicated; if only one value is
known it is listed in the mean column, if 2 values are available they are listed as Min and Max. CV = coeffi-
cient of variation (in %), Max = maximum value, mean =arithmetic mean, Min = minimum value, n = sample
size, SO =standard deviation

Character Species mean SO CV Min Max n

Body, length aul 285.3 38.7 13.6 245 370 15
au2 179.1 22.6 12.6 145 230 15
cit 99.0 11.3 11.3 75 112 11
mul 180 200 '/
mu2 120.0 13.4 11.2 100 146 13
mw 129.4 14.7 11.3 105 155 22
mu4 175.8 40.2 22.8 117 246 25
pri 84.7 10.0 11.9 74 102 9
qui 107.0 100 120 '/
qu2 86.6 6.3 7.2 77 100 10
tet 89.3 8.1 9.0 77 105 8
Body, width aul 97.3 10.3 10.6 80 120 15
au2 67.6 10.7 15.9 50 83 15
cit 33.5 4.8 14.2 27 41 11
mu2 46.0 5.1 11.0 37 56 13
mw 47.7 5.3 11.2 36 56 22
mu4 49.1 7.8 15.9 33 66 25
pri 36.0 2.8 8.3 29 38 9
quI 47.0 40 57 '/
qu2 40.8 4.3 10.5 34 49 10
tet 46.1 8.5 18.4 34 59 8
Adoral membranelles, number aul 57.3 4.8 8.4 50 64 15
au2 48.1 5.5 11.4 43 58 15
cit 33.6 2.0 6.0 31 37 11
roul 37 41 '/
mu2 41.1 3.5 8.6 33 47 13
mw 34.8 3.9 11.1 24 40 22
mu4 34.7 2.9 8.3 30 45 25
pri 30.2 3.2 10.7 27 36 9
qui 36.0 35 38 '/
qu2 40.0 3.4 8.4 35 47 10
tet 33.4 2.7 8.0 28 36 8
Right marginal row, number of cirri aul 36.1 3.6 10.1 29 41 15
au2 33.8 3.9 11.7 27 40 15
cit 21.1 2.2 10.3 18 25 11
mul 23 I
mu2 25.0 1.8 7.2 22 28 13
mw 23.0 1.7 7.5 20 25 22
rou4 20.1 1.9 9.5 18 24 25
292 SYSTEMATIC SECTION

Table 20 Continued

Character Species mean SD CV Min Max n

Right marginal row, number of cirri pri 20.3 2.0 9.8 17 24 9
qui 22.0 12 23 ?
qu2 20.3 0.9 4.7 18 21 10
tet 17.0 3.1 18.3 14 24 8
Left marginal row, number of cirri aul 38.1 3.7 9.7 31 42 15
au2 35.8 4.2 11.8 31 45 15
cit 21.5 2.1 9.6 17 24 II
mul 22.0 I
mu2 27.4 2.3 8.5 24 31 13
mu3 24.7 2.6 10.5 20 29 22
mu4 25.0 1.5 6.2 22 28 25
pri 15.6 1.5 9.7 13 18 9
qui 19.0 17 20 ?
qu2 18.3 2.2 12.1 IS 22 10
tet 16.4 2.7 16.3 13 21 8
Dorsal kineties, number cit 5.8 0.4 6.9 5 6 II
mul 6.0 ?
mu2 6.0 0 0 6 6 13
mu3 6.1 0.3 4.8 6 7 22
mu4 6.0 0 0 6 6 25
pri 6.0 0 0 6 6 10
qu2 6.0 0 0 6 6 8
tet 6.0 0 0 6 6 8

Morphogenesis of Cyrtohymena museorum is described in great detail by Voss

(1991b; Fig. 94r-z, 95a-s; Table 4). It proceeds rather similarly to in other oxytrichids.
In the middle stages the paroral and endoral are arranged in parallel. They cross in late
stages, and the Cyrtohymena-specific figure is attained only after cell fission (Fig.
95p-s). The de novo formation of primordia V and VI of the proter is possibly an autapo-
morphy of the Urosomoida-Cyrtohymena-Notohymena group (Fig. 25a, Table 4).

o c cur r e n c e and e colo g y: Common and sometimes rather abundant in terres-

trial mosses and soil; vel)' likely confined to terrestrial habitats. Locus classicus, that is,
sample site of type population, not fixed. KAHL (1932) discovered Cyrtohymena museD-
rum in mosses of Germany and California, USA. According to Article 75 (f) of the IcZN
(1985), the place of the origin of the neotype becomes the type locality of the nominal
species. ForSSNER (1982), who provided the neotype, found C. museorum in the soil of an
alpine pasture at Hochmais beside the Glockner Road, Salzburg, Austria. A detailed de-
scription of this site (new locus classicus), dominated mainly by Rumex sp. and Urtiea
sp., is given in ForSSNER (1981; site SO 3).
Further records substantiated by illustrations: Austria (FOISSNER & ForSSNER 1988);
litter of a beech forest near Bergheim, Salzburg, Austria (BERGER & FOISSNER 1989a); soil
 Cyrtohymena 293

of a mixed woodland in Melbtal near Bonn, Gennany (BUITKAMP 1977a, 1979; locus clas-
sicus of the synonym Oxytricha rubra); terrestrial moss in Ruhpolding, Gennany (Voss
1991 b); leaf litter of a beech forest in Gennany (BONKOWSKI 1996); humus underneath
mosses near the village of Boldogkovara1ja, Hungary (GEU. ERT 1956; locus classicus of
the synonym Steinia dubia); rain-wet mosses near Espegrend, Blomsterda1en, Norway
(REUTER 1963a, b); infusion of dry mosses from the park of the University of Yaounde,
Cameroon (DRAGESCO 1970).
Records not substantiated by illustrations: garden soil in Upper Austria (PETz et al.
1988); agricultural soil and dry mosses in Czechoslovakia (TIRJAKovA 1988, TIRJAKovA &
MATIS 1987a); dry mosses, leaf litter, inundated meadows, agricultural soi~ and soil infu-
sions in Gennany (DINGFELDER 1962, LElll..E 1989, WENZEL 1953); soil from Japan (TAKA-
HASID & SUHAMA 1991); soil and mosses from the Antarctica (SUOZUKI 1979). Possibly,
Cyrtohymena muscorum is not represented in the Australian fauna because BLATIERER &
FOISSNER (1989) did not record it from any of their 21 samples from various terrestrial
habitats.
TIRJAKovA (1992b) and TIRJAKovA & MATIS (1987a) recorded C. muscorum from
freshwater in Czechoslovakia, BUCK (1961) from various, usually heavily polluted run-
ning waters in Nordwiirttemberg, Gennany, and WANG (1977) from lentic water bodies of
the Tibetan Plateau. Probably, they observed Rubrioxytricha species, which are also red-
dish but confmed to freshwater habitats.
Feeds on fungal spores, diatoms, heterotrophic flagellates, testaceans (Schoenbornia
sp.), and ciliates (Odontochlamys sp., Opercularia sp., Gonostomum sp.). Voss (1991b)
cultured C. muscorum in mineral water (Vittel); every second or third day a few drops of
a suspension of yeast cells and Chlorogonium elongatum were added. Biomass of 106
large specimens about 225 mg (FOISSNER 1987a); REUTER (1963b) calculated an average
biomass of only 55 mg.

Cyrtohymena curina (BERGER & FOISSNER, 1987) FOISSNER, 1989 (Fig.

96a-e, Table 20)

1932 Steinia (Histrio) inquieta (STOKES, 1887) - KAHL, Tierwelt Ot\., 25: 613 (Fig. 96e; misidentification).
1985 Steinia citrina FOISSNER - FOISSNER, PEER & AoAM, Mitt. Ost. bodenk. Ges., 30: 109 (nomen nudum).
1987 Steinia citrina BERGER & FOISSNER, Zoo\. Jb. Syst., 114: 225 (Fig. 96a-d; one slide of holotype and 1
slide of paratype specimens are deposited in the OberOsterreichische Landesmuseum in Linz, Upper
Austria).
1989 Cyrtohymena citrina (BERGER & FOissNER, 1987) - FOISSNER, Sber. Ost. Akad. Wiss., 196: 239.

Nom e n c I at u rea n d t a x 0 nom y: FOISSNER et al. (1985) published the name

without indication; thus, it was a nomen nudum until BERGER & FOISSNER (1987a) pro-
vided the original description. The combination Cyrtohymena citrina was already used
by BLATTERER & FOISSNER (1988, p 70), but FOISSNER (1989) must be credited with the
authorship because he established Cyrtohymena. KAHL (1932) classified Steinia as subge-
 294 SYSTEMATIC SECTION

,
,
",
",
,
~

,
", ,
,
"
"
"
"~ •
tI ' •
•••

Fig. 96a-d Cyrtohymena citrina (from BERGER & FOISSNER 1987a. a, b, from life; c, d, protargol
impregnation). a, b) Ventral and lateral view, a = 130 f.!Dl, b = 125 f.!m. The yellow to orange-yellow cortical
granules are shown in (b) only. c, d) Ventral and dorsal infraciliature, c, d = 100 f.!m. Note that all postoral
ventral cirri are arranged close to the buccal vertex; p 293.

nus of Oxytricha; the correct name in his revision is thus 0. (Steinia) inquieta. KAHL
(1932) erroneously identified his granule-bearing population with C. inquieta, which has,
however, no cortical granules according to the original description (STOKES 1887b).
Cyrtohymena citrina should not be confused with C. primicirrata (transverse cirri
distinctly displaced anteriad) and C. australis (significantly larger, namely 250-400 I.I.m).
Possibly these three yellowishly granulated species form a clade within Cyrtohymena.

M 0 r p hoi 0 g y and b i 0 log y: In life 120-150 I.I.m, after KAHL (1932) 100 to
170 I.I.m long. Slender, sometimes somewhat S-shaped, left margin slightly convex, right
nearly straight; according to KAHL (1932) margins distinctly converging posteriorly (Fig.
96e). Both ends more or less broadly rounded. Body flexible and moderately contractile.
Macronuclear nodules in life 25 x 14 I.I.m, after protargol impregnation 11-21 times
7-10 I.I.m. 1--4, usually 2, in life about 7 x 5 I.I.m sized micronuclei. Contractile vacuole
 Cyrtohymena 295

Fig. 96e-j Ventral view of some Cyrlohymena species (from life). e) Cyrlohymena cilrina (from KAm 1932),
150 11m. Differs from the type population in that the postoral ventral cirrus V13 is displaced somewhat poste-
riad; p 293. f) Cyrlohymena gracilis (from KAm 1932), 100 11m. Note the single micronucleus between the
two macronuclear nodules; p 305. g) Cyrlohymena granuiala (from KAm 1932), 100 11m; p 306. h) Cyrtohy-
mena saprope/ica (from KAm 1932), 100 11m. Note the single micronucleus between the two macronuclear
nodules and the distinctly anteriorly displaced transverse cirri; p 306. i) Cyrlohymena lorrenlicoia (from
SRAMEK-HUSEK 1957), 120 11m; p 307. j) Cyrlohymena marina (from KARL 1932), 130 !lm; p 323.

with inconspicuous collecting canals. Cells yellow at low magnification due to yellow to
orange-yellow cortical granules, about 0.5-1.5 11m across, mainly arranged around cirri
and dorsal cilia (Fig. 96b); according to KAHL (1932) granules brownish (possibly due to
weak illumination).
Adoral zone of membranelles about 35 % of body length. Undulating membranes
consist of 2 or 3 rows of basal bodies. Frontal cirri in life about 20 11m long, bases
slightly enlarged. All three postoral ventral cirri arranged close to buccal vertex (Fig. 96a,
c), according to KAHL (1932) posterior one (cirrus V/3) displaced somewhat posteriad
(Fig. 96e). Distance between posterior transverse cirrus and end of cell 3-6 11m (mean =
3.5 11m; n = 11), thus transverse cirri protrude distinctly beyond posterior end of cell. Left
marginal row J-shaped, nearly overlapping to right body margin. Right row almost
straight, terminates at level of pretransverse ventral cirri. Dorsal kinety 1 anteriorly al-
most unshortened, kinety 4 begins at mid-body. Invariably 3 caudal cirri.
296 SYSTEMATIC SECTION

o c cur r e nee and e colo g y: Not very common in terrestrial and freshwater
habitats. Locus classicus is the litter and upper soil layer of a very shallow soil of a goat
pasture between Nauplion and Tripolis, Peloponnesus, Greece (BERGER & FOISSNER
1987a). FOISSNER et al. (1985) found C. citrina in soils in the Tullnerfeld, Lower Austria.
KARL (1932) isolated his population from katharobic waters with Utricularia and Hotto-
nia. I found C. citrina in the detritus of an aquarium (24°C; together with Sty/onychia
pustu/ata, Holosticha monilata, Euplotes patella, Vorticella campanula, Mesodinium pu-
lex, Chilodonella acuta, Frontonia leucas, Urocentrum turbo), and between leaves in the
clean Illach River, Bavaria, Germany, in October (see FOISSNER 1997a). FOISSNER (1995b,
1997b, and pers. comm.) found C. citrina in soil from Japan, the Dominican Republic,
Costa Rica, and Peru.
Feeds on diatoms (Pinnularia sp.), flagellates (Anisonema sp.), fungi, and ciliates
(Cyclidium sp., Vorticella sp., Pseudocohnilembus sp.). Biomass of 106 individuals about
54 mg (FOISSNER 1987a).

Cyrtohymena australis FOISSNER, 1995 (Fig. 97a-m, Table 20)

1995 Cyrtohymena australis FO/sSNER, Arch. Protistenk., 145: 70.

T a x 0 nom y: This species, Cyrtohymena citrina, and C. primicirrata very likely form
a clade of their own within Cyrtohymena, having yellowish cortical granules as apomor-
phy. Autapomorphies of C. australis are the enormous size and the increased number of
dorsal kineties.

M 0 r p hoi 0 g y and b i 0 log y: In life 25~00 x 60-100 11m. Lanceolate, right

margin usually slightly concave, left approximately convex, widest in buccal area, gradu-
ally narrowed and broadly rounded at both ends. Dorso-ventrally flattened up to 2: 1.
Very flexible. Macronuclear nodules ellipsoid, in life about 45 x 20 11m, in middle third
of cell left of median, with many tiny nucleoli. 2-8 micronuclei (4-5 on average; n = 15),
almost spherical, often distant from macronuclear nodules, after protargol impregnation
~ x 2.7-3.5 11m (n = 15). Contractile vacuole slightly in front of mid-body at left mar-
gin, with 2 long collecting canals extending anteriorly and posteriorly. Cells yellow at
low magnification due to brilliant citrine colour of cortical granules. Granules globular,
0.4-1.0 11m across, arranged mainly around cirral bases and dorsal bristles but also in
loose rows between cirri and bristles (Fig. 97g, h, m), do not impregnate with protargol.
Cytoplasm colourless, with some fat globules, 2-4 11m across, and many small and large
food vacuoles. Moves rapidly to and fro.

Fig. 97a-i Cyrtohymena australis (from FOISSNER 1995b. a, b, g, h, from life; c-f, i, protargol impregnation). ~
a, b) Ventral and right lateral view, 305 Jim. c-t) Details of oral structures. The anterior portion of the paroral
consists of short kineties and has fan-shaped fibres attached (c). The adoral membranelles are of different size
 Cyrtohymena 297

~.:
~ proximal f

6':··0. 9

and their fine structure varies depending on the location within the membranellar zone. g) Surface view show-
ing arrangement of cortical granules (circles) on ventral side. b) Dorsal view of broad specimen showing pat-
tern produced by cortical granules. i) Ventral infraciliature. Arrowheads mark distal and proximal end of endo-
ral. Small arrows denote the rather inconspicuous pretransverse ventral cirri. Long arrow marks distal end of
adoral zone of membranelles, which is very near to right frontal cirrus (IW3). AZM = adoral zone of mem-
branelles, Ma = macronuclear nodule, Mi = micronucleus; p 296.
 298 SYSTEMATIC SECTION

, • ,
, ,
, ,, ('
,, ,, ('
,, ,/

, (

, ,
, ,
,,
,
, ,
1
?
, ,
, ,
, ,
, ,
, ,
,
#

, ,.

.-
f
f

••
k
Fig. 97j, k Cyrtohymena australis (from FOISSNER 1995b. j, k, protargol impregnation). Ventral and dorsal in-
fraciliature, 260 Jim. This specimen has only I pretransverse ventral cirrus. Note the increased number of dor-
sal kineties and the sometimes increased number of basal bodies per bristle unit. Arrow marks caudal cirri;
p296.

Adoral zone of membrane lIes about 34 % of body length, bases of largest mem-
branelies in life about 20 !lm wide. Buccal cavity large and deep. Paroral almost semicir-
cular with 2 fan-like bundles of fibres at anterior end, in large specimens composed of
oblique rows having 4-6 cilia each in anterior and dikinetids in posterior half. Endoral
hook-like, extends diagonally across buccal cavity and crosses (optically) paroral in pos-
 Cyrtohymena 299

Fig. 971, m Cyrtohymena australis (from FOISSNER 1995b. I, protargol impregnation; m, from life). I) Ventral
infraciliature of anterior third. m) Dorsal side of posterior third showing arrangement of cortical granules
within (arrows) and between (arrowhead) dorsal kineties. AZM = adoral zone of membrane\les, e = endoral,
Ma = macronuclear nodule, Mi = micronucleus, p = paroral, PVC = postoral ventral cirri; p 296.

terior third, very likely composed of dikinetids. Pharyngeal fibres inconspicuous. Frontal
cirri 20-25 ~m long, slightly (cirrus 11113) to distinctly (cirrus Ill) enlarged (Fig. 97j).
Usually 3, rarely only 2 postoral ventral cirri; posterior postoral ventral cirrus (V/3) ar-
ranged rather close to other postorals (Fig. 97i, j). Usually 2, rarely only 1 pretransverse
ventral cirrus. Transverse cirri (very rarely up to 7) in life 30 ~m long, displaced slightly
anteriorly. Marginal cirri in life 20-25 ~m long, rows distinctly separated posteriorly, gap
occupied by caudal cirri right of cell median. Marginal, transverse, and caudal cirri form
conspicuous fringe at posterior end. Dorsal cilia in life about 5 ~m long, arranged in 5-7
rows almost as long as body and few shortened rows at margins of anterior body half.
Bristle complexes composed of 2-6 basal bodies, only 1 or 2 of which are, however, cili-
ated (Fig. 97k).
300 SYSTEMATIC SECTION

Morphogenesis commences with the proliferation of basal bodies at the postoral ven-
tral cirri and the uppermost 2 transverse cirri (Costa Rica population; ForSSNER 1995b).

o c cur r e n c e and e colo g y: Probably widespread in soils of South America.

Locus classicus is soil from the Amazon rain forest near the town of Iquitos, Peru (Forss-
NER 1995b, 1997b). FOISSNER (1995b) found two other populations of C. australis, one in
Costa Rica (a tropical dry forest in Santa Rosa National Park, about 5 Ian east of the
ranch house "La Casona", near a small path to the Pacific Ocean) and another at Cura~ao,
a small island at the north coast of Venezuela, where it occurred in a rather saline sample
collected in a garden in the town of Salina.
Feeds on fungal spores, testate amoebae (Trinema lineare, Euglypha spp.), ciliates
(Leptopharynx costatus, Drepanomonas pauciciliata), and cysts of Polytoma sp. Bio-
mass of 106 individuals about 350 mg.

Cyrtohymena primicirrata (BERGER & FOISSNER, 1987) FOISSNER, 1989

(Fig. 20e, 98a-o, Table 20)

1970 Steinia platystoma EHRBo-STEIN, 1859 - DRAGESCO, Annis Fac. Sci. Univ. fed. Cameroun, (Numero
hors-serie): 119 (Fig. 98b-j; misidentification).
1984 Steinia inquieta (STOKES, 1887) - FOISSNER, Stapfia, 12: 115 (Fig. 98a-g; misidentification; 1 slide of
protargol-impregnated material is deposited in the OberOsterreichische Landesmuseum in Linz, Upper
Austria).
1987 Steinia primicirrata BERGER & FoISSNER, Zoo!. ]b. Syst., 114: 227, 228 (pro Steinia inquie/a sensu
FOISSNER 1984).
1987 Steinia ultricirrata BERGER & FOISSNER, 1987 - FOISSNER, Progr. Protistol., 2: 127 (nomen nudum; see
FOISSNER & FOISSNER 1988, P 91).
1989 Cyrtohymena primicirrata (BERGER & FOISSNER, 1987) - FOISSNER, Sber. Ost Akad. Wiss., 196: 239.

T a x 0 nom y: FOISSNER (1984) identified his population with cortical granules as

Steinia inquieta (STOKES), considering the redescription of KAlIL (1932) as decisive.
However, STOKES (1887b) mentioned neither a cortical granulation nor a colouring of the
cytoplasm in the original description of this species. Thus, BERGER & ForSSNER (1987a)
established S. primicirrata for S. inquieta sensu FOISSNER (1984). The infraciliature, espe-
cially the distinctly anteriorly displaced transverse cirri of Steinia platystoma sensu DRA-
GESCO (1970) agrees very well with the illustration by FOISSNER (1984; cpo Fig. 98f and
98i); although DRAGESCO (1970) did not mention a cortical granulation (very likely be-
cause he made no live observations), I consider the DRAGESCO and FOISSNER populations
as conspecific.

Fig. 98a-g Cyrtohymenaprimicirrata (from FOISSNER 1984. a-e, from life; f, g, protargol impregnation). a, c, ....,
e) Ventral, dorsal, and lateral view, a = 120 )lm, c = 115 )lm, e = 155 )lm. The yellow, 0.5-0.8 )lm sized corti-
cal granules are shown in (c) only. b) Cytoplasmic crystals. d) Pellicle with cortical granules. f, g) Ventral and
dorsal infraciliature, 85 )lm; p 300.
 Cyrtohymena 301

'-
--... '" "', \.
'-, \.
r..
\' (

..
-".

.•
"- \.

,
--, <..
\.
... ... '.. -' OJ

~.

'-

/.
<-...

" " " , ,

- .-
,
"- "-

-. -..
~
•
~~

" " ............

~.
• <'
"l: ~::.
-;-:: ':\: ~.

"
o ••
..:' ....
0 ••
'\<"
,..
".
.. ~

..<too ..
.;; .. ~" ~:: "';;:: ~<>: ~:

... 0;: ,,~. .'0

/0'

...
.~ .....
.,.0 • .. ~

..0
 302 SYSTEMATIC SECTION

• • •

. -.
.. . .. • •
•

•
••
••
• • •

Fig. 98h-j Cyrtohymena primicirrata (from DRAGESCO 1970. h-j, protargol impregnation). h-j) Ventral and
dorsal infraciliature, h,j = 75 11m, i = 70 11m; p 300.
 Cyrtohymena 303

Fig. 98k, I Cyrtohymena primicirrata (original scanning electron micrographs, kindly supplied by W. FOlSS-
NER). k) Ventral view, 95 11m. Arrow marks a cilium of dorsal kinety 1. I) Buccal area with prominent parorai;
endorai is arranged in deep buccal cavity (arrow). Arrowheads mark special cilia at right margin of adoral
membranelles, originally designated "paroral" cilia by STERK! (1878; see general section for details). Be = buc-
cal cirrus, p = paroral; p 300.

The granule-bearing Steinia inquieta sensu KAHL (1932) is classified as C. cUrina

(Fig. 96e). According to FOISSNER (1984), the pellicle of S. primicirrata is only slightly
flexible, which is atypical for a member of the Oxytrichinae. Thus, this character has to
be checked and morphogenetic data are needed to clarify the position of this species.

M 0 r ph 0 log y and b i 0 log y: In life about 90-130 x 35-50 ~m, 66-105 ~m af-
ter fixation (DRAGESCO 1970). Elliptical; slightly converging posteriorly, broadest at level
of buccal vertex, right margin convex, left usually slightly concave, both ends broadly
rounded. Macronuclear nodules in life about 20 x 12 ~m, after protargol impregnation
 304 SYSTEMATIC SECTION

1 3 ~
2 ' . 5

Fig.98m-o Cyrtohymena primici"ata (originals kindly supplied by W. FOISSNER. m, n, protargol impregna-

tion; 0, scanning electron micrograph). m, 0) Ventral and dorsal infraciliature. Note the supernumerary fron-
toventral cirrus (arrowhead) and the somewhat unusual arrangement of the postoral ventral cirri (cp. with Fig.
98t). Arrow marks posterior end of dorsal kinety 6. 0) Pellicle of ventral surface. The small pores are very
likely caused by ejected extrusomes (cortical granules). 1-5 = dorsal kineties 1-5; p 300.

10-15 x 7-9 11m. Usually 2 (rarely 3), after protargol impregnation 2.6-3.3 J-lm sized,
spherical micronuclei. Contractile vacuole with short collecting canals. Pellicle only
slightly flexible (check; see taxonomy). Cortical granules shiny yellow, so that specimens
appear distinctly yellowish even at low magnification; granules about 0.5-0.8 11m across,
usually arranged in groups of 3-10 along cirri, adoral membranelles, and especially
around dorsal cilia (Fig. 98c, d). Cytoplasm colourless, in posterior half of cell usually
many 1-7 11m sized yellowish cytoplasmic crystals.
Adoral zone of membranelles about 38 % of body length. Bases of largest mem-
brane1les in life about 8 11m broad. Pharyngeal fibres easily recognisable in life. Paroral
consists of 3 or 4 rows of basal bodies, terminates distally distinctly in front of endoral,
which consists of2 or 3 rows of basal bodies. From paroral many fibres run dorsally (Fig.
98f). Frontal and transverse cirri moderately enlarged, in life about 20 11m and 30 11m
 Cyrtohymena 305

long respectively. Five (very rarely 6) conspicuously anteriorly displaced transverse cirri,
that is, distance between posterior-most transverse cirrus and end of cell after protargol
impregnation 9-14 11m (mean = 11 11m; n = 9). Dorsal kinety 5 only slightly shortened
posteriorly, kinety 6 about 113 of body length. Invariably 3, in life about 20 11m long very
motile caudal cirri insert dorsally exactly in gap between marginal rows.

o c cur r e n c e and e colo g y: Rare in terrestrial and freshwater habitats. Locus

classicus is the upper soil layer (0-2 cm) of a flood plain, dominated by Phalaris au-
rundinacea and Phragmites australis in Vogelsang near the village of GrafenwOrth,
Lower Austria (BERGER & FOISSNER 1987a). A detailed description of this site is given by
FOISSNER et al. (1985; Profi15). DRAGESCO (1970) found his population in freshwater habi-
tats in Yaounde, Cameroon. Recently, W. FOISSNER (pers. comm.) isolated C. primicir-
rata from a pond in a forest near a lake (Grabensee) in Salzburg, Austria (Fig. 98k-o).
However, according to FOISSNER (1987a) C. primicirrata prefers terrestrial habitats. Feeds
on diatoms, fungal spores, testate amoebae (Trinema lineare), and ciliates. Biomass of
106 individuals about 77 mg (FOISSNER 1987a).

Cyrtohymena gracilis (KARL, 1932) FOISSNER, 1989 (Fig. 96f)

1932 Steinia graCilis KAHL, Tierwelt Dtl., 25: 614 (Fig. 961).
1989 Cyrtohymena gracilis (KAHL, 1932) - FOISSNER, Sber. Ost. Akad. Wiss., 196: 239.

Nom e n c I at u rea n d t a x 0 nom y: KAHL (1932) classified Steinia as subgenus

of Oxytricha; the correct name in his revision is thus Oxytricha (Steinia) gracilis. Differs
from other species with granulation by the single micronucleus and the conspicuous long
caudal cirri. Little-known species, thus detailed redescription required. Possibly a syno-
nym of C. muscorum.

M 0 r p hoi 0 g y and b i 0 log y: In life about 100 x 33 11m. Slender oval; some
(degenerative?) specimens with nearly parallel margins. Body flexible and slightly con-
tractile. Cortical granules brownish, arranged in conspicuous longitudinal rows. Cyto-
plasm often dark by oblong crystals. Moves rapidly to and fro, swims slightly bent, under
rotation about longer axis of cell. Eighteen frontal, ventral, and transverse cirri. All cirri
very long. Transverse cirri conspicuously enlarged, about 50 % of their length protrude
beyond posterior end of cell. Three caudal cirri 114-113 of body length.

o c cur r e n c e: Very rare in terrestrial and limnetic habitats. KAHL (1932) mentioned
no type location; he wrote "together with Steinia candens, but usually more abundant";
however, under S. candens he mentioned three locations, namely (i) Hamburg, Germany,
(ii) alpine mosses, and (iii) mosses from America. Records not substantiated by illustra-
tions: freshwater in Czechoslovakia (TIRJAKovA 1992b); small, eutrophic lake (Heiliges
Meer) in Germany (MOCKE 1979); mosses from a spring in Italy (STELLA 1947).
306 SYSTEMATIC SECTION

Cyrtohymena granulata (KARL, 1932) FOIssNER, 1989 (Fig. 96g)

1932 Steinia granulata lCAHL, Tierwelt Dtl., 25: 614 (Fig. 96g).
1989 Cyrtohymena granulata (KAHL, 1932) - FOlsSNER, Sber. Ost. Akad. Wiss., 196: 239.

N 0 ill e n c I a t u rea n d t a x 0 nom y: KAHL (1932) classified Steinia as subgenus

of Oxytricha; the correct name in the original description is thus 0. (Steinia) granulata.
Little-known species, detailed redescription necessary.

M 0 r p hoi 0 g y and b i 0 log y: In life about 100 x 25 llm. Margins usually al-
most in parallel, left often slightly convex. Moderately contractile. Each macronuclear
nodule with a single micronucleus. Cortical granules lacking. Cytoplasm always with
dark (often they appear greenish) globules, either near mid-body or in the periphery and
at posterior end of cell. Full set (18) of frontal, ventral, and transverse cirri; latter pro-
trude distinctly beyond posterior end of cell. Three caudal cirri, hardly longer than mar-
ginal cirri.

o c cur r e n c e: Locus classicus of this rather seldom species is Wisconsin, USA,

where KAHL (1932) found C. granulata very abundantly and persistently in a terrestrial
(?) moss carpet. MAnoNI & GHETII (1977; not substantiated by an illustration or descrip-
tion) counted 40 indo cm-2 in a heavily polluted running water in Parma, Italy, together
with Paramecium, Colpidium, and Glaucoma. Further record: soil of paddy fields in Ja-
pan (TAKAHASHI & SUHAMA 1991).

Cyrtohymena sapropelica (KARL, 1932) FOIssNER, 1989 (Fig. 96h)

1932 Steinia saprope/ica lCAHL, Tierwelt Dtl., 25: 613 (Fig. 96h).
1989 Cyrtohymena sapropelica (KAHL, 1932) - FOlsSNER, Sber. Ost. Akad. Wiss., 196: 239.

Nom e n c I at u rea n d t a x 0 nom y: KAHL (1932) classified Steinia as subgenus

of Oxytricha; the correct name in the original description is thus Oxytricha (Steinia)
sapropelica. Little-known species, detailed redescription necessary.

M 0 r p hoI 0 g y and b i 0 log y: In life about 100 !lm long. Outline somewhat
variable, usually slender oval and right anteriorly slanted; left body margin slightly con-
vex. Body moderately contractile. Without conspicuously coloured extrusomes (KAHL
1932), that is, cortical granules very likely absent. Adoral zone of membranelles short
with a high peristomial lip and high undulating membranes. Eighteen or 17 (?) frontal,
ventral, and transverse cirri; latter distinctly displaced anteriad so that they do not pro-
trude beyond posterior end of cell. Three distinctly elongated caudal cirri.
 Cyrtohymena 307

o c cur r e n c e and e colo g y: Very rare, limnetic. KAHL (1932) discovered C.

saprope/ica in the sapropel (see also MAUCH 1976); unfortunately, KAHL fIxed no type lo-
cation. CHORIK & VIKOL (1973) recorded it from a cooling plant in Moldova. Feeds on
ciliates (KAHL 1932).

Cyrtohymena torrenticola (SRAMEK-HuSEK, 1957) FOISSNER, 1989

(Fig. 96i)

1957 Steinia torrentico/a SRAMEK-HuSEK, Vl!st.~!. zoo!. SpoJ., 21: 4, 21 (Fig. 96i).
1989 Cyrtohymena torrenticola (SRAMEK-HusEK, 1957) - FOISSNER, Sber. Ost. Akad. Wiss., 196: 239.

T a x 0 nom y: Little-known species, thus detailed redescription necessary. The slender

outline is reminiscent of Urosoma, which has, however, a different oral apparatus.

M 0 r p hoi 0 g y and b i 0 log y: In life about 120 x 22 !lm. Anterior half of cell
with more or less parallel margins and broadly rounded, posterior half continuously con-
verging and very narrowly rounded. Macronuclear nodules in life about 21 !lm long, ar-
ranged slightly right of median. No cortical granules mentioned. Adoral zone of mem-
branelles about 35 % of body length. Three postoral ventral cirri, according to the fIgure,
the anterior-most is located in front of cytostome (misobservation?). Five transverse cirri
arranged rather close to posterior end of cell. According to SRAMEK-HuSEK (1957) three
short but conspicuous caudal cirri (obviously not shown in the fIgure).

o c cur r e n c e: Locus classicus is Cerny potok at Stare Vody, an oligo- or betameso-

saprobic brook of a river system ("Kleine March") in Czechoslovakia (SRAMEK-HUSEK
1957). Not found since.

Cyrtohymena sp. (Fig. 101h, i)

1932 Steinia candens KARL, Tierwelt Dt!., 25: 613, in part: Fig. 116 18,21; not Fig. 1205, 12b.

T a x 0 nom y: KAHL (1932) provided four illustrations of C. candens. In two of them

the body margins are more or less parallel and at least two micronuclei are present; they
are classifIed as C. candens. The remaining two forms are anteriorly slightly to distinctly
narrowed and clearly have only one micronucleus between the two macronuclear nodules
(Fig. 10lh, i). Because of the latter difference, especially, I separate the populations with
one micronucleus from C. candens. Detailed description necessary.

M 0 r p hoI 0 g y and b i 0 log y: In life about 150-180 !lm long. Anterior portion
slightly narrowed and sometimes inconspicuously curved to the left. Ratio of length to
308 SYSTEMATIC SECTION

width about 2.8: 1. Body very likely flexible. Macronuclear nodules left of median with
one micronucleus in between. Cortical granules lacking, cytoplasm colourless.
Adoral zone of membrane lIes about 40 % of body length. Transverse cirri protrude
distinctly beyond posterior end of cell. Dorsal cilia in life about 4 ~m long. Three dis-
tinctly elongated caudal cirri.

o c cur r e n c e: KAlIL (1932) found Cyrtohymena sp. in terrestrial mosses from the
Alps and from North America.

Cyrtohymena aestuarii (MARGALEF LOPEZ, 1945) comb. nov. (Fig. tole, f)

1901 Oxytricha plarystoma EHRao. - Roux, Mem. Inst. natn. genev., 19: 102 (Fig. 10 Ie; misidentification).
1945 Oxytricha (Steinia) candens KAHL Vat. aestuarii MARGALEF L6PEZ, Boln R. Soc. esp. Hist. nat., 43: 377
(Fig. IOlf).

Nom e n c I a t u rea n d t a x 0 nom y: As the name for the variety was published
before 1961, the change in rank does not affect the authorship (IeZN 1985; Articles 45 g
(ii), 50 c (i)). Both populations have only 4 transverse cirri, suggesting identity with C.
tetracirrata. However, this species is very likely confined to terrestrial habitats (FOISSNER
1987a, own observations), whereas the Raux (1901) and MARGALEF LoPEZ (1945) popula-
tions were from fresh and brackish water respectively; I thus avoid a synonymisation. De-
tailed redescription necessary.

M 0 r ph 0 log y and b i 0 log y: After MARGALEF LOPEZ (1945) in life about

200 ~m long, according to Raux (1901) only 90-110 x 28-33 ~m. Body margins more
or less in parallel, both ends broadly rounded. Slightly flexible. Cortical granules not
mentioned. Cytoplasm colourless, with many spherical inclusions. Adoral zone of mem-
branelles about 40 % of body length. Buccal area large. Arrangement and number of cirri
in frontal area obviously incorrectly observed by MARGALEF LOPEZ (1945; Fig. 101t).
Transverse cirri protrude distinctly beyond posterior end of cell. About 17 right and
13-14 left marginal cirri, which is rather low for such a large species (misobservation?).

o c cur r e n c e and e col 0 g y: Locus classicus is a saline pond in the estuary of

the Lobregat River in Barcelona, Spain (MARGALEF LOPEZ 1945). Raux (1901) found his
population in a clean, stagnant water body near Geneva, Switzerland. Probably feeds on
protozoa.

Cyrlohymena candens (KAHL, 1932) FOISSNER, 1989 (Fig. 99a-m, Table 20)

1932 Steinia candens KAHL, TierweltDtl., 25: 613, in part: Fig. 1205, 12h; not Fig. 116,8.2, (Fig. 99g, h).
1966 Steinia simplex DRAGESCO, Protistologica, 2: 88 (Fig. 99f; new synonym).
 Cyrtohymena 309

--.
1

(1

'"
..."
.A

, .A
....
, f'

..
A
".,

, ..
"
• ,.
c •• • d e

Fig. 99a--e Cyrtohymena candens (from FOISSNER 1982. IK, from life; d, e, protargol impregnation). a-c)
Ventral, dorsal, and lateral view showing, among other things, contractile vacuole with collecting canals (b), a,
b = 135 J.lm. d, e) Ventral and dorsal infraciliature, d = liS J.lm, e = 120 J.lm. CC = caudal cirri; P 308.

1975 Sieinia candens KAm., 1932 - GROUERE, Protistologica, II: 495 (Fig. 99i).
1977 Oxytricha platysloma EHRB. - BUITKAMP, Acta Protozool., 16: 272 (Fig. 99j; misidentification).
1982 Steinia candens KAm., 1932 - FOISSNER, Arch. Protistenk., 126: lOS (Fig. 991H:; authoritative rede-
scription; 1 slide of protargol-impregnated material is deposited in the OberOsterreichische Landesmu-
seum in Linz, Upper Austria).
1986 Steinia candens KAm., 1932 - DRAGESCO & DRAGESCo-KERNas, Faune tropicaIe, 26: 481, in part: Fig.
142F,G,J; not Fig. 142H,1.
1987 Steinia candens KAm., 1932 - BERGER & FOISSNER, Zool. Jb. Syst., 114: 227, 228.
1989 Cyrtohymena candens (KAm., 1932) - FOISSNER, Sber. Ost. Akad. Wiss., 196: 239.
1989 Cyrtohymena simplex (DRAGESCO, 1966) - FOISSNER, Sber. Ost. Akad. Wiss. 196: 239.
1994 Cyrlohymena candens - FOISSNER, Katalog des OberOsterreichischen Landesmuseums Linz, 71: 192
(colour-micrograph of a protargol-impregnated specimen).

Nom e n cIa t u rea n d t a x 0 nom y: KAHL (1932) classified Steinia as subgenus

of Oxytricha; the correct name in the original description is thus Oxytricha (Steinia) can-
dens. "Oxytricha caudens KAHL" in CAIRNS (1966a, p 61), NING et al. (1993, p 2), and
YANG (1989, P 157) is an incorrect subsequent spelling.
Synonymy with C. inquieta cannot be excluded; see relevant chapter in C. inquieta
for discussion of this problem. Steinia simp/ex is obviously somewhat superficially de-
scribed (Fig. 99f). In my opinion, DRAGESCO (1966b) erroneously interpreted a pretrans-
310 SYSTEMATIC SECTION

.,
·
O: ,-
•
•
It

Fig. 99f-j Cyrtohymena candens (f, from DRAGESCO 1966b; g, h, from KAHL 1932; i, from GROLIERE 1975; j,
after BUITKAMP 1977b. f, i,j, protargol impregnation; g, h, from life). C, i, j) Ventral infraciliature, C= 215 ~m, i
= 21 0 ~m, j = ISO ~m. Arrowheads in G) mark caudaI cirri. Ciliary pattern (for example, lack oC postoraI ven-
traI cirri) in (f) unreliable. g, h) VentraI views, g = 180 ~m, h = ISO ~m; p 308.

verse ventral cirrus as sixth transverse cirrus, and furthermore he very likely overlooked
the anterior two postoral ventral cirri.
The undulating membranes of Steinia platystoma sensu BUITKAMP (1977b) show the
typical Cyrtohymena pattern; since BUITKAMP did not mention a cortical granulation and
because of the size (200-250 !!m), I synonymise it with C. candens (Fig. 99j). The rede-
scriptions of CHARDEZ (1971, 1981; Fig. 223d, 2341) are insufficient. KAHL (1932) pro-
vided four illustrations of the present species; two of these are anteriorly convergent and
have only one micronucleus, indicating that they are a different species (see Cyrtohymena
sp.;Fig.lOlh,i).
 Cyrtohymena 311

Fig. 99k Cyrtohymena candens (original scanning electron micrograph, kindly supplied by W. FOISSNER).
Buccal area with prominent, semicircularly curved paroral running proximally between two distinct folds (ar-
rowheads). BC = buccal cirrus, e = endoraI, FC = frontal cirri, p = paroraI; p 308.
 312 SYSTEMA TIC SECTION

Morphology and biology:

In life 150-250 x 60-80 J1m; 150-200 x
60-80 J1m (KAHL 1932), 185-260 x 55
to 65 J1m (Bouin fIxed specimens; GRO-
L1ERE 1975), about 150 J1m long (FoISs-
NER 1982), 200-250 J1m long (in life?;
BUITKAMP 1977b). Length of synonym,
S. simp/ex in life (?) 240 J1m on average
(DRAGESCO 1966b). Broad elliptical,
margins sometimes nearly in parallel,
left margin often straight, right convex.
Both ends broadly rounded. According
to I<AHL (1932) slightly contractile, ac-
cording to FOISSNER (1982) very flexible.
Macronuclear nodules after protargol
impregnation 35 Jim long (GROLIERE
1975). 2-5 spherical micronuclei. Con-
Fig. 991, m Cyrtohymena candens (original scanning tractile vacuole with long collecting ca-
electron micrographs, kindly supplied by W. FOISS- nals, excretion pore slit-like (Fig. 99m,
NER). Ventral and dorsal side. Arrow in (m) marks ex- arrow). Cortical granules lacking, cyto-
cretion pore of contractile vacuole. b = dorsal kineties, plasm colourless, according to I<AHL
p = curved parora1; p 308.
(1932) shiny due to many cytoplasmic
crystals. Movements usually hasty, stays
close to soil particles.
Adoral zone of membrane lies between 35 % (Fig. 99d) and 55 % (Fig. 99i) of body
length; the latter value is very high and rather atypical for a member of the Oxytrichinae.
About 45 adoral membranelles (BUITKAMP 1977b). Buccal area very wide and deep. Ac-
cording to BUITKAMP (1977b) frontal cirri consist of 10 x 12 cilia. About 16-23 left and
17-28 right marginal cirri. After FOIsSNER (1982) and according to scanning electron mi-
crographs (W. FOISSNER, pers. comm.) 6 dorsal kineties, according to GROLIERE (1975)
 Cyrtohymena 313

and BUITKAMP (1977b) only 5 (Fig. 9ge); possibly they overlooked the rather short kinety
6. Three distinctly elongated caudal cirri.
Resting cyst spherical, with 3-5 !.lm long spines, about 40 !.lm across. Cyst wall dis-
tinctly yellowish (W. FOISSNER, pers. comm.). ALoNSO (1975) and ALoNSO & PEREZ-SILVA
(1966a, b, 1967, 1969) studied the polytene chromosomes in the macronuclear anlage.

o c cur r e n c e and e colo g y: Common in soil and in benthic freshwater habitats.

Locus classicus is Volksdorf, Hamburg, Germany, where KAlIL (1932) discovered C.
candens in mosses from a shaded wall. Records substantiated by illustrations: soil of al-
pine pastures in the Glockner area, Austria (FOISSNER 1982, FOISSNER & FOISSNER 1988);
puddles at the shore of Lake Geneva (= locus classicus of the synonym, Steinia simp/ex)
and pond in France (DRAGESCO 1966b, GROLIERE 1975); soil of the savannah of Larnto,
Ivory Coast (BUITKAMP 1977b); bark of a tree near Ross Highway, about 50 km east of
Alice Springs, Australia (BLATIERER & FOISSNER 1988; Fig. 99k-m are also from an Aus-
tralian population); soil in Tasmania (FOISSNER 1997b).
Records not substantiated by illustrations: Belgium (CHARDEZ 1987); rivers in Bul-
garia (DETCHEVA 1981, 1991); agricultural soil, submerged and terrestrial mosses, and
freshwater from Czechoslovakia (TIRJAKovA 1988, 1992b, TIRJAKovA & MATIS 1987a, b);
soil of a beech wood in Denmark (STOUT 1968); in Staufen and in a pond in Germany
(ALoNSO & PEREZ-SILVA 1966a, MOCKE 1979); beta- to alphamesosaprobic running waters
in Italy (MAnoN! 1980, MAnoN! & GHETTI 1981a, STELLA & CAMPEA 1948); Segovia and
Lobregat River, Spain (ALoNSO & PEREZ-SILVA 1966a, b, GRACIA et al. 1989); freshwater
habitats in the Tibetan Plateau (WANG 1977); China (NING et al. 1993, YANG 1989); soil
in Costa Rica (FOISSNER 1995b); Rio Bella, Peru (CAIRNS 1966a); saline soils south-east of
Adelaide, Australia (POMP & WILBERT 1988).
Feeds on heterotrophic flagellates, amoebas, and ciliates (FOISSNER 1982, GROLIERE
1975). Biomass of 106 individuals about 150 mg (FOISSNER 1987a). DETCHEVA (1983a)
found C. candens in oligosaprobic areas of the Maritza River, Bulgaria, under following
conditions: 0.3 % frequency, 17.4 °C, pH 7.4, 8.2 mg I-I O2 (87 % saturation), 49 mg 1-1
Ca2+, 6.3 mg I-I Mg2+, 192 mg I-I HC03-, 10 mg I-I SOl, 12.5 mg I-I Cl-, 0.05 mg I-I NH/,
0.4 mg }-1 N0 3-, 0.12 mg 1-1 N02-, 0.02 mg 1-1 Fe2+. For autecological data of terrestrial
populations, see FOISSNER (1981, 1985a), FOISSNER & PEER (1985), and FOISSNER et al.
(1985).

Cyrtohymena in quieta (STOKES, 1887) FOISSNER, 1989 (Fig. lOOa, b, 101~)

1859 Oxytricha platystoma. EHRBG. - STEIN, Organismus der Infusionsthiere I, p 190, in part: Tafel XII, Fig.
16,18; not Fig. 17, 19,20 (Fig. 100a, b).
1887 Histrio inquietus STOKES, Ann. Mag. nat. Hist., 20: 113 (Fig. lOla).
1888 Histrio inquietus STOKES - STOKES, 1. Trenton nat. Hist. Soc., 1: 296.
1972 Oxytricha inquieta (STOKES, 1887) KAHL, 1932 - BORROR, J. Protowol., 19: 13.
314 SYSTEMATIC SECTION

1974 Oxytricha candens (KARL) - PATSCH, Arb. Inst. landw. Zool. Bienenkd., 1: 62 (Fig. lOIc; very likely a
misidentification).
1975 Steinia macrostoma GROLIERE, Protistologica, 11: 495 (Fig. IOIb; new synonym).
1987 Steinia inquieta (STOKES, 1887) - BERGER & FOISSNER, Zool. Jb. Syst., 114: 227, 228.
1989 Cyrtohymena inquieta (STOKES, 1887) - FOISSNER, Sber. Ost. Akad. Wiss., 196: 239.
1989 Cyrtohymena macrostoma (GROLIERE, 1975) - FOISSNER, Sber. Ost. Akad. Wiss., 196: 239.

Nom e n c I a t u rea n d t a x 0 nom y: "Stienia macrostoma" in GROLIERE (1975, P

481) is an incorrect subsequent spelling of Steinia. KAHL (1932) provided C. inquieta
with brownish cortical granules, which is unjustified because STOKES (1887b) mentioned
neither cortical granules nor a colouring of the cytoplasm. Thus, C. inquieta sensu KAHL
(1932) is classified as C. citrina (Fig. 96e). Accordingly, C. inquieta is a species without
cortical granulation which differs from C. candens only in body size (BERGER & FOISSNER
1987a); Cyrtohymena inquieta is less than 150 11m long, C. candens becomes 150 to
250 11m long. This separation is of course somewhat arbitrarily and synonymy cannot be
excluded. However, there is further a character indicating the validity of these two spe-
cies, namely the absence of one postoral or pretransverse ventral cirrus in C. inquieta
(Fig. 10 I a). This is confirmed by protargol impregnation of Steinia macrostoma, which is
thus considered as new synonym of C. inquieta (Fig. 10 Ib).
Neither STOKES (1887b) nor GROLIERE (1975) wrote anything about the flexibility of
the cell. STOKES (1887b) assigned it to Histrio, indicating that C. inquieta has a rigid body
like Histriculus or Steinia. This would correlate with other apomorphies of the Stylony-
chinae, namely the long adoral zone of membranelles (about 50 % of body length) and
the lack of cortical granules. The micrograph of a protargol-impregnated specimen in
GROLIERE (1975; his Fig. 26) shows a distinctly curved parora! (unfortunately no distinct
endoral), indicating that the oral apparatus is Cyrtohymena- and not Steinia-like; thus, it
is not transferred to Steinia. Detailed reinvestigation needed.
Oxytricha platystoma sensu Gu & ZHANG (1992) is very likely identical with C. in-
quitea. STEIN (1859b) obviously confused Steinia platystoma (Fig. 172i, j) and a Cyrtohy-
mena species which is preliminarily classified as C. inquieta (Fig. I OOa, b).
Steinia candens var. depressa GELLERT and S. ovalis REUTER are preliminarily treated
as supposed synonyms of C. inquieta because they are less than 150 11m long (Fig. lOld,
g). It is impossible to separate them reliably from C. inquieta or C. candens with the data
available. Detailed investigations of populations from the type locations are needed to
elucidate the taxonomic status of these little-known species.
Steinia inquieta sensu REUTER (1963a) from rain-wet mosses is preliminarily classi-
fied as C. muscorum because it has dark brown granules (Fig. 94m). The DINGFELDER
(1962; Fig. 223c) and VUXANOVICI (1961; Fig. 234f-h) redescriptions of C. inquieta are
insufficient.

M 0 r ph 0 log y and b i 0 log y: In life about 95 !lm (STOKES 1887b), after protar-
gol impregnation about 90!lm long (PATSCH 1974), according to GROLIERE (1975)
95-140 x 30-50!lm (mean = 110 x 43 !lm) after protargol impregnation. Body margins
 Cyrtohymena 315

distinctly converging posteriorly, ante-

rior end broadly rounded. Macronu-
clear nodules after protargol impregna-
tion about 14 Ilm long, arranged
slightly left of median. At least two, af-
ter protargol impregnation about 4 Ilm
sized spherical micronuclei. Cortical
granules lacking!
Adoral zone of membrane lies
about 50 % of body length (see taxon-
omy), consists of 45-50 membranelles.
Buccal area very large. Frontal and
transverse cirri distinctly enlarged.
Both STOKES (1887b) and GROLIERE
(1975) draw only 4 postoral and pre-
transverse ventral cirri, possibly a typi-
cal character of C. inquieta (Fig. 10 1a,
b); however, the STEIN (1859b; Fig.
100a, b) and PATSCH (1974; Fig. 101c)
illustrations do not confirm this peculi-
arity, indicating that these populations
are not conspecific. About 20 right and
14-16 left marginal cirri. According to Fig. lOOa, b Cyrtohymena inquieta (after STEIN
1859b). Ventral and dorsal view of ventral ciliature
GROLIERE (1975) only 5 dorsal kineties
from life, sizes not indicated; p 313.
and no caudal cirri; probably he over-
looked the usually rather short outer
dorsomarginal row and misinterpreted
the caudal cirri as marginal cirri.

o c cur r e n c e and e colo g y: Very likely confined to freshwater habitats. Locus

classicus is a pond with Lemna in North America, probably near Trenton, New Jersey,
where STOKES lived. GROLIERE (1975, 1977, 1978) discovered the synonym, Steinia mac-
rostoma, in a pond in France. PATSCH (1974) found it in small running waters near Bonn,
Germany. Records not substantiated by illustrations: terrestrial mosses from Belgium
(CHARDEZ 1983, 1987); once in the Bulgarian Strouma River (DETCHEVA 1979c; incor-
rectly spelled as Steinia inqueta); freshwater habitat in the Tibetan plateau (WANG 1977);
Brush Lake, Ohio, USA (RIDDLE 1905); Pantanal flood plains, Brazil (HARoIOM & HECK-
MAN 1996). Feeds on bacteria, flagellates, and ciliates (GROLIERE 1975).

Sup p 0 sed s y non y m s: Steinia candens var. depressa GELLERT, 1942, Acta Scien-
tiarum Mathematicarum et Naturalium, 8: 28 (Fig. IOld). Remarks: GELLERT (1942)
found this 100-120 Ilm sized population without cortical granules in the algae layer of
 316 SYSTEMATIC SECTION

Fig. lOla--c Cyrtohymena inquieta (a, after STOKES 1887b; b, from GROLIERE 1975; c, from PATSCH 1974. a,
from life; b, c, protargol impregnation). a) Ventral view, 70 llm. Note the lack of one postoral ventral or pre-
transverse ventral cirrus, which is confirmed by protargol impregnation (Fig. 1OIb). Possibly, this species be-
longs to Steinia, as indicated by the long adoral zone of membranelies and the rigid body (see text). b, c) Ven-
tral infraciliature, b = 115 lllll, c = 90 llm; p 313. d Cyrtohymena candens var. depressa, a supposed synonym
ofe. inquieta (from GELLERT 1942). Ventral infraciliature after sublimate fixation and opalblue staining after
 Cyrtohymena 317

tree bark in Hungary. Buccal cavity less broad than in C. inquieta and C. candens. Syn-
onymy with C. inquieta uncertain because full set (3) of postoral ventral cirri present.
FOISSNER (1989, p 239) transferred it to Cyrtohymena: C. candens var. depressa
(GELLERT, 1942) FOISSNER, 1989.

Steinia ova/is REUTER, 1961, Acta zool. fenn., 99: 19 (Fig. 101g). Remarks: FOISSNER
(1989, p 239) transferred it to Cyrtohymena, C. ova/is (REUTER, 1961). This species is
reminiscent of C. inquieta and C. candens; because it is less than 150 ).lm long, I assign it
to the C. inquieta group, which has, however, only 2 postoral ventral cirri. In life about
130-140).lm long; slender oval; adoral zone ofmembranelles about 1/3 of body length;
peristomial lip anteriorly rectangularly curved; only the three posterior-most transverse
cirri protrude beyond posterior end of cell; probably 3 slightly enlarged caudal cirri. Lo-
cus classicus is the groundwater from the northern sandy beach of the Langskllr Island in
Finland. REUTER (1961) found it 0.1-0.2 m (?) from the sea, indicating that S. ovalis is a
marine interstitial form. Not found since.

Cyrtohymena tetrac;rrata (GELLERT, 1942) FmssNER, 1989 (Fig. 102a-k,

Table 20)

1942 Steinia tetracirrata GELLERT, Acta Scientiarum Mathematicarum et NaturaIium, 8: 25 (Fig. 102a, b).
1972 Oxytricha tetracirrata GELLERT, 1942 - BORROR, 1. Protozoot., 19: 14.
1987 Steinia tetracirrata GELLERT, 1942 - BERGER & FOISSNER, Zoot. Jb. Syst., 114: 225 (Fig.l02c, e, h, i;
authoritative redescription; 1 slide of protargol-impregnated specimens is deposited in the OberOsterrei-
chische Landesmuseum in Linz, Upper Austria).
1989 Cyrtohymena tetracirrata (GELLERT, 1942)- FOISSNER, Sber. ()st. Akad. Wiss., 196: 239.

T a x 0 nom y: Very likely C. tetracirrata is confined to terrestrial habitats. Oxytricha

candens var. aestuarii MARGALEF L6PEZ, 1945 and o. platystoma sensu Roux (1901) also
have only 4 transverse cirri (Fig. 101e, f). However, they were collected from brackish
and, respectively, freshwater habitats, which is the main reason I do not synonymise them
with C. tetracirrata. The non-terrestrial populations are preliminarily classified as C. ae:;-
tuarii MARGALEF LOPEZ, 1945.
Cyrtohymena tetracirrata has an almost rigid body, a long adoral zone (40 % of
body length), and cirrus V/3 is distinctly displaced posteriad indicating that it is not in-
volved in primordia formation. These character states suggest that it is (like C. quadrinu-
cleata and C. inquieta) not an Oxytrichinae, but a Stylonychinae; however, morphoge-
netic and molecular biological data are needed to verify this hypothesis.

~ BRESSLAu, 100 11m; p 315. e, f Cyrtohymena aestuarii (e, after Roux 1901; f, after MARGALEF L6PEZ 1945).
Ventral views in life, e = 95 11m, f= 210 11m; p 308. g Cyrtohymena ovalis, a supposed synonym ofe. inqui-
eta (after REUTER 1961). Ventral view in life, about 130 11m; p 317. b, i Cyrtohymena sp. (from KARL 1932).
Ventral views in life, h = 180 11m, i = 150 11m. Note single micronucleus between macronuciear nodules;
p 307.
318 SYSTEMATIC SECTION

,
~

,
~

,
,

!
I ;
e !. I
~ I "
" :/ '.
:~ "
, ,.'
: I
'.
,
\ /
" (
':
I
\

,
# • # ':
':
,
I
I

I
I

.,
,
# #
,
\
( :'
# \ :
I

#
# •.t , ,
: I
I
:

,
\

•••
\
I
9 # ,I
\ ,
/ ,I
,I
••
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,I I
~ '.

..• ,
I '
!. ( .'
1. f. ( I
II'
h
 Cyrtohymena 319

Fig. 102j, II. Cyrtohy-

mena tetracirrata
(originals, kindly sup-
plied by W. FOISSNER).
j) Ventral infracilia-
ture after protargol im-
pregnation. k) Cyst in
life, bright field;
p 317.

j k

M 0 r ph 010 g y and b i 010 g y: In life about 150-170 x 60-70 J.1m, according to

GELLERT (1942, 1956) only 90-110 J.1m long. Body margins more or less parallel, right
one sometimes slightly convex; some specimens posteriorly slightly broader than anteri-
orly; both ends broadly rounded. Body almost inflexible and fragile, according to
GELLERT (1942) flexible. Macronuclear nodules in life about 20-30 x 14-16 J.1m, after
protargol impregnation 14-18 x 10-11 J.1m (mean = 17 x 10 J.1m; n = 8), arranged about
in median of cell (Fig. 102a, f, i). Distance between macronuclear nodules after protargol
impregnation 7-18 J.1m (mean = 13 J.1m). Nucleoli large, easily recognisable even in life.
2-3, in life about 5 J.1m sized, spherical micronuclei. Contractile vacuole with distinct col-
lecting canals. Cortical granules lacking. Cytoplasm colourless, many about 12-17 J.1m
sized food vacuoles, some with crystalline content.
Adoral zone of membranelles about 40 % of body length. Bases of largest mem-
branelles in life about 11-13 J.1m broad. Undulating membranes consist of 3--4 rows of
basal bodies. Frontal cirri distinctly enlarged, in life about 25-30 J.1m long. Arrangement
of frontoventral cirri usually as in Figures 102a, f, sometimes cirrus VII3 slightly dis-
placed posteriad (Fig. 102h, arrow). Invariably 4, in life about 30 J.1m long, slightly to dis-
tinctly enlarged transverse cirri (Fig, 102a, f, h, j). Marginal cirri in life about 20-25 J.1m
long, left row terminates in median of cell, right one ends slightly subterminally. Dorsal

+- Fig.l02a-i Cyrtohymena tetracirrata (a, b, after GELLERT 1942; c, e, h, i, from BERGER & FOISSNER 1987a; d,
f, g, originals. a, b, sublimate fixation and opalblue staining; c-e, g, from life; t: h, i, protargol impregnation).
a, b, C, h, i) Ventral and dorsal infraciliature, a, b, h, i = 80 J.Im, f= 105 J.Im. Arrowheads in (f) mark the 3 cau-
dal cirri, arrow in (f, h) denotes frontoventral cirrus Vl/3, which is in a somewhat unusual position in (h). c-e)
Ventral, dorsal, and right lateral view, c = 160 J.Im, d, e = 150 J.Im. g) Part of the cyst wall; height of protuber-
ances = 2.5-4 J.Im; p 317.
320 SYSTEMATIC SECTION

kinety 4 somewhat shortened anteriorly, kinety 5 about half of body length, kinety 6 con-
sists of about 6 basal body pairs (Fig. 102i). Usually 3, rarely 4, in life about 30 ~m long
caudal cirri on dorsal kineties I, 2, and 4; arranged dorsally in gap between marginal
rows (Fig. 102f, i).
Resting cyst spherical, with many 2.5--4.0 ~m high protuberances (Fig. 102g, k). Dia-
meter without tubercles 42-56 ~m. Cyst wall 1-2 ~m thick, distinctly yellowish. Cyto-
plasm slightly yellowish.

o c cur r e n c e and e colo g y: Probably confmed to terrestrial habitats. Locus

classicus is a tree bark grown with algae (Pleurococcus vulgaris) in Hungary (GEILERT
1942). Unfortunately, the paper is written in Hungarian so that I could not fmd the sam-
ple site; possibly it is near Tihany, where GEll.ERT lived and worked. GELLERT (I 956)
found C. tetracirrata in mosses and tree barks near the village of Boldogk5varalja, Hun-
gary, and BERGER & FOISSNER (1987a) in a salt soil with rush, about 50 m from the sea in
Nauplia Bay, Greece. I found it in compost heaps near the village of Garsten (Upper Aus-
tria) and Salzburg (Fig. 102d, t) and in terrestrial mosses from a gorge (Glasenbach-
klarnm) in Salzburg, Austria.
Feeds on bacteria, flagellates (Anisonema sp.), and ciliates. Can be cultured in Eau
de Volvic (France) with yeast as food. Biomass of 106 individuals about 150 mg (FOISS-
NER 1987a).

Cyrtohymena quadrinucleata (DRAGESCO & NJINE, 1971) FOISSNER, 1989

(Fig. 21c, 103a-f, Table 20)

1971 Steinia quadrinucleata DRAGESCO & NJINE, Annis Fac. Sci. Univ. fed. Cameroun, 7-8: 129 (Fig.
103a).
1984 Steinia quadrinucleata DRAGESCO & NJINE, 1971 - FOISSNER, Stapfia, 12: ll8 (Fig. 103b-f; authorita-
tive redescription; 1 slide of protargol-impregnated specimens is deposited in the OberOsterreichische
Landesmuseum in Linz, Upper Austria).
1986 Steinia quadrinucleata DRAGESCO et NJINE, 1971 - DRAGESCO & DRAGESCO-KERNElS, Faune tropicale,
26: 482.
1989 Cyrtohymena quadrinucleata (DRAGESCO & NJINE, 1971) - FOISSNER, Sber. Ost. Akad. Wiss., 196: 239.

Nom e n c I at u rea n d t a x 0 nom y: Steinia quadrinucleatus in NJINE (1977,

P 232) is an incorrect subsequent spelling. DRAGESCO & NJINE (1971) and DRAGESCO &
DRAGESCO-KERNEIS (1986) mentioned a third undulating membrane, which is a misobser-
vation and/or a misinterpretation of the protargol preparations. Some character states
(body almost rigid, adoral zone of membranelles more than 40 % of body length) indicate
that Cyrtohymena quadrinucleata (and c. tetracirrata and C. inquieta) does not belong
to the Oxytrichinae, but to the Stylonychinae. Furthermore, the posterior-most postoral
ventral cirrus (V/3) is possibly not involved in primordia formation (according to an un-
published scanning electron micrograph of a population from Namibia, kindly supplied
 Cyrtohymena 321

Fig.l0Ja-d Cyrtohymena quadrinucleata (a, from DRAGESCO & NIINE 1971; b-d, from FOISSNER 1984. a, pro-
targol impregnation; b-d, from life). a) Ventral infraciliature, 105/lm. b-d) Ventral, dorsal (with contractile
vacuole), and lateral view, b = 140 /lm; p 320.

by W. FOISSNER), another, certainly plesiomorphic character of the Stylonychinae (Fig.

25a, b); however, detailed morphogenetic and molecular biological data are needed for
fmal classification.

M 0 r p hoI 0 g y and b i 0 log y: In life about 140 x 55 !lm (FOISSNER 1984). Or-
thogonal, anteriad often slightly converging; both ends broadly rounded. Pellicle only
slightly flexible (FOISSNER 1984; see taxonomy). Macronuclear nodules in life
15-20 x 10 !lm, after protargol impregnation 9-16 x 7-10 !lm (n = 10), sometimes ar-
ranged in 2 indistinct pairs left of median (Fig. 103b). 2-4 in life about 3 !lm sized,
spherical micronuclei. Contractile vacuole without conspicuous collecting canals. Pellicle
and cytoplasm colourless. Cortical granules lacking. Posterior half of cell usually with
322 SYSTEMATIC SECTION

) }
}
i\
}
l . i
) {
) }
) }

)
~
,
,,
}
) l
)
)
'I (
,
,
(

*
-* *
~ ('

-
~ (

#
* i
~
(

i
* --
:'

•
•,.••
.---
.•
~ :'

~
I r
# ,.. I. h

# (

• • •
('
#
'"'
e

Fig. t03e, f Cyrtohymena quadrinucleata (from FOISSNER 1984. e, t: protargo\ impregnation). Ventral and
dorsal infraciliature, 80 J.lm; p 320.

nwnerous 0.5-5.0 11m sized, yellowish crystals, so that specimens appear dark at low
magnification and bright field illwnination. Moves rapidly to and fro.
Adoral zone of membrane lies about 47 % of body length (see taxonomy). Bases of
largest membranelles in life about I 1 11m broad. Pharyngeal fibres easily recognisable in
life. Paroral consists of 3 or 4 rows of basal bodies, with many dorsally running fibres
(Fig. I03e); endoral composed of 2 or 3 rows. Frontal cirri distinctly enlarged, in life
about 20 /lm long. Transverse cirri slightly enlarged, in life about 28 /lm long, distinctly
protruding beyond posterior end of cell, somewhat fringed distally. Marginal cirri in life
15 /lm long, rows not confluent posteriorly. Dorsal kineties 3 and 4 slightly shortened an-
 Cyrtohymena 323

teriorly. Three, in life about 25 ~m long, enlarged caudal cirri; arranged dorsally, almost
exactly above gap of marginal cirri (Fig. 103 f).

o c cur r e n c e and e colo g y: Not vel)' common in terrestrial and freshwater

habitats. Locus classicus is a leaf-littered pond near Yaounde, Cameroon (DRAGESCO &
NJINE 1971; see also Noo 1977). ForssNER (1984) found C. quadrinucleata in the upper
soil layer of a conventionally farmed field in the town of Salzburg, Austria. W. ForssNER
(pers. comm.) found it also in soils from Namibia and Antarctica. Further record: soil of
rain forest in Brazil (FOISSNER 1997b). Biomass of 106 specimens about 81 mg (FOISSNER
1987a). Feeds on ciliates.

Cyrtohymena marina (KARL, 1932) FOISSNER, 1989 (Fig. 96j)

1932 Steinia marina KAHL, Tierwelt Dtl., 25: 614 (Fig. 96j).
1989 Cyrtohymena marina (KAHL, 1932) - FOISSNER, Sber. liSt. Akad. Wiss., 196: 239.

Nom e n c I a t u rea n d t a x 0 nom y: KAm.. (1932) classified Steinia as subgenus

of Oxytricha; the correct name in the original description is thus Oxytricha (Steinia) ma-
rina, which is a primary homonym of Oxytricha (Oxytricha) marina KAm.., 1932; the lat-
ter received a replacement name. Little-known species; detailed redescription thus neces-
sary.

M 0 r ph 0 log y: Length in life 100--150 ~m. Long elliptical, that is, 4-5 times as long
as broad. Flexible and soft, slightly contractile. Cortical granules sparse and delicate, ar-
ranged in short rows (colour? probably inconspicuous). Adoral zone of membranelles
about 114 of body length.

o c cur r e n c e and e colo g y: Vel)' rare in marine habitats. Locus classicus is the
North Sea near Sylt, Germany, where KAm.. (1932) discovered C. marina in a mesosapro-
bic culture with Oscillatoria. Records not substantiated by illustrations: brackish water
(1-18 %0 salinity, 22-27°C) of Black Sea in Bulgaria (DETCHEVA 1980, 1981, 1983d).
REINNARTH (1979) found a rather similar species (Steinia cf. marina) in the mud-water in-
terface of freshwater habitats near Bonn, Germany; vel)' likely confused with another
species.

Insufficient redescriptions

Steinia candens KAm.. 1930 - CHARDEZ, 1981, Revue verviet. Hist. nat., 38: 53 (Fig.
223d). Remarks: Neither the number and the arrangement of the cirri nor the morphology
of the oral apparatus indicate that the identification is correct. Freshwater in Belgium.
324 SYSTEMATIC SECTION

Steinia caudens KARL - CHARDEZ, 1971, Revue verviet. Hist. nat., 28: 38 (Fig. 2341). Re-
marks: Incorrect subsequent spelling. The illustration does not show the typical Cyrto-
hymena-like buccal area. Freshwater in Belgium.

Steinia inquieta (STOKES, 1887) - VUXANOVICI, 1961, Studii Cerc. BioI., 13: 439 (Fig.
234f-h). Remarks: Cytoplasm with yellowish, 0.5-1.0 11m sized granules; cortical gran-
ules dark. The identification cannot be accepted because Cyrtohymena inquieta has no
cortical granules. Lake in Bucharest, Romania.

Steinia inquieta (STOKES 1887) KARL, 1932 - DINGFELDER, 1962, Arch. Protistenk., 105:
620 (Fig. 223c). Remarks: In life about 150 11m long. DINGFELDER (1962) identified his
population according to KARL (1932), however, he did not mention ifhis population has a
cortical granulation or not. During warm season in road pools in Bavaria, Germany.

Notohymena BLATIERER & FOISSNER, 1988

1988 Notohymena BLAITERER & FOISSNER, Stapfia, 17: 70 - Type (original designation): Notohymena rubes-
cens BLAITERER & FOISSNER, 1988.
1997 Notohymena BLAITERER & FOISSNER, 1988 - BERGER & FOISSNER, Arch. Protistenk., 148: 146.

C h a r act e r i sat ion: Adoral zone of membranelles formed like a question mark.
Undulating membranes in Notohymena pattern. Frontoventral cirri -in V-shaped pattern.
Postoral ventral cirri in dense cluster behind buccal vertex. Two pretransverse ventral and
5 or 4 transverse cirri. One right and 1 left row of marginal cirri. Six dorsal kineties. Cau-
dal cirri present. Primordia V and VI of proter originate de novo. Dorsal morphogenesis
in Oxytricha pattern.

Add i t ion a I c h a r act e r s: Long elliptical, both ends usually moderately broad
rounded. 2-3:1 flattened dorsoventrally. Very flexible and slightly contractile. Two ellip-
soidal macronuclear nodules arranged slightly left of median. Contractile vacuole some-
what in front of mid-body, during diastole with two, usually short collecting canals. Corti-
cal granules present (N. rubescens, N. antarctica, N. australis); Notohymena selvatica
was described without live observation, this character is thus not mentioned in the origi-
nal description. Pellicle and cytoplasm colourless, cytoplasm usually with many I-311m
sized globules and cytoplasmic crystals. Usually rapid movement. Buccal area deep, thus
appears bright at low magnification. Frontal cirri slightly to distinctly enlarged. Arrange-
ment offrontoventral cirri as in Figure 104f. Left marginal row usually terminates at pos-
terior end at mid-line, right one at about level of posterior transverse cirri. Transverse
cirri protrude distinctly beyond posterior end of cell, arranged in a rather oblique row.
Usually 6 dorsal kineties, including 2 dorsomarginal rows.
 Notohymena 325

T a x 0 nom y: The characterisation is according to BERGER & ForSSNER (1997); for de-
tailed description of patterns mentioned, see chapter 2 in the general section. The cladisti-
cally significant character of Notohymena, namely the hook-shaped distal end of the paro-
ral, is recognisable only after protargol impregnation (Fig. 104h).
The morphogenesis of the type species, N. rubescens, shows a conspicuous feature,
namely the new fonnation of anlagen V and VI of the proter (Fig. 104t, u; Voss 1991a),
which is considered as the autapomorphy of the Urosomoida-Cyrtohymena-Notohymena
group (Fig. 25a, Table 4).

Key to species

Species identification in Notohymena unconditionally requires live observation because

almost all (see taxonomy) species have cortical granules, that is, identification based only
on protargol impregnation is insufficient.

1 Length in life about 170-190 ~m; 4 transverse cirri (Fig. 109) . N. selvatica (p 344)
Length in life 80-140 ~m; usually 5 transverse cirri ........................ 2
2 Cortical granules ruby-coloured (Fig. 104e, g) ............. N. rubescens (p 325)
Cortical granules yellow or yellow-green to orange-green .................... 3
3 6-11 caudal cirri (Fig. 107a, e, 108a, e) .................... N. australis (p 340)
3 caudal cirri (Fig. 106a, e) ............................... N. antarctica (p 334)

Notohymena rubescens BLATIERER & FOISSNER, 1988 (Fig. 20b, 22c,

104a-z, 105a-n, Tables 3, 4, 21)

1988 Notohymena rubescens BLATfERER & FOISSNER, Stapfia, 17: 71 (Fig. 104a-h; the slide of holotype-
specimens is deposited in the OberOsterreichische Landesmuseum in Linz, Upper Austria).
1991 Notohymena rubescens BLATIERER & FOISSNER, 1988 - Voss, Arch. Protistenk., 140: 219 (Fig. 104i-z,
105a-n).

T a x 0 nom y: Differs from Notohymena antarctica mainly in the colour of the cortical
granules; live observation is thus indispensable for distinction. In life, N. rubescens
should not be confused with the terricolous species Oxytricha rubripuncta (Fig. 79a-e)
and Cyrtohymena muscorum (Fig. 94a-n), which also have red granules. However, in
Oxytricha rubripuncta the undulating membranes are only slightly curved and in Cyrto-
hymena muscorum - which has the typical Cyrtohymena pattern of undulating mem-
branes - the granules occur both close underneath the pellicle and in the cytoplasm. Fur-
thennore, both species are usually larger than Notohymena rubescens. The reddish and
brownish coloured Rubrioxytricha species are confmed to freshwater.
 326 SYSTEMATIC SECTION

, )
, \ .-
, )
,
,
, , , (i
i
, \

, . r\ )

.
,
.... \l\
I
\ \
\.
... , \
\.
,
,
,
, u·
• '.
• \
\.
\. !
..
\\ ' c

,
,
cG ,
, .".

d e ~
.".
)
Fig. 104a-f Notohymena rubescens (from BLAlTERER & FOISSNER 1988. a, d, e, from life; b, c, f, protargol im-
pregnation). a) Ventral view, 100/lm. b, c) Ventral and dorsal infraciliature, b, c = 90 /lm. d, e) Lateral and
dorsal view, d = 110 /lm, e = 105/lm. f) Ventral infraciliature of anterior half of cell, bar division = 10 /lm.
Arrow marks genus-specific hook at anterior end of parora\. cG = ruby-coloured cortical granules; p 325.
 Notohymena 327

9
Fig. l04g, h Notohymena rubescens (from BLATIERER & FOISSNER 1988. g, from life; h, protargol impregna-
tion). g) Dorsal surface with the ruby-coloured cortical granules. h) Ventral infraciliature. Arrow marks hook at
anterior end ofparoral, the autapomorphy of Notohymena; p 325.

M 0 r p hoi 0 g y and b i 0 log y: The Voss (1991a) population from Germany

agrees very well with the type population from Australia. In life 90-100 x 30-35 Ilm.
Left margin convex, right straight to slightly convex. Macronuclear nodules of type popu-
lation after protargol impregnation 9-15 x 6-9 J1m (mean = 11.8 x 7.3 J1m; n = 10), ac-
cording to Voss (1991a) 8-20 x 7-11 Ilm (mean = 14.3 x 9.5 J1m; n = 25), with many
spherical nucleoli. Distance between macronuclear nodules after protargol impregnation
4--15 Ilm (mean = 10.5 Ilm; n = 25). 1--4, usually 2, after protargol impregnation 3 Ilm
sized, spherical micronuclei. Cortical granules 1.0-1.4 Ilm across, ruby-coloured; speci-
mens thus appear conspicuously reddish at low magnification and bright field illumina-
tion; granules arranged in small groups around cirri and dorsal cilia, on ventral side ar-
ranged in short rows (Fig. 104e, g, i).
Adoral zone of membrane lies about 34 % of body length, bases of largest mem-
branelles in life approximately 7 Ilm wide. Anterior portion of peristomial lip hooked
(Fig. 104a, i, arrow). Undulating membranes probably composed of2 rows of basal bod-
ies each. Number of frontal (3), buccal (I), frontoventral (4), postoral ventral (3), pre-
transverse ventral (2), transverse (5), and caudal cirri (3) invariable at sample sizes of 10
 328 SYSTEMA TIC SECTION

Fig. 104i-k Notohymena rubescens (from Voss 1991a i, bright field with oblique illumination;j, phase con-
trast; k, protargol impregnation). i) Ventral view. Arrow marks hooked anterior end of peristomiallip (that is
not the hooked end of the paroral!). The general appearance of the oral apparatus is reminiscent of Cyrtohy-
mena. j) Cyst. k) Ventral infraciliature of anterior half of a morphogenetic stage. Arrow marks reorganizing
undulating membranes of proter. Ma = anterior macronuclear nodule, sG = cortical granules; p 325.

Fig. 1041--0 Notohymena rubescens (from Voss 1991a. h>, protargol impregnation). Ventral infraciliature ~
and nuclear apparatus of very early and early morphogenetic stages, bar = 30 J.lm. The formation of the oral
primordium commences near the left transverse cirrus (I). Next, it extends anteriorly up to the postoral ventral
cirri. See text for details. OP = oral primordium; p 325.
 Notohymena 329

"
"
"
"
q
"
c "
"
()

"
""
"
"
q
~ "
()
IJ OP ()

o~ t:'
0
0
<:>
"
0
0
~8 t:'

v
~
~
"
" ~
~
~
'"
'" \~
C1 ,~ "
C1
\~ ""
C1 " I)

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0 II

"
0
i "()
r:J
~
'"
()

~
""
()

I
() 11
~
•
()
~ "
"
()
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r:J

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¢
oJ 0
) t:1

~ 0: ,:1
" 0
0
c
0 0
; 0
C1
" ¢
0
o
p
~ 00 a
n
"
<:1
0
P: 0
 330 SYSTEMATIC SECTION

~~
~
c=
/?E
\

o9° iff!!i' o
" Cr~
~ ~ ~ " " ~ 0\
"
~"
r::s;;
d . §;f;
()
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I

'-.::.....
~

~
"\
"

-'1,
¢ .:.
(J ¢'f;.~. ~

:I&. " -'~ "

()
<>
" ()

" ()
" "~
\~.

" " r.:.l.~

gj
"
tJ

~ ~
"
()
rtf
~ t37
"
tJ
:it' §
e

I
E
"
()
?i'
() ~ "
" 0
~
()
~

0
~ " \]
aa
" ~
0
" ~

~oo
" ~8 p
<7 ()

¢
c ()
0" q

Fig. l04p, q Notohymena rubescens (from Voss 1991a p, q, protargol impregnation). Ventral infraciliature of
early morphogenetic stages, bar = 30 J.lm. Arrows mark parental cirri (V/3 in p; V/4 in q) which are modified
to primordia See text for details; p 325.

(BLATIERER & FOISSNER 1988) and 25 (Voss 1991a). Frontal, ventral, and marginal cirri in
life about 14 J.1m, transverse cirri about 20 J.1m long. Bases of frontal and transverse cirri
distinctly enlarged. Dorsal cilia in life about 3 J.1m long. Dorsal kineties 1,4, and 6 anteri-
orly slightly, kineties 5 and 6 posteriorly distinctly shortened (Fig. 104c, w).
Cyst spherical, mean diameter in life 33 J.1m (SD = 2.2 J.1m, CV = 6.7 %, Min =
28 J.1m, Max = 36 J.1m, n = 25). Cytoplasm heavily granulated. Cyst wall covered with a
2 J.1m thick, mucous layer. Ectocyst smooth (Fig. 104j).
M 0 r ph 0 g e n e sis of cell division is described in great detail by Voss (l991a;
Fig. 104k-z, 105a-n). It commences with the proliferation of basal bodies close to the
left transverse cirrus to form the oral primordium (Fig. 1041-p). Three frontal-ventral-
transverse streaks of the opisthe separate from the oral primordium and 3 from the post-
oral ventral cirri (Fig. 104q-v). In the proter, anlage I originates from the parental undu-
lating membranes, anlage II from the buccal cirrus, anlagen III and IV from left (III12)
and right (IV/3) posterior frontoventral cirri, and primordia V and VI originate de novo
(Fig. 104r-v, x, Z, 105a, Table 4). The genus-specific hook at the anterior end of the pa-
roral is formed by addition of basal bodies, whereby the undulating membranes are ar-
 Notohymena 331

.
•
\.

~ ~

•
()

()

0 "
Q "
Q
"
Q
"
<)
r:
Q
0 "
() ~ 0 <7

()
Q8 <> 0
s

Fig. l04r, S Notohymena rubescens (from Voss 1991a r, s, protargol impregnation). Ventral infraciliature of
middle morphogenetic stages, bar = 30 J.lm. Arrows mark parental cirri (buccal cirrus and rvJ3 in r; III12 in s)
which are modified to primordia See text for details; p 325.

ranged side by side (Fig. I04k, x, z, lOSa, b, e, f). They cross (optically) in late stages
and their characteristic arrangement is attained just after cell fission, similarly to in Cyrto-
hymena muscorum. Dorsal morphogenesis is in the plesiomorphic Oxytricha pattern (Fig.
l04w, y, lOSe, d, g, i, 1).

o c cur r e nee and e colo g y: Probably restricted to terrestrial habitats; not very
common. Locus classicus is a tropical forest between Cairns and Innisfail, Eastern Aus-
tralia, where BLATIERER & FOISSNER (1988) discovered Notohymena rubescens in barks
(grown with lichens and mosses) from a secondary pine forest (Scots Pine, Pinus silves-
Iris); they screened 21 soil samples from Australia, but N rubescens occurred only once.
Voss (l991a) found his population in an air-dried moss and soil sample from the park of

Fig.104t-w Notohymena rubescens (from Voss 1991a t-w, protargol impregnation). t-v) Ventral infracilia- -)
ture of middle morphogenetic stages, bar = 30 J.lm. Arrows mark de novo formation of streaks V and VI in pro-
ter (t, u) and modification of anterior-most right marginal cirrus to right marginal primordium ofproter (v). w)
Dorsal infraciliature of a middle morphogenetic stage, bar = 30 J.lm. 2-6 = frontal-ventral-transverse cirri pri-
mordia II-VI; p 325. Figures on p 332.
332 SYSTEMATIC SECTION

"
" "
" " "
" " "
" " "
" "
" "
"
()

" "
" " "
" " "
" "
"o ()
" " aa "
°
o 0 o () a
08
()
<:>
()
I)
'0 0 0°
0 o
0'" u
104t

I
/ /,
'("~
~,
,

, 1~~Jl~"
...;'~j::',:,..-
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,i ~o~:.i ~....- .
"" . . l ...

.
I)
0 ()
I) 0
a "
" 'LO
v v
,
0 w
0 0
 Notohymena 333

~
~

a 0

..
~
,;:
\) 0
0° . ,
CJ
<1 ' y
X
'."

a ~ 0 (;J

o 0 0
oD0
0 0
~
CO <:> Q

0 <:> ~ c>
n
•• 0
Z
~
DQ " "" a
Fig. l04x-z, l05a NotohymelUl rubescens (from Voss 1991a. Protargol impregnation). Ventral and dorsal
ciliature of middle and late morphogenetic stages, bar 30 11m. See text for details. l-{) = primordia I-VI; p 325.
334 SYSTEMATIC SECTION

,
\

o
0
0
00
0 0
0

0 \)
0
"
"
"
r:>

b
I
Fig. IOSb, c Notohymena rubescens (from Voss 1991a b, c, protargol impregnation). Ventral and dorsal in-
fraciliature of a late morphogenetic stage, bar = 30 !lm. Parental structures white, new black; p 325.

the "Ketteler Hof" at the village of Haltem, Gennany, as well as in terrestrial mosses
from St. Jean sur Mayenne, France.
Feeds on fungal spores, heterotrophic flagellates, and naked amoebas. Voss (1991a)
cultured N. rubescens in mineral water (Vittel), adding a few drops of a dense suspension
of yeast cells and Chlorella sp. every second or third day. Biomass of lO6 individuals
about 100 mg.

Notohymena antarctica FOISSNER, 1996 (Fig. 106a-e, Table 21)

1996 Notohymena antarctica FOISSNER, Acta Protozool., 35: 109.

T a x 0 nom y: Notolzymena antarctica is very similar to N. australis as concerns size,

shape, and cortical granules. However, it has fewer adoral membranelles and marginal
cirri and, more importantly, only 3 caudal cirri. The unusually high number, namely 6-8,
of caudal cirri in N. australis is confInned in both the type material and a Gennan popula-
tion.
 Notohymena 335

........
~

,.........
~

'"\J'::..••
S
~
::
cc- -i· - - .; ::
••
••••

cc- -\-'

-
, o?,
/
d

Fig. 105d-f Notohymena rubescens (from Voss 1991a. d-f, protargol impregnation). d, e) Dorsal and ventral
infraciliature oflate morphogenetic stages, bar = 30 Ilm. Parental structures, new black. 1) Basal body pairs are
added to form genus-specific, broadened anterior end of paroral. CC = new caudal cirri, DP = primordia of
dorsomarginal kineties; p 325.

M 0 r p hoi 0 g y: In life 80-110 x 30-40 Jlm. Prolate ellipsoidal, right side straight or
slightly concave, left rather distinctly convex, both ends broadly rounded, flattened up to
3:1. Flexible like, for example, Oxytricha granulifera. Macronuclear nodules ellipsoidal,
after protargol impregnation 11-18 x 7-8 Jlm (mean = 12.9 x 7.9 Jlm; n = 11), rather
close (mean = 7 11m) together in middle third of cell to left of mid-line. Usually 1 globu-
lar (3-4 x 2.3-3 11m, after protargol impregnation) micronucleus attached to each macro-
nuclear nodule; rarely 3 micronuclei. Contractile vacuole in mid-body at left margin, with
two inconspicuous collecting canals. Cytopyge in posterior end between transverse cirri
and left marginal cirral row; faecal balls contain yellowish globules like those found in
the cytoplasm. Pellicle colourless, flexible; cortical granules arranged in groups around
cirral bases and dorsal bristles (Fig. 106c), yellow to yellow-green, give cell yellowish
colour at low magnification and bright field illumination, do not stain with protargoL Cy-
toplasm colourless, contains some 1-4 Jlm sized, yellowish fat globules and rather many
336 SYSTEMATIC SECTION

DP-:·.:
..
: .. ,.
" · f"
··.•""
..
...
) I
\\'I/~ "
#
.j
\0
• • v . "f) ~ "

...'• v.;::,
tl1==
~

DP
}Jj
II
()
==~~
;::,l
f •

· .-" ,
• i ~:
,,,~" ,.'It·t~ E

I
~ :

I
~
~
t
..1•
I" 0
: . ..--
- 0
Q

-. • c o a 0

0:;'. 9 08 ~
0
h
'. 0

•'f•
or •
.
Fig. l05g-j Notohymena

..,(J\-
c \\\""- rubescens {from Voss
1991a g-j, protargol im-
~ ~
, pregnation). Dorsal and
f s'
~ ,

.. !
ventral infraciliature of
"'/ ~

...
very late morphogenetic
~.
. stages, bar = 30 J.lm. Ar-

I
~:

I
o

."" rows in (g) mark fragmen-

tation of dorsal kineties 3.
..
, •
'"
•
0 :
: a
0 Parental structures white,
\. o
\) 0
g c
" new black. DP = primor-
dia of dorsomarginal kine-
C! •
ties; p 325.
 Notohymena 337

.,.:~~ .
I,
o.
=--
§:;:~
~ :

.,
I •.' ~:
:
:
II
II
. .
~:
§i '"'"

.·
II •

'II" •
'I"; •, :"
#
~ I "." ....
' "c •
f
J •

..,~\
. "
<> .\\\,~ .' .
'-,~-
i ~~
"" ,, ~~
~.
," ~ ~

I
~II
•• II

",,
II II

, •
'"
II
II

'",
',c •II"
'0 D
DO k .
........
"

, "
,- "
"
, •
•
•
• ,
I

t
•
•
" II
• • t •
m
•• ••
n
• •..
t t

Fig. lOSk-n Notohymena rubescens (from Voss 1991a. Protargol impregnation). Ventral and dorsal infra-
ciliature of very late morphogenetic stages and postdividers, bar = 30 flm. Note that the genus-specific hook at
the anterior end of the paroral is formed just in the postdivider. Parental structures white, new black; p 325.
 338 SYSTEMATIC SECTION

Fig. l06a-e Notohymena antarctica (from FOISSNER 1996b. a-c, from life; d, e, protargol impregnation). a)
Ventral view, 95 11m. b) Cytoplasmic crystals. c) Yellow cortical granules around dorsal cilia d, e) Ventral and
dorsal infraciliature, 85 11m. Arrow marks hook of paroral, arrowhead denotes right frontal cirrus (IlIl3); p 334.

1-3 J!m long crystals, mainly in posterior half (Fig. 106a, b). Scrabbles rather quickly
amongst soil particles.
Adoral zone of membranelles 37 % of body length. Buccal field rather large and
deep, anterior portion semicircularly curved, similarly to in Cyrtohymena. Paroral and en-
doral conspicuously curved, intersect optically in mid-portion, paroral distinctly longer
than endoral, its distal end hooked (autapomorphy of Notohymena!), both very likely
composed of basal body pairs. Arrangement of frontal-ventral-transverse cirri as in Figure
l06d. Constantly 3 frontal cirri, 1 buccal cirrus, 4 frontoventral cirri, 3 postoral and 2
 Notohymena 339

Table 21 Morphometric data of Notohymena antarctica (ant, from FOISSNER 1996b), N. australis (aul, from
FOISSNER & O'DoNOGHUE 1990; au2, from GSCHWrND 1991), N. rubescens (rul, from BLATIERER & FOISSNER
1988; ru2, from Voss 1991a), and N. selvatica (sel, from HEMBERGER 1985). All data are based on protargol
impregnated specimens. All measurements in micrometres. ? = sample size not indicated. CV = coefficient of
variation (in %), Max = maximum value, mean = arithmetic mean, Min = minimum value, n = sample size, SD
= standard deviation

Character Species mean SD CV Min Max n

Body, length ant 85.5 5.8 6.7 70 91 11
aul 81.0 8.9 ll.5 66 95 12
au2 133.4 13.8 10.3 108 160 22
rul 84.1 3.3 4.0 77 88 10
ru2 104.1 15.5 14.9 81 136 25
sel 170 190 ?
Body, width ant 30.4 2.1 7.0 28 34 II
aul 31.4 4.5 14.4 22 29 12
au2 58.1 9.3 12.7 42 72 21
rul 33.7 2.9 8.5 30 38 10
ru2 36.4 6.0 16.5 26 51 25
sel 60 70 ?
Adoral zone of membranelles, length ant 32.4 2.2 6.8 28 35 II
aul 31.1 2.7 8.8 27 35 12
au2 49.3 6.6 13.3 38 60 22
rul 29.2 2.0 7.0 26 32 10
ru2 32.2 4.4 13.7 26 44 25
seJl 49.0
Adoral membranelles, number ant 30.2 1.9 6.4 27 33 II
aul 39.0 3.6 9.3 30 44 12
au2 39.0 1.9 4.9 35 43 19
rul 26.6 1.3 4.8 24 28 10
ru2 30.4 2.1 6.9 26 34 25
sel 25 30 ?
Right marginal row, number of cirri ant 16.8 1.2 6.9 15 19 11
aul 29.5 3.9 13.3 24 38 12
au2 36.0 2.3 6.5 31 40 19
rul 18.3 1.3 7.3 16 20 10
ru2 17.9 1.2 6.7 15 20 25
sel 18 21 ?
Left marginal row, number of cirri ant 17.9 1.3 7.3 IS 24 15
aul 28.2 2.5 8.8 24 32 12
au2 36.3 2.9 8.1 31 44 20
rul 17.0 1.3 7.8 15 19 10
ru2 19.6 1.2 6.1 17 22 25
sel 17 21 ?
Dorsal kineties, number ant 6.0 0 0 6 6 II
aul 6.3 0.5 7.3 6 7 12
au2 6.3 0.6 9.0 6 8 19
rul 6.0 0 0 6 6 10
ru2 6.0 0 0 6 6 25
sel 6.0 ?

I From Figure 109.

340 SYSTEMATIC SECTION

pretransverse ventral cirri, and 5, slightly enlarged transverse cirri (n = 11). Frontal and
transverse cirri about 20 !lm, marginal cirri about 15 !lm long. Gap between posterior end
of marginal rows indistinct because left row extends to mid-line of cell and is indistinctly
separated from caudal cirri. Dorsal cilia in life about 3 !lm long. Dorsal kineties 1 and 4
slightly shortened anteriorly, kineties 2 and 3 as long as body, kinety 5 terminates behind
mid-body, kinety 6 consists of about 5 basal body pairs only and ends in front of mid-
body. Invariably (n = 11) 3 caudal cirri, about 20 !lm long.

o c cur r e n c e and e colo g y: Terrestrial. Locus classicus is a sheltered north-

facing slope on Signy Island, South Orkney Islands, Antarctica (lat 600 40'S, long
45°40'W), where FOISSNER (1996b) discovered Notohymena antarctica in a grass sward
from Deschampsia antarctica patch (PH 4.4). Feeds on ciliates, heterotrophic flagellates,
and, possibly, on bacteria.

Notohymena australis (ForSSNER & O'DONOGHUE, 1990) comb. nov. (Fig.

107a-g, 108a-e, Table 21)

1988 Notohymena australis (FOISSNER & O'OoNOGIRJE im Druck) nov. comb. - BLATIERER & FOISSNER, Stap-
tia, 17: 70 (Fig. 107t: g; see nomenclature).
1990 Oxytricha australis FOISSNER & O'OoNOGIRJE, lnvertebr. Taxon, 3: 689 (Fig. 107a-e; the slide ofholo-
type specimens is deposited in the South Australian Museum).
1991 Notohymena australis (FOISSNER & O'OoNOGIRJE im Druck) BLATIERER & FOISSNER 1988 - GscHWIND,
Diplomarbeit, p 68 (Fig. 108a-e).

Nom e n c I at u rea n d t a x 0 nom y: BLATIERER & ForssNER (1988) made the new
combination before the original description, which was in press for several years, was
available. This obviously invalid nomenclatural act is corrected here.
At superficial live observation, N. australis is easily confused with Holosticha
multistilata because the colour of the cortical granulation is vel)' similar (FOISSNER &
O'DONOGHUE 1990). In terms of the increased number of caudal cirri N. australis is remi-
niscent of the euplanktonic Pseudostrombidium planctonticum, which has, however, dif-
ferent undulating membranes and a tapered body (Fig. 147a-j). Notohymena australis is
also easily confused with Cyrtohymena citrina, which is vel)' similar in all characters ex-
cept the undulating membranes, which are in Cyrtohymena pattern.

M 0 r p hoi 0 g y: In life 80-140 x 30-45 !lm (ForssNER & O'DONOGHUE 1990), after
GSCHWIND (1991) 120-150 x 50 !lm. Body margins sometimes slightly converging poste-
riorly (Fig. 107b). Macronuclear nodules in life about 14 x 10 !lm, after protargol im-
pregnation 8-17 x 7-11 !lm (mean = 11.7 x 8.4 !lm; n = 12), located close together to far
apart in middle third of cell. 1-4, usually 3, in life about 3.0 x 2.5 !lm sized micronuclei,
in life of refractive bright appearance. Cortical granules mainly along cirral rows and dor-
 ~\"~ \.
\.. ')'f ( 6
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" # •
• '"""',
.- ..- 0 )
J.
b C "'- -,,~ d :/ e

Fig,107a-e Notohymena australis (from FOIsSNER & O'DONOGHUE 1990. a-c, from life; d, e, protargol impregnation). a) Ventral view, 110 ~m. b) Dorsal view showing
cortical granules and contractile vacuole, 100 ~m. c) Lateral view, 100 ~m. d, e) Ventral and dorsal infraciliature, 80 ~m. This specimen has two supernumerary postoral w
ventral cirri and only one pretransverse ventral cirrus! Genus-specific hook of the paroral is shown in Fig. 107t: g. CC = caudal cirri, I, 4, 6 = dorsal kineties; p 340. ~
 342 SYSTEMATIC SECTION

9
Fig. 107f, g Notohymena australis (from BLAlTERER & FOISSNER 1988). Ventral infraciliature of anterior half
of cells of type population, bar = IO J.lm. Only (g) shows genus-specific hook of paroral clearly (arrowhead);
p340

sal kineties, arranged in small groups; cells appear greenish at low magnification; gran-
ules do not impregnate with protargol.
Adoral zone of membranelles about 38 % of body length, bases of largest mem-
branelles in life about 7 !lm wide. Number of postoral and pretransverse ventral and
transverse cirri rather variable, therefore difficult to describe any individual as "typical".
Number of postoral ventral cirri: mean = 4.3, median = 4.0, SD = 1.6, CV = 36 %, Min =
3, Max = 7, n = 12; pretransverse ventral cirri: mean = 1.7, median = 2.0, SD = 0.2, CV =
35.5 %, Min = 1, Max = 3, n = 12; transverse cirri: mean = 5.2, median = 5.0, SD = 0.6,
CV = 11.8, Min = 4, Max = 6, n = 12. Marginal cirri in life about 10 !lm, frontal and
transverse cirri about 18 !lm, dorsal cilia about 2 !lm long. Dorsal kineties 1-3 almost un-
shortened (sometimes a fourth unshortened kinety occurs). Kinety 4 begins and kinety 5
terminates slightly behind mid-body; kinety 6 consists of2-5 basal body pairs only. 6-11
(mean = 7.5, median = 8, SD = 0.6, CV = 8.6 %, Min = 6, Max = 8, n = 12, FOISSNER and
O'DONOGHUE 1990; mean = 9.1, median = 9, SD = 1.0, CV = 11.5 %, Min = 8, Max = 11,
n = 19, GSCHWIND 1991) caudal cirri arranged as in Figures 107e, 108e, located dorsally

Fig.l08a-e Notohymena australis (from GSCHWIND 1991. a, c, from life; b, d, e, protargol impregnation). a, c) ~
Ventral and dorsal view (a = 140 J.lm, c = 120 J.lm) showing, among other things, cortical granules and defeca-
tion. b, d, e) Ventral and dorsal infraciliature, d, e, = 120 J.lm, bar = 30 J.lm. Arrows mark broadened (hook-
shaped) anterior end of paroral, the sole autapomorphy of Notohymena; p 340.
 Notohymena 343

,,
,;.
."
t·
:
'"
I
Ii I

,
I
I
#

.-
.-.- { (
..
.- {
(
" (
~
~ :I
~ .
( . { .(

..
~
"",, , \
,, ,
( !

" ,. • " ,
,., .'
d e
344 SYSTEMATIC SECTION

above gap between marginal rows, appearing in life as conspicuous seam (Fig. 107a,
108a).

o c cur r e n c e and e col 0 g y: Locus classicus is a small pond in metropolitan

Perth, Western Australia. Recently, W. FOISSNER (pers. comm.) found Notohymena aus-
tralis in an alphamesosaprobic area of the Amper River, Bavaria (GSCHWIND 1991). In
cultures N. australis feeds on bacteria, rice starch, and ciliates (Dexiostoma campy/um,
Glaucoma scintillans).

Notohymena selvatica (HEMBERGER, 1985) BLATIERER & FOISSNER, 1988

(Fig. 109, Table 21)

1985 Oxytricha selvatica HEMBERGER, Arch. Protistenk., 130: 406 (Fig. 109; 1 slide of type specimens is de-
posited in the Institut filr landwirtschaftliche Zoologie und Bienenkunde of the University of Bonn,
Germany).
1988 Notohymena selvatica (HEMBERGER, 1985) - BLATrERER & FOISSNER, Stapfia, 17: 70.

T a x 0 nom y: HEMBERGER (1985) mentioned some resemblance to "Oxytricha musco-

rum KAHL" (I assume that he meant Cyrtohymena muscorum; note that KAHL described 4
different 0. muscorum!) and "Oxytricha ru-
bra BUITKAMP", which I consider as a synonym
of C. muscorum. Cyrtohymena muscorum has
a cortical granulation; thus, it is unclear if this
resemblance refers only to the general appear-
ance of the oral apparatus, or if Notohymena
selvatica also has reddish cortical granules.
Reinvestigation required.
#
• M 0 r p hoI 0 g y: In life (?) 170-190 times
.. 60-70 Ilm. Slightly tapered posteriorly. 2-3
micronuclei. Cortical granules not mentioned,

o
possibly present (see taxonomy). Adoral zone
of membranelles about 27 % of body length.
Postoral ventral cirri arranged in line. Pre-
transverse ventral cirri very close to distinctly
enlarged, about 20 Ilm long transverse cirri;
rather invariably 4 transverse cirri. Marginal
cirri 14-18 Ilm, dorsal cilia 6 Ilm long. Four

Fig. 109 Notohymena selvatica (after HEMBERGER 1985).

Ventral infraciliature and nuclear apparatus after protar-
gol impregnation, 180 11m.
 Urosomoida 345

dorsal kineties ofhody length (very likely kineties 1-4) and 2 shortened kineties (dorso-
marginal rows). Caudal cirri 30 ~m long.

o c cur r e n c e: Locus classicus is a soil from a woodland in Peru; not found since.
Biomass of 106 individuals about 560 mg (ForSSNER 1987a).

Urosomoida HEMBERGER in FOISSNER, 1982

1982 Urosomoida HEMBERGER, 1981- FOISSNER, Arch. Protistenk., 126: 115 - Type (original designation; see
also nomenclature): Uroleptus agilis ENGELMANN, 1862.
1985 Urosomoida HEMBERGER, Arch. Protistenk., 130: 415 (see nomenclature).
1997 Urosomoida HEMBERGER in FOISSNER, 1982 - BERGER & FOISSNER, Arch. Protistenk., 148: 148.

C h a r act e r i sat ion: Adoral zone of membranelles formed like a question mark.
Undulating membranes in Oxytricha pattern. Frontoventral cirri in V-shaped pattern.
Postoral ventral cirri in dense cluster behind buccal vertex. One (or 2) pretransverse ven-
tral cirrus and less than 4 transverse cirri. One right and 1 left row of marginal cirri. Four
dorsal kineties. Caudal cirri present. Primordia V and VI of proter originate de novo.
Dorsal morphogenesis in Urosomoida pattern.

Add i t ion a 1 c h a r act e r s: Outline usually long elliptical or posteriorly converg-

ing, sometimes slightly S-shaped. Flattened about 2: 1. Very flexible. Two or 4 macronu-
clear nodules. Contractile vacuole about in mid-body (Urosomoida agilis, U. perthensis)
or slightly ahead of it (u. agilijormis, U. dorsiincisura, U. minima?); during diastole usu-
ally with short and inconspicuous collecting canals. Pellicle and cytoplasm colourless.
Cytoplasm with 5-10 ~m sized food vacuoles, usually many cytoplasmic crystals, and up
to 4 ~m sized, yellowish shining globules. Some species with cortical granules. Move-
ment slow or moderately rapid.
Adoral zone ofmembranelles 114-113 of body length. Bases of all cirri of about same
size. Marginal cirri in life 10 ~m (u. minima, U. agi/is) to 15 ~m (other species), trans-
verse cirri 15-20 ~m, and caudal cirri 10 ~m (u. minima) and 15-25 ~m (other species)
long. Transverse cirri protrude distinctly beyond posterior end of cell. Dorsal kinety 1
slightly to distinctly shortened anteriorly (not known for U. minima). Dorsal cilia 3 to
4 ~m, in U. minima 4-6 ~m long. Common in freshwater and terrestrial habitats.

Nom e n c I a t u rea n d t a x 0 nom y: The nomenclature of this genus is rather

complicated. ForSSNER (1982), who described two new Urosomoida species, assigned it
to HEMBERGER, referring to his unpublished thesis (HEMBERGER 1982, P 262). The nomen-
claturally valid paper describing the new taxa of this thesis, was only published three
years later (HEMBERGER 1985). Thus, Urosomoida HEMBERGER, 1985 would contain two
species which were already described in 1982, namely U. agilijormis ForSSNER, 1982 and
U. dorsiincisura ForSSNER, 1982. This is of course impossible! According to Article 50a
346 SYSTEMATIC SECTION

of the IcZN (1985), "HEMBERGER in FOISSNER, 1982" is the author of Urosomoida because
FOISSNER (1982) published the diagnosis proposed by HEMBERGER (1982) verbatim. Fur-
thennore, FOISSNER (1982) fixed Uroleptus agilis as type species, as proposed by HEM-
BERGER (1982). Urosomoides in CORLISS (1994, p 15) and SMALL & LYNN (1985, P 460,
461) and Urosomoidea spp. in FUNKE (1986, P 72) are incorrect subsequent spellings.
The characterisation is according to BERGER & FOISSNER (1997); for a description of
the patterns mentioned therein, see chapter 2 in the general section. The flexible body, the
involvement of postoral ventral cirrus V/3 in primordia fonnation, the short adoral zone
of membranelles, and the cortical granules in some species assign Urosomoida unambi-
guously to the Oxytrichinae (Fig. 25a). Together with Notohymena and Cyrtohymena, it
shares the character that primordia V and VI of the proter are fonned de novo (Table 4,
Fig. I 13k). The autapomorphies of Urosomoida are (i) the usually distinct reduction of
the number of pretransverse ventral cirri and transverse cirri (convergence to some Uro-
soma species) and (ii) lack of fragmentation of dorsal kinety 3 (convergence to Urosoma
and some Oxytricha species).
Urosoma differs from Urosomoida in the arrangement of the frontoventral cirri (lon-
gitudinal row against V-shaped pattern) and in the shape of the undulating membranes
and the adoral zone of membranelles (Gonostomum pattern against Oxytricha pattern).
Oxytricha differs from Urosomoida in that it usually has a full set (7, sometimes 6) of
pretransverse ventral and transverse cirri and more (usually S or 6) dorsal kineties. How-
ever, some transitions exist between these taxa so that an unequivocal classification is not
always possible (for example, Urosomoida antarctica, U. granulifera, Oxytricha is-
fandica, O. auripunctata, O. fonga); morphogenetic data are needed for a more accurate
classification of these species.
Concerning the ventral ciliature, most Urosomoida species differ almost exclusively
in the number of pretransverse ventral and transverse cirri (Table 22). The infraciliature
of U. minima deviates in some points (for example, only 3 dorsal kineties) from the rather
unifonn organisation, indicating that the classification of this species is uncertain.

Key to species

Species identification unconditionally requires live observation because some species

have cortical granules. The differences between Urosomoida and Oxytricha are
sometimes indistinct, thus refer also to the Oxytricha key if identification is unsuc-
cessful.

1 Usually 4, rarely up to 6 macronuclear nodules (Fig. lISa, c) .................... .

. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. U. dorsiincisura (p 362)
2 macronuclear nodules ............................................... 2
2 Invariably only I micronucleus between macronuclear nodules (Fig. 1I4a, e) ..... .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . U. perthensis (p 362)
 Urosomoida 347

Usually with at least 1 micronucleus near each of the two macronuclear nodules (for
example, Fig. 110e, g) ................................................ 3
3 Cortical granules present (Fig. 11 Oh) .................................... 6
Cortical granules absent .............................................. 4
4 3 dorsal kineties ....................................... U. minima (p 366)
4 dorsal kineties (for example, Fig. 11OeI, 112c) ........................... 5
5 1-4, usually 3 transverse cirri, including pretransverse ventral cirri (Fig. 112a, b) .. .
· ................................................ , U. agiliformis (p 356)
Usually 4 or 5 transverse cirri and 2 pretransverse ventral cirri (Fig. 44a, c, 116d) 8
6 (3) Body distinctly tapered posteriorly; usually 3 pretransverse ventral and trans-
verse cirri (Fig. 110a-j) ................................... U. agi/is (p 347)
Body long elliptical; 5 or 6 pretransverse ventral and transverse cirri (Fig. 63a, e,
Illa,e) ........................................................... 7
7 Cortical granules colourless, irregularly arranged in a few groups; 2 caudal cirri (Fig.
111 b) ............................................ U. granulifora (p 352)
Cortical granules orange-yellow; usually 3, sometimes 4-5 caudal cirri (Fig. 63) '"
· .......................................... Oxytricha auripunctata (p 194)
8 (5) Body broadly elliptical; 2 postoral ventral cirri (Fig. 116a, b, d) .............. .
· .................................................. U. antarctica (p 365)
Body long elliptical; 3 postoral ventral cirri (Fig. 44a, c) .. Oxytricha fonga (p 158)

Urosomoida agilis (ENGELMANN, 1862) HEMBERGER in FOISSNER, 1982

(Fig. 110a-q, Tables 3, 4, 22)

1862 Uro/eptus agilis ENGELMANN, Z. wiss. Zool., 11: 386 (Fig. l1Oa).
1882 Uroleptus agilis ENG. - KENT, Manual infusoria II, p 781.
1932 Uroleptus agilis ENGELMANN, 1862 - KAHL, Tierwelt Otl., 25: 550.
1972 Oxytricha agi/is (ENGELMANN, 1862) - BORROR, J. Protozool., 19: 13.
1975 Oxytricha agi/is ENGELMANN - BUITKAMP, Acta Protozool., 14: 68 (Fig. 11 OJ-q).
1982 Urosomoida agilis (ENGELMANN, 1862) - FOISSNER, Arch. Protistenk., 126: 115 (Fig. llOe-i; authorita-
tive redescription; neotype slides are deposited in the OberOsterreichische Landesmusewn in Linz, Up-
per Austria).
1985 Urosomoida agilis (ENGELMANN) - HEMBERGER, Arch. Protistenk., 130: 415.
1986 Oxytricha agilis (ENGELMANN, 1862) BORROR, 1972 - DRAGESCO & DRAGESCO-KERNEIS, Faune tropicale,
26: 465.
1987 Urosomoida agi/is (ENGELMANN, 1862) HEMBERGER, 1985 - BERGER & FOISSNER, Zool. Jb., 114: 235
(Fig. llOb, c; one slide of protargol-impregnated specimens is deposited in the OberOsterreichische
Landesmuseum in Linz, Upper Austria).

Nom e n c I at u rea n d t a x 0 nom y: For the complicated nomenclature, see rele-

vant chapter of genus section. Utrofeptus agilis in ENGELMANN (1862, P 387, footnote I)
is an incorrect spelling of Uroleptus. Urosomoida agilis is very similar to some Urosoma
species as concerns the tapered posterior end. However, the V-shaped arrangement of the
348 SYSTEMATIC SECTION

(
(

, , ,
/
/
/
(
(

,/

Fig. 110a-d Urosomoida agilis (a, from ENGELMANN 1862; b, c, from BERGER & FOISSNER 1987a; d, original
from a terrestrial population from the Schlossalm area in Salzburg. a, b, from life; c, d, protargol
impregnation). s, b) Ventral views, a = 195 jlm, b = 120 jlm. C, d) Ventral and dorsal infraciliature, c, d =
90 11m. Arrow marks pretransverse ventral and transverse cirri. 1-4 = dorsal kineties 1-4; p 347.

frontoventral cirri and the oral apparatus of the Oxytricha type allow an unambiguous
separation. Uroleptus agilis sensu EDMONDSON (1906; Fig. 234e), sensu SMITII (1914; Fig.
233i), and sensu TAl (1931) are Uroleptus species because mid-ventral cirri are present

M orp hoi 0 gy an d b i 0 logy: In life 80-160 x 30-35!lm (BERGER & ForssNER

1987a, FOISSNER 1982), after ENGELMANN (1862) 75-240 !lm (rather high variability!), af-
ter BUITKAMP (1975) 130 !lm long; the BULLINGTON (1925, without illustration) population
is 214-238 x 71 !lm, indicating misidentification. Usually spindle-shaped, anteriorly
broad, posteriorly very narrowly rounded or even distinctly pointed. Slightly contractile.
Macronuclear nodules in life 10-14 x 7 !lm. Usually 2, sometimes up to 5, in life about
4 x 2 !lm sized micronuclei. Anterior collecting canal of contractile vacuole with a vesi-
cle. Cortical granules about 0.5 !lm sized, colourless to yellowish, rarely reddish, ar-
ranged mainly around bases of cirri and basal body pairs of dorsal cilia (Fig. 110h). Pos-
terior portion of cell often densely filled with up to 5 !lm sized, yellowish, cytoplasmic
crystals, so that cells appear dark at low magnification and bright field illumination. Cy-
 Urosomoida 349

9 h
Fig. llOe-i Urosomoida agilis (from FOissNER 1982. e, h, i, from life; f, g, protargol impregnation). e, h, i)
Ventral, dorsal, and left lateral view, e = 135 ~m, h, i = 120 ~m. Figure 110h shows the cortical granules and
the contractile vacuole with collecting canals. C, g) Ventral and dorsal infraciliature, 95 ~m. sO = colourless,
rarely yellowish or reddish cortical granules; p 347.

topyge, through which vacuoles with about 3-4 ,..m sized crystals may be discharged,
near posterior end of cell (Fig. 11 Ob).
Bases of largest membranelles in life about 5-7 ,..m wide, cilia of distal mem-
branelles about 15 ,..m long. Buccal cirrus somewhat behind anterior end of undulating
membranes. Distance between marginal cirri distinctly wider posteriorly than anteriorly.
Transverse cirri rather inconspicuous. Cyst about 50,..m across, with smooth surface
(ENGELMANN 1862).
M 0 r p hog e n e sis commences with the proliferation of basal bodies between
the postoral ventral cirri and the left marginal row (Fig. 11 Ok). The primordia V and VI
of the proter are formed de novo, as in U. agiliformis. This is obviously the sole synapo-
morphy of the Urosomoida-Notohymena-Cyrtohymena-group (Fig. 25a). See legend to
Figure 11 OJ-q and Table 4 for further details.
 350 SYSTEMATIC SECTION

::.
\
.f
0° 0
0 ..

.,
~
go Q

Qo
u:
0
OP
••1-~ ,!:-.

0: 8:

Fig. 110j-o Urosomoida agilis (from BUITKAMP 1975.j-o, ventral infraciliature and nuclear apparatus ofmor-
phogenetic stages after protargol impregnation). j) Non-dividing specimen, 120 J.lm. k) Very early stage. Oral
primordium originates between postoral ventral cirri and left marginal row. I) Early stage. The buccal cirrus, 2
frontoventral cirri (lII/2 and IV/3), and the postoral ventral cirri are modified to primordia. m, n) Middle
stages. Primordia V and VI (arrowheads) of proter originate de novo. Both in the proter and in the opisthe 6
frontal-ventral-transverse cirri streaks are recognisable. New marginal rows are formed. 0) Late stage showing
formation of new dorsal kineties 4 (dorsomarginal rows, arrowheads). OP = oral primordium; p 347.
 Urosomoida 351

a
G
p r
Fig. nop, q Urosomoida agi/is (from BUITKAMP 1975. p, q, protargol impregnation). Ventral infraciliature
and nuclear apparatus of very late morphogenetic stages. Parental structures, new black; p 347.

Fig. nOr Urosomoida minima (from HEMBERGER 1985. r, protargol impregnation). Ventral infraciliature and
nuclear apparatus, 80 J.lm; p 366.

o c cur r e n c e and e colo g y: Common in terrestrial habitats and in freshwater.

Locus classicus is a saline lake near Eisleben, Germany (ENGELMANN 1862); however, sa-
linity of this lake is low (KAHL 1932). Terrestrial records, largely substantiated by illustra-
tions: Austria (FOISSNER & ForSSNER 1988, ForSSNER et al. 1990); pasture and mixed wood-
land near Bonn, Germany (BUITKAMP 1975, 1977a, 1979); Pisang-Peak, Himalayas
(FOISSNER 1986a); savannah of Lamto, Ivory Coast (BUITKAMP 1977b); rain forest in Bra-
zil (FOISSNER 1997b).
Records from freshwater habitats not substantiated by illustrations: Belgium (CHAR-
DEZ 1987); drinking-water of Briinn, Czechoslovakia (SRAMEK-HuSEK 1953); saline lakes
near Halle, Germany (ZACHARIAS 1888); beta- to alphamesosaprobic running waters in
Germany and Italy (ForSSNER et al. 1992b, MAnoN! 1980, MAnoN! & GHETfI 1980,
198Ia); bog in Switzerland (MERMOD 1914); Iowa, USA (SHAWHAN et al. 1947); ooze ofa
lake of the Devil's Lake complex in North Dakota, USA (EDMONDSON 1920, HAMMER
1986); Tahiti (EDMONDSON 1910).
Records from terrestrial habitats not substantiated by illustrations: with a constancy
of 15-22 % in various agricultural soils in Czechoslovakia (TIRJAKovA 1988); soil infu-
sions and ridges near Erlangen, Germany (DINGFELDER 1962); Italy (LUZZATfI 1938); alka-
352 SYSTEMA TIC SECTION

line soils (salt content 12--45 g 1-1) from arid regions in southern Australia (POMP & WIL-
BERT 1988); Antarctica (SUDZUKI 1979>-
Feeds on bacteria, diatoms, amoebas, testaceans (Trinema Iineare); inorganic soil
particles were also ingested. BUITKAMP (1975) used diluted soil extract as culture
medium, adding a grain of polished rice to support bacterial growth. According to Forss-
NER (1985a) and ForSSNER et al. (1985), Urosomoida agilis is an indicator of mull-like
moder. Biomass of 106 specimens about 30 mg (FOISSNER 1987a). MATIS (1975) found it
in road pools near Bratislava, Czechoslovakia, at 16.0-28.5 DC, pH 5.8-6.4, and
7.6-10.8 mg 1-1 O2 •

Urosomoida granulifera FOISSNER, 1996 (Fig. 111 a-f, Table 22)

1996 Urosomoida granulifera FOISSNER, Acta Protozool., 35: 1I5 (type slides are deposited in the OberOster-
reichische Landesmuseum in Linz, Upper Austria).

T a x 0 nom y: ForSSNER (1996b) did not explain why he classified this species in Uroso-
moida and not in Oxytricha; very likely because the ventral and dorsal infraciliature is
more or less identical with that of "Urosomoida agiliformis" sensu GANNER et al. (1987a,
b). However, the GANNER population is obviously very closely related to the type species
of Oxytricha, O. granulifera, as indicated by morphogenetic details (primordium V of the
proter originates from postoral ventral cirrus V/4, and primordium VI from posterior pos-
toral ventral cirrus V/3). Furthermore, in Urosomoida granulifera the number of pre-
transverse ventral and transverse cirri is not distinctly decreased as in ''typical'' Uroso-
moida species (u. agilis, U. agiliformis), and 4 dorsal kineties are also known from some
"Oxytricha" species. Consequently, morphogenetic data are needed to clarify the system-
atic position of U. granulifera.

M 0 r ph 0 log y and b i 0 log y: In life about 70-100 x 20-30 ~m. Shape highly
variable, slenderly to broadly elliptical or parallel-sided, sometimes slightly fusiform,
both ends narrowly to broadly rounded. Dorsoventrally flattened up to 2: 1. Usually 2,
very rarely 3 distinctly ellipsoidal macronuclear nodules in middle third of cell to left of
median, after protargol impregnation 12-20 x 5-7 ~m (mean = 15.6 x 5.7 ~m; n = 12).
Micronuclei ellipsoidal, after protargol impregnation 3.0 x 2.4 ~m on average, number
highly variable (2-8; mean = 3.1; n = 12), however, most specimens have two. Contrac-
tile vacuole in mid-body, with 2 inconspicuous collecting canals extending anteriorly and
posteriorly. Cortex colourless. Cortical granules inconspicuous because colourless, min-
ute (about 1 ~m across) and sparse, form small, irregularly arranged patches composed of
up to 30 granules. Cytoplasm colourless, contains many small, yellowish crystals. Move-
ment moderately rapid, scrabbling amongst soil particles.
Adoral zone of membranelles about 34 % of body length, bases of largest mem-
branelles in life about 7 ~m wide. Buccal cavity narrow and flat, right half and posterior
 \
: \
/
\'
,#
/
I ,,, ,/
.... _,
~
"\. ··\'f ,., ,/
I ,,
., ~ ~
, 1;;
c
1ft "-0\"
., ~:"".':
..
'J.~.'.; ,/ ~
., -/- c
, \ ~
., -----
/I
., \., 7
,
,
., I
Os d
~. l' r
\;'\

e .}f
Fig. 111a-f Urosomoida granulifera (from FOISSNER 1996b. a-d, from life; e, f, protargol impregnation). a, c, d) Ventral and lateral view, 80 Jim. Short canals are
formed during diastole of contractile vacuole (c). b) Dorsal view showing spherical, about 1 Jim sized, colourless cortical granules arranged in irregular groups. e,1)
Ventral and dorsal infraciliature, 100 Jim. Arrowheads mark right transverse cirrus and anterior end of dorsal kinety 1; P 352.
w
VI
W
354 SYSTEMATIC SECTION

Table 22 Morphometric data of Urosomoida agiliformis (afl, from FOISSNER 1982; at2, from FOISSNER &
ADAM 1983a), U. agilis (agl, from BUITKAMP 1975; ag2, from FOISSNER 1982; agJ, from BERGER & FOISSNER
1987a), U. antarctica (ant, from FOISSNER 1996b), U. dorsiincisura (dor, from FOISSNER 1982), U. granulifera
(gra, from FOISSNER 1996b), U. minima (min, from HEMBERGER 1985), and U. perthensis (per, from FOISSNER &
Q'DoNOGHUE 1990). All data after protargol impregnation. All measurements in micrometres. ? = sample size
not indicated; if only I value is known it is listed in the mean column, if 2 values are available they are listed
as Min and Max. CV = coefficient of variation (in %), Max = maximum value, mean = arithmetic mean, Min
= minimum value, n = sample size, SD = standard deviation
Character Species mean SD CV Min Max n
Body, length afl 70.4 4.0 5.6 64 77 12
a12 69.5 7.2 10.4 55 87 25
agl 130.0 1
ag2 94.0 15.3 16.3 79 124 8
ag3 86.1 5.3 6.2 78 96 15
ant 61.6 5.4 8.9 52 70 13
dor 81.3 8.5 10.4 70 98 10
gra 73.9 9.2 12.4 59 90 12
min 75 90 ?
per 42.6 4.4 10.2 38 51 10
Body, width afl 20.6 2.2 10.8 17 24 12
at2 21.7 3.2 14.9 16 28 25
agl 42.0
ag2 17.9 2.1 11.7 14 21 8
agJ 25.3 4.2 16.7 17 32 15
ant 27.3 3.3 12.1 22 33 13
dor 24.0 2.0 8.3 20 27 10
gra 21.5 3.1 14.6 17 29 12
min 15.0 ?
per 18.8 2.1 11.2 15 22 10
Adoral membranelles, number afl 21.7 0.8 3.9 20 23 12
a12 22.2 1.3 6.0 20 26 25
agl 25.0 ?
ag2 27.9 1.5 5.5 25 30 8
ag3 25.7 1.9 7.6 23 30 15
ant 19.5 0.8 4.0 18 21 13
dor 31.6 2.2 7.0 28 36 10
gra 23.7 1.9 7.9 21 27 12
min 14 15 ?
per 16.6 1.0 5.8 15 18 10
Right marginal row, number of cirri afl 20.1 1.0 4.7 19 22 12
a12 24.1 2.3 9.7 20 29 25
agl 20 22 ?
ag2 22.4 2.0 8.9 19 25 8
agJ 22.7 2.5 10.8 17 27 15
ant 17.0 1.2 7.2 14 19 13
dor 34.1 1.9 5.6 32 39 10
gra 24.1 3.7 15.2 16 30 12
min II 13 ?
per 19.5 1.9 9.7 17 22 10
 Urosomoida 355

Table 22 Continued

Character Species mean SD CV Min Max n

Left marginal row, number of cirri afl 19.6 1.6 8.2 16 22 12
af2 21.6 2.0 9.2 18 25 25
agl 24 26 ?
ag2 22.4 2.2 10.0 19 26 8
ag3 23.7 3.5 14.9 20 31 15
ant 19.9 2.1 10.3 17 24 13
dor 29.7 2.4 8.1 26 34 10
gra 23.3 3.2 13.7 17 29 12
min 12 16 ?
per 16.4 2.1 12.6 14 21 10
Postoral ventral cirri, number afl 3.2 0.6 18.6 3 5 12
af2 3.0 0.4 15.0 2 5 25
agl 3.0 ?
ag2 2.6 0.7 26.5 3 8
ag3 3.1 0.3 8.8 3 4 15
dor 3.0 0 0 3 3 10
gra 2.9 2 3 12
min 3.0 ?
per 3.1 0.3 10.2 3 4 10
Transverse cirri, number afl 3.1 0.3 9.7 3 4 12
(including pretransverse af2 2.9 0.6 20.8 4 25
ventral cirri) agl 3.0 ?
ag2 3.0 0 0 3 3 8
ag3 3.0 0 0 3 3 15
anti 5.1 5 6 13
dor 2.8 0.4 14.3 2 3 10
gral 3.9 3 4 12
min 5.0 ?
per 5.0 0 0 5 5 10
Caudal cirri, number afl 2.2 0.4 17.2 2 3 12
af2 2.1 0.5 24.8 1 4 25
agl 3.0 ?
ag2 3.0 0 0 3 3 8
ag3 3.0 0 0 3 3 IS
ant 2.0 0 0 2 2 13
dor 2.2 0.4 17.2 2 3 10
gra 2.0 0 0 2 2 12
min 3.0 ?
per 3.0 0 0 3 3 10

I Pretransverse cirri (invariably 2) not included!

third of adoral zone covered by hyaline lip. Paroral and endoral almost straight, extend
side by side diverging posteriorly (that is, without distinct Oxytricha pattern). Pharyngeal
fibres inconspicuous. Frontal, transverse, and caudal cirri in life about 18 J.lID long, other
cirri about 12 J.lID long. Marginal rows open at posterior end, gap occupied by posterior-
 356 SYSTEMATIC SECTION

"-

r '"

1\
"
\

a
,
t.

b
(j AAc d e
Fig. 112a-e Urosomoida agiliformis (from FOISSNER 1982. a, d, e, from life; b, c, protargol impregnation). a,
d, e) Ventral, dorsal, and lateral view, a = 110 Ilm, d, e = 85 Ilm. b, c) Ventral and dorsal infraciliature, 75 Ilm.
Arrow in (b) marks frontoventral cirrus III/2, arrowheads in (c) denote caudal cirri.

most transverse cirrus and caudal cirri. Ventral and dorsal infraciliature (Fig. III e, t) as
in Oxytricha fonga. Dorsal cilia in life about 4 ~m long.

o c cur r e n c e and e colo g y: Locus classicus is the Livingstone Island, South

Shetland Islands, Antarctica, where FOISSNER (l996b) discovered Urosomoida granulif-
era in Drepanocladus uncinatus moss. Possibly feeds on bacteria.

Urosomoida agiliformis FOIssNER, 1982 (Fig. 112a-e, 113a-v, Tables 3,

4,22)

1982 Urosomoida agiliformis FOISSNER, Arch. Protistenk., 126: 117 (Fig. 112a-e; I slide of holotype and 1
slide of paratype specimens are deposited in the OberOsterreichische Landesmuseum in Linz, Austria).
1983 Urosomoida agiliformis FOISSNER, 1982 - FOISSNER & ADAM, Zool. Anz., 211: 161 (Fig. 113a-v).
1987 Urosomoida agiliformis FOISSNER, 1982 - GANNER, FOISSNER & ADAM, Ann. Sci. nat., Zool., 8: 200, in
part: populations PI (= population of FOISSNER 1982) and P2 (= population of FOISSNER & ADAM
1983a); not populations P3 and P4 (see taxonomy).
1996 Urosomoida agiliformis FOISSNER, 1982 - WANG & SONG, 1. Ocean Univ. Qingda, 26: 179, Fig. lA-J
(reinvestigation of morphogenesis).
 Urosomoida 357

" l () -op
" '~--,,-

" " "

"
" " "
" " "
" " "
" "
"
" "

•
. ()o :,/- ----oP
d
a c

Fig. 113a-f Urosomoida agili-

{ormis (from FOISSNER & ADAM
1983a a-f, protargol impregna-
<>
" tion). a, b) Ventral and dorsal in-
" fraciliature of a non-dividing
specimen, 65 Ilm. c-t) Ventral
and dorsal infraciliature of very
; early morphogenetic stages, c, d =
o
:-o--op 80 Ilm, e, f = 85 Ilm. The macro-
nuclear nodules are conspicuously
enlarged and of irregular shape (t).
OP = oral primordium, 1-4 = dor-
e sal kineties 1-4; p 356.
358 SYSTEMATIC SECTION

"
"
"

"
"
"
"
"
"
{ "
I "
"
"
"
I " "
I
"
"
: •.t

113g

ARM R-J
I
"

"
"
"
"
"
"
ARM R--\
I
\
\,
"
"
"
"

•• ..'
k m
 Urosomoida 359

Nom e n cia t u rea n d t a x 0 nom y: Urosomoides agiliformis in SMALL & LYNN

(1985, P 461) is an incorrect subsequent spelling of Urosomoida. Two features strongly
suggest that the GANNER et al. (1987a, b) populations P3 and P4 are not conspecific with
U. agiliformis; firstly, the number of pretransverse ventral and transverse cirri is signifi-
cantly higher (means = 6.0 and 5.7 respectively) in P3 and P4 than in the populations PI
and P2 described by FOISSNER (1982) and FOISSNER & ADAM (1983a) (means = 3.0 and
2.9 respectively); secondly, the primordia V and VI of the proter do not originate de
novo, but they are derived from the postoral ventral cirri V/4 and V/3 respectively, identi-
cally to in the type species of Oxytricha, O. granulifera (Table 4). The latter character is,
at the present stage of knowledge, the only autapomorphy of the Oxytricha-Allotricha
group (Fig. 25a). Thus, P3 and P4 of GANNER et al. (1987b) are classified in Oxytricha
(see O. longa).

M 0 r p hoi 0 g y: In life about 80-100 x 20-30 /lm. Margins almost in parallel, both
ends rounded. Macronuclear nodules after protargol impregnation 9-18 x 4-7 /lm (mean
= 15 x 5 /lm; n = 12), usually ellipsoidal, rarely broadly spindle-shaped, conspicuously
voluminous and often connected by a thin thread (Fig. II2c). Cytoplasm often with
many, 3---6 /lm sized, yellowish crystals surrounded by tiny granules. Buccal cirrus at an-
terior end of undulating membranes. Dorsal kineties 1 and 2 distinctly shortened anteri-
orly, kinety 4 terminates slightly ahead of mid-body. Distance between basal body pairs
of kineties posteriorly distinctly wider than anteriorly.
M 0 r p hog e n e sis of cell division commences with the proliferation of basal
bodies at three sites, namely twice left of the postoral ventral cirri and once left of the
transverse cirri (Fig. 113c). For details, see Figures 113a-v and Table 4. According to
FOISSNER & ADAM (1983a), 1, 2, 2, 3, 3, and 3 cirri originate from the frontal-ventral-
transverse cirri streaks I-VI; however, BERGER et al. (1985a) assumed that streak VI pro-
duces 4 cirri, resulting in the following pattern: 1,2,2,2,3,4.

o c cur r e n c e and e colo g y: Common in terrestrial habitats and in the ooze and
aufwuchs of freshwater. Locus classicus is the soil of a beech forest near the village of
Baumgarten, Lower Austria (for details see FOISSNER et al. 1985, Profil 7). FOISSNER &

~ Fig. 113g-m Urosomoida agiliformis (from FOISSNER & ADAM 1983a. g-m, protargol impregnation). g-j)
Ventral and dorsal infraciliature of early morphogenetic stages, g-j = 84 )lm. k, I) Ventral and dorsal infracilia-
ture of a middle morphogenetic stage, 80 )lm. This specimen has 5 dorsal kineties. Arrow marks new (de novo)
anlage of primordia V and VI ofproter. m) Ventral infraciliature ofa middle stage, 80 )lm. AD = primordia of
dorsal kineties, AO = primordium of cirri and undulating membranes of opisthe, ARMR = primordia of right
marginal rows, PVC = postoral ventral cirrus, 1-{) = frontal-ventral-transverse cirri primordial-VI; p 356.

Fig. 113n-r Urosomoida agiliformis (from FOlssNER & ADAM 1983a n-r, protargol impregnation). n) Dorsal ~
infraciliature of a middle morphogenetic stage (same as Fig. 113m), 80 )lm. o-r) Ventral and dorsal infracilia-
ture of late morphogenetic stages, 0, p = 70 )lm, q, r = 75 11m. Parental structures white, new black. AD4 = pri-
mordium of dorsal kinety 4 (dorsomarginal row), CC = caudal cirri; p 356. Figures on p 360.
360 SYSTEMATIC SECTION

4( '.
'-.

-,,,
.cClD<;>

'" '" ~
AD-·
::"
.,- I'/~i
~ ""

! ~
.~
~
i'i'
~

"
...........
" , ....
"
, " ;:
'~
· 'iSo s

\': ('1
/1f: :s::
....-"'.- 0

" .'i §-
'.
:

~
\

.
0

0° 0
~

113n 0 p

-
,
.....
-
,,
.
. --
,,
,
, _" cc--:
o

.
.....•
..
.
.•
••
• ##
#
#

...
•,,
#

cc--·
# ,
.
',' ..
q • • r
 Urosomoida 361

• •

..-
II
II
II
- ••

II
II
II
II
II

..... .
II
II

....
, ~
..
'. ~

..
.
-
...
f.
•

s o

.
II
II
II
II
",
l§§ . ~

§.

i::: :
II
s:: ..
II
r l§:
..
II

.....
II
,~ Fig. 113s-v Urosomoida agili-
II
{ormis (from FOISSNER & ADAM
II
II
II
. 1983a S-V, protargol impregna-
tion). s, t) Ventral and dorsal infra-
II
II ciliature of a very late morphoge-
II
netic stage, 75 )!m. Above the ar-
...
II
II rowhead in (t) the micronucleus

. .. .0
did not impregnate. u, v) Ventral
and dorsal infraciliature of an opis-
#
the, 40 )!m. Parental structures
• 0 v white, new black; p 356.
362 SYSTEMATIC SECTION

ADAM (1983a) found U. agiliformis in the soil of an alpine pasture near the village of
Heiligenblut, Austria. Further records: soils from alpine pastures in the Gasteiner and
Fuscher valleys in Salzburg, and in a meadow in Aigen in Styria, Austria (FOISSNER
1985a, 1987b, FOISSNER & PEER 1985, FOISSNER et al. 1990); beta- to alphamesosaprobic
brooks and rivers in Austria and Germany (BLATIERER 1994, FOISSNER & MOOG 1992,
FOISSNER et al. 1992a, b); limed and fertilized spruce forest soils near UIm, Germany
(LEHLE 1989, 1994); leaflitter of beech forests in Germany (BONKOWSKI 1996); forest hu-
mus in France (PALKA 1991); soil from the Pisang-Peak, Himalayas (FOISSNER 1986a);
sand from the Coral Pink Sand Dunes near the Zion National Park, Utah, USA (FOISSNER
1994b); soil from Qingdao, China (WANG & SONG 1996); soils from South America, Aus-
tralia, and Tasmania (FOISSNER 1997b). Feeds on bacteria and ciliates. Biomass of 106 in-
dividuals about 30 mg (FOISSNER 1987a).

Urosomoida perthensis FOISSNER & O'DONOGHUE, 1990 (Fig. 114a-e, Ta-

ble 22)

1990 Urosomoida perthensis FOISSNER & O'DoNOGHUE, Invertebr. Taxon, 3: 692 (1 slide of holotype speci-
mens is deposited in the South Australian Museum).

M 0 r p hoi 0 g y: In life 50-70 x 20-30 !lm. Long elliptical, not contractile. Macronu-
clear nodules spherical to slightly ellipsoid, after protargol impregnation 7-8 x 5-6!lm
(mean = 7.3 x 5.7 !lm; n = 10). Micronucleus after protargol impregnation approximately
2.7 x 2.4 /lm (n = 10). Contractile vacuole lacks collecting canals. Cortical granules ab-
sent, endoplasm colourless, but heavily granulated.
Bases of largest membranelles in life about 4 !lm wide. Buccal cavity narrow but
deep, right anterior margin raggedly curved to adoral zone of membranelies. Undulating
membranes lying almost over each other. Cirri about 15 !lm long and thick in relation to
body size. Bases of pretransverse ventral cirri distinctly smaller than those of all other
cirri. Marginal rows distinctly separated posteriorly. Dorsal kinety 1 distinctly, kinety 2
slightly shortened anteriorly; kinety 1 in middle part with a conspicuous gap due to lack
of about 2 basal body pairs; kinety 4 terminates about at frrst third of cell. Prominent cau-
dal cirri at posterior end of kineties 1-3.

o c cur r e n c e and e col 0 g y: Locus classicus is a small pond in the town of

Perth, West Australia. W. FOISSNER (pers. comm.) found Urosomoida perthensis also in
soil from South Africa. In cultures, it feeds on rice starch and bacteria.

Urosomoida dorsiincisura FOISSNER, 1982 (Fig. 115a-e, Table 22)

1982 Urosomoida dorsiincisura FOISSNER, Arch. Protistenk., 126: 119 (1 slide of holotype and 1 slide of
paratype specimens are deposited in the OberOsterreichische Landesmuseum in Linz, Upper Austria).
 )

,
\
,, ,
, (
~
,, 2
c
tI ~
{ c
tI , §:
')
tI
tI
~
c ~

# "
# "
# •
ft
d

Fig. 114a-e Urosomoida perthensis (from FOISSNER & O'DoNOGHUE 1990. a-c, from life; d, e, protargol impregnation). a-c) Ventral, dorsal, and lateral view, a =
55 11m, b, c = 50 11m. Note the single micronucleus (black dot) between the two macronuclear nodules. d, e) Ventral and dorsal infraciliature, 40 11m; p 362. W
0\
W
364 SYSTEMATIC SECTION

OF

Fig. 115a-e Urosomoida dorsiincisura (from FOISSNER 1982. a, d, e, from life; b, c, protargol impregnation).
a, e) Ventral and left lateral view, a = 145 11m, e = 140 11m. b, c) Ventral and dorsal infraciliature, 80 11m. Ar-
row in (b) marks frontoventral cirrus llIl2, arrowhead in (c) denotes posterior end of dorsomarginal kinety. d)
Dorsal view showing contractile vacuole with inconspicuous collecting canals and 0.5 11m sized, yellowish, re-
ticularly arranged cortical granules, 140 11m. DF = dorsal furrow; p 362.

Tax 0 nom y: Differs from the equally sized and also quadrinucleate Oxytricha is-
landica (Fig. 68a-f) by the lower number of transverse cirri, by the somewhat higher
number of adoral membranelles, the presence of cortical granules (Fig. II5d), and the
more posteriorly located frontoventral cirrus IIII2 (Fig. ll5b, arrow). Oxytricha auri-
punctata has a similar cortical granulation, but only 2 macronuclear nodules.

M 0 r p hoi 0 g y: In life about 100-150 x 25-40 Jlm. Margins nearly parallel, both ends
broadly rounded. Dorsal surface with a distinct furrow in left anterior part of cell (Fig.
115d). Macronuclear nodules ellipsoidal to bean-shaped, after protargol impregnation
8-11 x 4-7 Jlm, rather invariably arranged in two distinctly separated groups, each group
usually with 1 micronucleus. Pellicle conspicuously crenellated at bases of cirri. Cortical
granules about 0.5 Jlm sized, yellowish, usually reticular arranged (Fig. II5d).
Bases of largest membranelles in life about 7 Jlm wide. Buccal cavity moderately
deep. Caudal cirri vigorously motile, arranged dorsally slightly right of median. Dorsal
kinety 4 terminates about in mid-body (Fig. II5c, arrowhead).
 Urosomoida 365

o c cur r e n c e and e colo g y: Probably confmed to terrestrial habitats. Locus

classicus is an intensively cultivated field near Zwentendorf, Lower Austria (Profit 3 in
FOISSNER et al. 1985). FOISSNER & PEER (1985) found U. dorsiincisura in the marginal
area of a graded ski-trail in the Gasteiner Valley, Salzburg, Austria. TIRJAKovA (1988) re-
corded it from agricultural soils in Czechoslovakia and W. FOISSNER (pers. comm.) in soil
from South Africa.
Feeds on bacteria, amoebas, and ciliates. Biomass of 106 specimens about 38 mg
(FOISSNER 1987a).

Urosomoida antarctica FOISSNER, 1996 (Fig. 116a-e, Table 22)

1996 Urosomoida antarctica FOISSNER, Acta Protozool., 35: 115 (type slides are deposited in the OberOster-
reichische Landesmuseum in Linz, Upper Austria).

T a x 0 nom y: As stated by FOISSNER (1996b), the classification of this species is uncer-

tain and requires morphogenetic data. The almost full set of 17 frontal-ventral-transverse
cirri (1 postoral is lacking!) is reminiscent of Oxytricha, whereas the simple dorsal infra-
ciliature (obviously the Urosomoida pattern) indicates that it might belong to Uroso-
moida.

M 0 r ph 0 log y and b i 0 log y: In life about 60-75 x 25-35 ~m. Broadly ellipti-
cal, posteriad usually slightly broadened, both ends broadly rounded. Dorsoventrally flat-
tened up to 2: 1. Macronuclear nodules distinctly ellipsoidal, in middle third of cell left of
median, after protargol impregnation 11-17 x 6-10 ~m (mean = 13.5 x 7.6 ~m; n = 13).
Micronuclei slightly ellipsoidal, after protargol impregnation 2.8 x 2.4 ~m on average,
number rather variable (1-3), if only one present, usually attached to anterior macronu-
clear nodule. Contractile vacuole without distinct collecting canals. Cortical granules
lacking. Cytoplasm contains many 2-3 ~m sized, colourless fat globules, some small
vacuoles with yellowish, crystalline content, and many 4-6 ~m sized food vacuoles.
Moves slowly.
Adoral zone of membrane lIes about 32 % of body length, bases of largest mem-
branelles in life about 6 ~m wide. Buccal cavity narrow and flat. Paroral and endoral
membrane almost straight, slightly intersecting optically. Pharyngeal fibres inconspicu-
ous. All cirri strikingly thin; frontal, transverse, and caudal cirri in life about 18 ~m, all
other cirri 12 ~m long. Ventral and dorsal infraciliature, see Figure 116d, e. Invariably
only 2 postoral ventral cirri. Right marginal row terminates at about level of pretransverse
ventral cirri, left one i-shaped, leaving a broad gap which is occupied by caudal cirri.

o c cur r e n c e and e colo g y: Locus classicus is Garwood Valley, South Victoria

Land, Antarctica, where FOISSNER (l996b) discovered U. antarctica in three small sam-
ples which were bulked: "salt" soil from near Garwood glacier; dry soil from "flood
 366 SYSTEMATIC SECTION

o ~r
#
\..,
\
\, \, \,
;

'-. \,
b
;

, \ \,
" \
""
c
...., .•
••
d
"
'-"'
• e
Fig. 116a-e Urosomoida antarctica (from FOISSNER 1996b. a-c, from life; d, e, protargol impregnation). a-c)
Ventral views and right lateral view, 70 11m. d, e) Ventral and dorsal infraciliature, 63 11m. Note that U. ant-
arctica has only two postoral ventral cirri and two caudal cirri; p 365.

plain" of melt-stream; rock flour from glacial outflow stream. No other ciliates recorded
in this sample! Feeds on bacteria and, possibly, also on heterotrophic flagellates and
small naked amoebae.

Urosomoida minima HEMBERGER, 1985 (Fig. 11 Or, Tables 3, 22)

1985 Urosomoida minima HEMBERGER, Arch. Protistenk., 130: 416 (type slides are deposited in the Institut
rur landwirtschaftliche Zoologie und Bienenkunde der Universitllt Bonn, Germany).

T a x 0 nom y: The classification in Urosomoida is uncertain because there are supernu-

merary frontoventral cirri, less than 4 dorsal kineties, and the oral apparatus is not exactly
in the Oxytricha pattern. Redescription necessary.

M 0 r p hoI 0 g y: After protargol impregnation 75-90 /lm long. Long elliptical, poste-
rior end slightly tapered. Three distal membranelles separated from proximal portion of
adoral zone of membranelles by a distinct gap. Buccal area very small. Undulating mem-
branes nearly straight and clearly not intersecting. Both right and left marginal row begin
 Gonostomum 367

at about level of cytostome and tenninate distinctly ahead of posterior cell end. Oral pri-
mordium originates left of postoral ventral cirri.

o c cur r e n c e and e colo g y: Locus classicus is an infusion of soil from a gar-

den near Bonn, Germany. HEMBERGER (1982, 1985) found U. minima also in the soil of a
woodland from near Bonn. Not found since. Biomass of 106 individuals about 12 mg
(FOISSNER 1987a).

Insufficient redescriptions

Ciliate sp. - MOTE, 1954, Proc. Iowa Acad. Sci., 61: 588, Plate VIII, Fig. 30a (Fig.
225u). Remarks: MOTE (1954) found this hypotrich in a prairie soil in Iowa, USA. In
terms of the body shape and the (insufficiently described) ciliature it could be a Uroso-
moida.

Uroleptus agi/is EHR. - SMITH, 1914, Kans. Univ. Sci. Bull., 9: 164 (Fig. 233i). Remarks:
"Urpleptus agilis ENG." in the legend to the figure is an incorrect subsequent spelling.
Obviously a Uroleptus because a mid-ventral row is present. In life 83-250 J.lm long.
Limnetic habitat in Kansas, USA.

Uroleptus agilis EHRBG. - TAl, 1931, Sci. Rep. natn. Tsing Hua Univ., 1: 50, PI. XIV,
Fig. 6. Remarks: A Uroleptus because a mid-ventral row is present. In life about 160 J.lm
long. Stream on east campus ofTsing Hua University, China.

Uroleptus agilis (?) ENG. - EDMONDSON, 1906, Proc. Davenport Acad. Sci., 11: 98 (Fig.
234e). Remarks: A Uroleptus because a mid-ventral row is present. Freshwater in Iowa,
USA.

Gonostomum STERKl, 1878

1878 Gonostomum STERK!, Z. wiss. Zoot., 31: 57 - Type (original designation): Oxytricha affinis STEIN,
1859.
1882 Plagiotricha KENT, Manual infusoria II, p 771 - Type (original designation): Oxytricha strenua ENGEL-
MANN,1862.
1932 Gonostomum STERK!, 1878 - KAHL, TierweltDtt., 25: 597.
1982 Gonostomum STERK!, 1878 - FOiSSNER, Arch. Protistenk., 126: 74.
1984 Gonoslomum STERK!, 1878 - MAEDA & CAREY, Bull. Br. Mus. nat. Hist., 47: 8.
1997 Gonostomum STERK!, 1878 - BERGER & FOiSSNER, Arch. Protistenk., 148: 145.

C h a r act e r i sat ion: Adoral zone of membranelles and undulating membranes in

Gonostomum pattern. Frontoventral cirri roughly in V-shaped pattern. Postoral ventral
cirri right of adoral zone. Usually fewer than 4 transverse cirri. One right and 1 left row
368 SYSTEMATIC SECTION

of marginal cirri. Three dorsal kineties. Caudal cirri present. Primary primordia. Dorsal
morphogenesis in Gonostomum pattern.

Add i t ion a I c h a r act e r s: Body flattened about 2:1, flexible. Contractile vacuole
about in mid-body, slightly to distinctly displaced inwards. Cortical granules present,
however, difficult to discern; cytoplasm colourless. Adoral zone of membranelles 40 to
50 % of body length, distally not overlapping on right margin. Buccal area flat and nar-
row. Left frontal cirrus displaced posteriorly, sometimes slightly larger than middle and
right one. Buccal cirrus at level of rather short paroral. Three, Gonostomum affine rarely
with 4 dorsal kineties, each with one mobile caudal cirrus; kinety 1 slightly shortened an-
teriorly; dorsal cilia in life about 3 Jlm long, except in G. gonostomoida, where they are
5 Jlm, posteriorly even up to 9 Jlm long. Feed mainly on bacteria. Common in terrestrial
habitats, rarely in freshwater.

Nom e n c I at u rea n d t a x 0 nom y: STERK! (1878) established Gonostomum un-

ambiguously for Oxytricha affinis, whereas KENT (1882) designated O. strenua clearly as
type species of Plagiotricha. Obviously erroneously, JANKOWSKI (1979) was of contrary
opinion. Plagiotricha KENT, 1882 is a homonym of Plagiotricha BORY DE SAINT-VINCENT
in LAMOUROUX et al. (1824). I do not know the original description of "Gonostomum mer-
eschkowskit' mentioned in LEPSI (1929b, p 297); very likely it is a nomen nudum, that is,
a new species without description.
Plagiotricha KENT was established for Oxytricha strenua because of the oblique ar-
rangement of the ventral cirri. KENT (1882) also included Gonostomum affine, that is,
KENT himself synonymised Plagiotricha and Gonostomum, considering the latter name as
unsuitable.
The characterisation of Gonostomum is according to BERGER & FOISSNER (1997); see
chapter 2 in the general section for a description of the patterns mentioned. The taxon-
omy is somewhat complicated because of the rather high variability in the number and the
arrangement of the frontal-ventral-transverse cirri, especially in the types species. Previ-
ously, Gonostomum was rather differently classified. For example, CoRUSS (1979) in-
cluded it in the Holostichidae, however, was uncertain and thought that it could belong to
the Oxytrichidae, a classification proposed by Sm.LER (1974b) and TUFFRAU (1987).
SMALL & LYNN (1985) established the Gonostomatidae comprising Gonostomum, Trache-
lochaeta, and Wallackia. Unfortunately, they made a curious nomenclatural mistake in
that they transferred the type of Gonostomum, G. affine, to Trachelostyla. According to
HEMBERGER (1982) and SONG (1990), morphogenesis of G. affine and G. strenuum pro-
ceeds typically oxytrichid, that is, 6 frontal-ventral-transverse cirri primordia occur (Fig.
120a-j, 122f-u). Furthermore, morphogenesis proved that G. affine has "postoral" ven-
tral cirri; however, during interphase they are not located in the postoral area, but right of
the adoral zone of membrane lIes (Fig. 117r).
Gonostomum develops long cirral streaks during early morphogenesis. Later, these
so-called primary primordia divide transversely to form secondary primordia, the usual
 Gonostomum 369

oxytrichid cirral anlagen in proter and opisthe. This type of primordia fonnation occurs
also in Urosoma emarginata (Fig. 127m-p) and Tachysoma pellionellum (Fig. 135j-m).
Urosoma even has the same type of oral apparatus (Gonostomum pattern), suggesting a
rather close relationship of these two taxa (BERGER & FOlSSNER 1997, FOlSSNER 1983a).
Gonostomum franzi FOlSSNER, 1982 was transferred to Kahliella CORLISS, 1960 be-
cause of deviating morphogenetic data: K. franzi (FOlSSNER, 1982) BERGER & FOlSSNER,
1988b; recently, EIGNER (1995) classified this species in Orthoamphisiella EIGNER &
FOISSNER, 1991 (0. franzl). The oral apparatus of Kahliella CORLISS, 1960, Wallackia
FOISSNER, 1976 and Cladotricha GAIEVSKAIA, 1925 looks like that of Gonostomum, indi-
cating parallelism or a close relationship of these taxa (BERGER & FOISSNER 1987a, BLAT-
TERER & FOlSSNER 1988, EIGNER 1995).
FOISSNER (1982) and I agree with the comprehensive list of synonyms for the type
species proposed by BUITKAMP (1977a), and Gonostomum parvum LEPSI, 1947 is a spe-
cies indetenninata (Fig. 228h). Thus, Gonostomum comprises only four species at the
present stage of knowledge.
So far it was generally assumed that the short undulating membrane is the endoral,
which consists of only few loosely arranged basal bodies (FOISSNER 1982). However,
scanning electron micrographs reveal that this is the paroral (Fig. 21d, e, 119b), which is
arranged on the ventral surface and not in the buccal cavity like the endoral.

Key to species

1 2 macronuclear nodules (for example, Fig. 117a) .......................... 2

10--17, usually 14 or 15 macronuclear nodules (Fig. 123a-t) ... G. kuehnelti (p 391)
2 Left of right marginal row 1 (rarely 2) frontoventral row(s); dorsal cilia 5-9 J.lm long
(Fig. 124a, b) ....................................... G. gonostomoida (p 392)
Continuous frontoventral row lacking; dorsal cilia about 3 J.lm long (for example,
Fig. 117d, 122d) .................................................... 3
3 4-6 migratory cirri (Fig. 122d; very important, invariably difference to G. affine);
number of "postoral" ventral cirri distinctly increased, fonning an oblique
row with 6-8 cirri at level of proximal portion of adoral zone of membranelles; left
marginal row tenninates behind transverse cirri (Fig. 122a, d) . G. strenuum (p 384)
Usually 2 migratory cirri (Fig. 117r); usually 3 "postoral" ventral cirri, arranged
rather irregularly ahead of level of buccal vertex; left marginal row tenninates at
level of transverse cirri (Fig. 117a, d, f, h, 0, r, 11ge) ............ G. affine (p 369)

Gonostomum affine (STEIN, 1859) STERK!, 1878 (Fig. 20c, 21d, e, 117a-z,
118a-t, 119a-f, 120a-j, Tables 3, 23, 27)

1859 Oxytricha affinis STEIN, Organismus der Infusionsthiere I, p 186 (Fig. 117j-I).
1865 Oxytricha affinis - QUENNERSTEDT, Acta Univ. lund., 2: 59 (Fig. 1181).
370 SYSTEMATIC SECTION

1878 Gonostomum affine - STERK!, Z. wiss. Zool., 31: 54,57, Tafel IV, Fig. 2.
1882 Plagiotricha (Gonostomum) affinis, STEIN sp. - KENT, Manual infusoria II, p 772.
1888 Stichochaeta affinis (STEIN) - GOURRET & ROESER, Arcbs BioI., 8: 187.
1897 Gonostomum affine (STEIN) - SVEC, Bull. int. Acad. tcheque Sci., 4: 34,46 (Fig. 118m).
1911 Gonostomum affine (STEIN) - GOODEY, Proc. R. Soc., 84: 169 (Fig. 1180).
1926 Gonostomum affine, ST. - KOFFMAN, Actazool., 7: 312 (Fig. 118t).
1929 Gonostomum andoi SlllBlNA, Proc. imp. Acad. Japan, 5: 156 (Fig. 117m).
1930 Gonostomum andoi SIllBlNA- SIllBlNA, J. imp. agric. Exp. Stn Nishigahara, 1: 200 (Fig. 117m).
1932 Gonostomum (Oxytricha) affine (STEIN, 1859) - KAHL, Tierwelt Dtl., 25: 598 (Fig. 117v-z).
1934 Gonostomum affine STEIN - GRANDORI & GRANDORI, Boll. Lab. Zool. agr. Bachic. R. 1st. sup. agr. Mi-
lano, 5: 287, Tavo1a XIII, fig. 270b.
1942 Gonostomum algicola GELLERT, Acta Sci. math.-nat. Univ. Kolozsvar, 8: 23 (Fig. 118j, k; new syno-
nym; according to W. FOISSNER [pers. comm.], a distinct species).
1952 Gonostomum affine STEIN 1859 - SRAMEK-HuSEK, Cs1ki1 BioI., 1: 183 (Fig. 117s).
1952 Gonostomum affine STEIN 1859 - SRAMEK-HUSEK, Chekh. BioI., 1: 373, 376 (Fig. 117s).
1953 Gonostomum affine - JIROVEC, WENIG, Forr, BARroS,WEISER & SRAMEK-HUSEK, Protozoologie, p 514
(Fig. 117s).
1953 Gonostomum affine (STEIN 1859) - WENZEL, Arch. Protistenk., 99: 109 (Fig. 118c-e).
1956 Gonostomum spirotrichoides GELLERT, Acta bioI. hung., 6: 347 (Fig. 118t).
1956 Gonostomum bryonicolum GELLERT, Acta bioI. hung., 6: 348 (Fig. 118g).
1956 Gonostomum ciliophorum GELLERT, Acta bioI. hung., 6: 348 (Fig. 118h).
1957 Gonostomum geleiiGELLERT, Annis Inst. bioI. Tihany, 24: 19 (Fig. 118b).
1957 Urosoma macrostoma GELLERT, Annis Inst. bioI. Tihany, 24: 21 (Fig. 118a; new synonym).
1963 Gonostomum strenuum (ENGElMANN) - LUNDIN & WEST, Free-living protozoa, p 67 (Fig. 117u; misiden-
tification).
1967 Gonostomum affine (STEINE) KARL - CHARDEZ, Revue ilcol. BioI. Sol, 4: 294 (Fig. 118s; incorrect spell-
ing of STEIN)
1972 Gastrostyla affine (STEIN, 1859) - BORROR, 1. Protozool., 19: 14.
1972 Trachelostyla bryonicolum (GELLERT, 1956) - BORROR, J. Protozool., 19: 15.
1972 Trachelostyla ciliophorum (GELLERT, 1956) - BoRROR, J. Protozool., 19: 15.
1972 Trachelostyla spirotrichoides (GELLERT, 1956) - BORROR, J. Protozool., 19: 15.
1972 Trachelostyla geleii (GELLERT, 1957) - BORROR, 1. Protozool., 19: 15.
1972 Trachelostyla macrostoma (GELLERT, 1957) - BORROR, 1. Protozool., 19: 15.
1973 Gonostomum affine STEIN, 1859 - LEPINIS, GELTZER, TSCIllBISOVA & GEPTNER, Key to soil protozoa,
p 154 (Fig. 118i).
1974 Gonostomum algicolum GELLERT - STILLER, Fauna Hung., 115: 93.
1974 Trachelostyla canadensis BUITKAMP & WILBERT, Acta Protozool., 13: 208 (Fig. 117p).
1977 Trachelostyla affine (STEIN) - BUITKAMP, Decheniana, 130: 123, 125 (Fig. 1170, q).
1978 Gonostomum affine - SMITH, Br. Antarct. Surv. Sci. Rep., No. 95: 75 (Fig. 118p).
1981 Gonostomum affine STEIN 1859 -CHARDEZ, Revueverviet. Hist. nat., 38: 53 (Fig. 117t).
1982 Gonostomum affine (STEIN, 1859) STERKI, 1878 - FOISSNER, Arch. Protistenk., 126: 77 (Fig. 117a-i;
authoritative redescription; 1 slide of neotype specimens is deposited in the OberOsterreichische Lan-
desmuseum in Linz, Upper Austria).
1984 Gonostomum affine (STEIN, 1859) STERKI, 1878 - MAEDA & CAREY, Bull. Br. Mus. nat. Hist., 47: 9.
1984 Gonostomum algicola GELLERT, 1942 - MAEDA & CAREY, Bull. Bf. Mus. nat. Hist., 47: 12.
1984 Gonostomum bryonicolum GELLERT, 1956 - MAEDA & CAREY, Bull. Br. Mus. nat. Hist., 47: 15.
1984 Gonostomum ciliophorum GELLERT, 1956 - MAEDA & CAREY, Bull. Br. Mus. nat. Hist., 47: 16.
1984 Gonostomum spirotrichoides GELLERT, 1956 -MAEDA & CAREY, Bull. Br. Mus. nat. Hist., 47: 13.
1985 Gonostomum affine - FLEURY, mODE, DEROUX & FRYD-VERSAVEL, Protistologica, 21: 509 (Fig. 119a).
1985 Gonostomum affine STEIN, 1859 - LOFI'ENEGGER, FOISSNER & ADAM, Oecologia, 66: 574 (Fig. 117n).
1985 Trachelostyla affine - SMALL & LYNN, Illustrated guide to the protozoa, p 461.
 Gonostomum 371

d e

• .
. )k
..
f :

, ,

, ~ ,
, ;
;

, ;
,
,
;
, ,
~

, ,
, ; , ,
, ,
~

,
, , , .
, .
,
'"
-TC
CC-
g h

Fig.1l7a-i Gonostomlltn affine (from FOISSNER 1982. IH:, from life; d, f-i, protargol impregnation; e, wet sil-
ver impregnation). a-c) Ventral, dorsal, and lateral view, IH: = 100 11m. d, f, h, i) Ventral infraciliature of
specimens from various populations, d, f= 65 11m, h = 70 11m, i = 50 11m. e, g) Silverline system and dorsal in-
fraciliature, e, g = 65 11m. CC = caudal cirri, FB = defecation ball, TC = transverse cirri; p 369.
372 SYSTEMATIC SECTION

•
.. • • •
•

O~
-i%!
•
::.::

::::
=: ,(}~ •
•
•
•
•
~ •
~
=:::
-..
•
,
::.:;
.
.
• ....
•• • 1 ,:

a
o
o 0
• p q
 Gonostomum 373

1986 Gonostomum affine (STEIN, 1859) STERKI, 1878 - DRAGESCO & DRAGESCO-KERNEIS, Faune tropicale, 26:
453.
1988 Gonostomum affine - BERGER & FOISSNER, Arch. Protistenk., 136: 67, 74 (Fig. 117r).
1995 Gonostomum affine (STEIN, 1859) - SHIN & KIM, Korean J. Zoo!., 38: 161 (Fig. 119d-t).
1997 Gonostomum affine - BERGER & FOISSNER, Arch. Protistenk., 148: 132, 133 (Fig. 20c, 21d, e).

Nom e n c I at u rea n d t a x 0 nom y: Gonostomum affinis in CU'ILER (1920,

P 136) and Oxytricha affinis in GRANDORI & GRANDORI (1934, P 287) are incorrect subse-
quent spellings. Gonostomum affine shows a high intra- and inter-population variability
in body size and ventral infraciliature (FOISSNER 1982), strongly suggesting that this very
common soil ciliate is a sibling species complex. At the present state of knowledge, how-
ever, I agree with the comprehensive list of synonyms proposed by BUITKAMP (1977a),
and consider G. algicola and Urosoma macrostoma as further synonyms. MAEDA & CA-
REY (1984) recognised most of these species using, for example, the number of transverse
cirri for species distinction. The redescriptions by DELHEZ & CHARDEZ (1970; Fig. 228d)
and ROSA (1957a; Fig. 233b) are insufficient.
Some populations of Gonostomum affine show the typical number of 4 frontoventral
cirri and 3 "postoral" ventral cirri (Fig. 1170, r, 11ge, 120a). Conversely, G. strenuum
has a distinctly higher number of cirri (Fig. 121, 122d). However, there transitions exist
between these two patterns (for example, Fig. 117f, m, 1180 and G. affine sensu NIESSEN
1984); thus, synonymy ofG. affine and G. strenuum cannot be excluded.

M 0 r p hoI 0 g y and b i 0 log y: Size rather variable: in life 85-116 ~m long and
3.5-4 times as long as broad (STEIN 1859b), 50-80 ~m (SRAMEK-HuSEK 1952a, b),
63-125 ~m (mean = 83 ~m, n = 26; WENZEL 1953), up to 160 ~m long (DINGFEIDER
1962). Outline long elliptical or spindle-shaped, broadest usually at level of buccal
vertex. Anterior end usually narrow, posterior broadly to narrowly rounded. Invariably 2
spherical or ellipsoidal macronuclear nodules slightly left of median, usually distinctly
separate from each other; nodules after protargol impregnation 8-23 x 4-11 ~m (mean =
13 x 6 ~m). 1-4, usually 2 spherical or ellipsoidal, after protargol impregnation about
1.3-4.0 ~m sized micronuclei. Contractile vacuole during diastole usually with an ante-
rior collecting canal (Fig. 117b), systole after 6-21 s (GELLERT 1956). Cortical granules
(mucocysts?) colourless, spherical, <1 ~m, loosely arranged in indistinct longitudinal
rows and granules, therefore difficult to discern (Fig. 119c; FOISSNER 1986b). Cytoplasm
colourless, with cytoplasmic crystals, greasily shining, 1-3 ~m sized globules, and about
5-10 ~m sized food vacuoles. Rapid movement, with jerky, relatively long interruptions;
swimming in lively rotation about longer body axis.

~ Fig. 1l7j-q Gonostomum affine (j-I, after STEIN 1859b; m, after SHlBUYA 1929; n, from LOFfENEGGER et aI.
1985; 0, q, from BUITKAMP 1977a; p, from BUITKAMP & WIlBERT 1974. j-n, from life; o-q, protargol impregna-
tion). j, II, m, n) Ventral views,j, k = 85-116 flm, m = 100 flm, n = 1\0 flm. I, q) Ventral view and ventral in-
fraciliature of postconjugational specimens, q = 140 flm. Arrowhead in (q) marks left, slightly enlarged frontal
cirrus. 0, p) Ventral infraciliature, 0 = 105 flm, p = 60 flm. Arrowheads in (0) mark left, slightly enlarged fron-
tal cirrus and caudal cirri. Arrows denote "postoral" ventral cirri; p 369.
 374 SYSTEMATIC SECTION

Fig. 117r-z, 118a, b Gonostomum affine (r, from BERGER & FOISSNER 1988b; s, from SRAMEK-HuSEK 1952a,
b; t, after CHARDEZ 1981; u, from LUNDIN & WE!IT 1963; v-z, from KAlIL 1932; a, b, from GELLERT 1957. r, pro-
targol impregnation; s-z (t?), from life; a, b, sublimate-glycerine-alcohol fixation and opalblue staining after
BRESSLAu). r) Ventral infraciliature, 60 J.lm. The cirri originating from the primordia I-VI are connected by
broken lines. The posterior cirrus of primordium IV (IV/2) and the two cirri of primordium V (V/3, Vl4) are
the "postoral" ventral cirri which are arranged right of the proximal portion of the adoral zone of
membranelles. Arrows mark the 2 migratory (= frontoterrninal) cirri. S-z, a, b) Ventral views showing variabil-
ity ofciliature causing a rather high number of synonyms (names, see list of synonyms), s, u, x-z= size not in-
dicated, t = 105 J.lm, v = 120 J.lm, w = 100 J.lm, a = 75 J.lm, b = 50 11m; p 369.
 Gonostomum 375

Endoral after protargol impregnation 7-11 ~m, paroral 12-16 ~m long (SHIN & KIM
1995). Frontal, ventral, and transverse cirri slightly enlarged, according to BUITKAMP
(1977a) left frontal cirrus larger than middle and right one (Fig. 1170, q). Nwnber and ar-
rangement of frontoventra1 and transverse cirri rather variable, cirri on frontal area some-
times form a short, slightly oblique row (for example, Fig. 117i-m). Figure 117r shows a
specimen where the oxytrichid lineage, that is, the typical nwnber and arrangement of
frontoventral and postoral ventral cirri, is well recognisable. The posterior cirrus of streak
IV and the two cirri of streak V form the "postoral ventral cirri", identical to, for instance,
in Oxytricha or Tachysoma, where these 3 cirri are, however, arranged just behind the
buccal vertex. All cirri about 12 ~m long. Dorsal cilia 3 ~m, according to BUITKAMP
(1977a) 3-7 ~m long; GELLERT (1956) described "long cilia" for Gonostomum bryonico-
fum and G. spirotrichoides. Very rarely 4 dorsal kineties.
Cyst spherical, about 33 ~m across, with a smooth, thin wall (Fig. 118n; CHAUDHURI
1929, SANDON 1927); according to an unpublished scanning electron micrograph (kindly
supplied by W. ForssNER), however, with small warts.
M 0 r p hog e n e sis of cell division is described by HEMBERGER (1982; Fig.
120a-j) and NIESSEN (1984). It proceeds rather similarly to in G. strenuum. The formation
of primary primordia is considered as synapomorphy of the Gonostomum-Urosoma-
Tachysoma pellionellum group. Conjugating stages and postconjugates usually slender
(Fig. 1171, q; CHAUDHURI 1929).

o c cur r e n c e and e colo g y: Very likely the most widely distributed and most
common, and often also the most abundant hypotrich in almost all terrestrial habitats
(soil, litter, mosses, lichens) all over the world (for example, ForssNER 1997b). Locus
classicus is the "St. Procop-Tal" near Prague, Czechoslovakia, where STEIN (1859b) dis-
covered Gonostomum affine in marshy water with decaying leaves of trees. Indicates ter-
restrial influence in limnetic habitats, for example, after inundation (ForssNER et al. 1995).
Records largely substantiated by illustrations: soils from Austria (AEsCIIT & ForSSNER
1992, 1993, ForssNER 1987b, ForssNER & ForssNER 1988, ForssNER et al. 1990, LOFrENEG-
GER et al. 1988, PETZ et al. 1988) and Belgiwn (CHARDEZ 1967, 1981); forest mosses and
pond in Czechoslovakia (SRAMEK-HuSEK 1952a,b, SVEC 1897); agricultural soils in Eng-
land (GOODEY 1911); dunghill in France (FLEURY et al. 1985); upper soil layer (0-5 cm) of
a pasture and a mixed forest near Bonn, Germany (BUITKAMP 1977a); mosses, coniferous
and deciduous litters, lichens, rotten wood, and moist layer of algae on a window of a
greenhouse (PH 4.8-5.2) in Germany (KAHL 1932, WENZEL 1953); soils in Hungary
(GELLERT 1942, 1956, 1957); Italy (GRANDORI & GRANDORI 1934); terrestrial and limnetic
habitats in Sweden (KOFFMAN 1926, QUENNERSTEDT 1865, 1869); USSR (LEPINIS et al.
1973); subtropical soil in China (SHEN et al. 1992); soil from the experimental farm in
Nishigahara near Tokyo, Japan (SHIBUYA 1929, 1930); terrestrial (?) habitats in Ullung Is-
land, Korea (SHIN & KIM 1995); prairie soil from Canada (BICK & BUITKAMP 1976, Burr-
KAMP & WILBERT 1974); limnetic habitats from the Upper Peninsula of Michigan, USA
376 SYSTEMATIC SECTION
 Gonostomum 377

Fig. llSn-t Gonostomum affine (n,

> after SANDON 1927; 0, after GooDEY
~2 1911; p, from SMIllf 1978; q, original
kindly supplied by W. FOISSNER; r,
OJ
o from LOFTENEGGER et aI. 1985; s, from
CHARDEZ 1967; t, after KOFFMAN 1926.
--21 n-<j, s, t, from life). u) Resting cyst,
-------10°C according to text 33 11m, according to
legend of figure only 17 11m across.
r o-q, s, t) Ventral views, 0, S = 95 Jim,
5 10 15 20 25 30 p = 85 11m, t = 80 11m. r) Growth
days curve at 10 °C and 21°C; P 369.

(LUNDIN & WEST 1963); mosses from USA (KAHL 1932); soils and moss peat of Sub-
antarctic and maritime Antarctic islands (SMITII 1978, 1981, 1982).
Records from freshwater habitats not substantiated by illustrations: Belgium (CHAR-
DEZ 1984); running waters in Bulgaria (DETCHEVA 1979b, 1981, DETCHEWA 1972, RUSSEV
et al. 1976, 1984); submerged and wet mosses and other sites in Czechoslovakia (TIRJAK-

Fig. USc-m Gonostomum affine (c-e, from WENZEL 1953; f-h, from GELLERT 1956; i, from UPINIS et aI.
1973;j, k, after GELLERT 1942; I, after QUENNERSTEDT 1865; m, after SVEC 1897. c-e, i, I, m, from life; f, subli-
mate fixation and nigrosin staining; g, h, j, k, sublimate fixation and opalblue staining after BRESSLAU). c-m)
Ventral views and dorsal view (k) of various synonyms, c, d = 95 11m, e = 75 11m, f= 100 11m, g, h = 70 Jim, i
= 75-115 Jim,j, k = 60-100 Jim, 1= size not indicated, m = 130 11m; p 369.
378 SYSTEMATIC SECTION

..
. . '

. '.

a . ..
c

Fig. 119a-c Gonostomum affine (a, from Fl£URY et aI. 1985; b, c, originals kindly supplied by W. FOISSNER. a,
protargol impregnation; b, scanning electron micrograph; c, from life). a) Ventral infraciliature, bar = 20 11m.
Arrow marks short paroral which consists of loosely arranged basal bodies. b) Cell from a soil from Mitten-
wald (Germany) in ventral view. Small arrow denotes posterior-most "postoral" ventral cirrus, large one para-
raI, which consists of few, loosely arranged cilia c) Dorsal view of a specimen from a Japanese population.
The <111m sized, colourless spherical cortical granules (mucocysts?) are arranged in indistinct rows and are
rather difficult to discern; p 369.

ovA 1992b, TIRJAKovA & MAns 1987a); Denmark (NIELSEN 1960); in May about
20 indo ml'( in the littoral of a pond near Randan and in mineral waters in France (GRO-
LIERE 1977, GROLIERE & NJINE 1973, CHAOUITE et al. 1990); ephemeral water bodies in
Gennany at pH 6.2-7.2 (DINGFELDER 1962); oligo- and mesosaprobic brook and rivers in
Bavaria, Gennany (FOISSNER 1997a, FOISSNER et al. 1992a, b); Schussen River in Ger-
many (WETZEL 1928a); caves in Gennany (GRIEPENBURG 1934, 1935) and Italy (BOSCOLO
1968, STELLA & SALVADOR! 1953; see also GITILESON & HOOVER 1969); pelagial ofa side-
branch of the Danube River in Hungary with 0.9 % dominance in late September (BEREC-
ZKY 1991); thennal waters in Hungary (DADAY 1891); abundant in polluted water in Po-
land (WRZESNIOWSKIEGO 1866); Romania (LEPSI 1932); Switzerland (MERMOD 1914, MEs-
 Gonostomum 379

o
() fJ
()
------
~

'----

JJ
0: ~

- ,, I
/

, , /
/
o J
D0
o I
o /
I
/
I
/
/
I I
e -- f
Fig.1l9d-f Gonostomum affine (from SHIN & KIM 1995. d, from life; e, f, protargol impregnation). d) Ventral
view, 82 ~m. e, 1) Ventral and dorsal infraciliature, 82 ~m. Note that in Gonostomum the postoral ventral cirri
(arrowheads) are arranged not behind but beside the adoral zone of membrane lies; p 369.

SIKOMMER 1954); Tibetan Plateau (WANG 1977); Turkestan (DADAY 1903, 1904); littoral
of Lake Baikal (GAJEVSKAJA 1927, GAJEWSKAJA 1933); China (HAN & HAo 1995, as G.
afxfine); at 24-27°C and pH 6-7 in Michigan waters, USA (CAIRNS & YONGUE 1966);
Brush Lake, Ohio, USA (RIDDLE 1905); Amazon River (CAIRNS 1966a); Madagascar
(SONDHEIM 1929),
Records from terrestrial habitats not substantiated by illustrations: Belgium (CHARDEZ
1983a); Bulgaria (DECHEVA 1966, 1968, 1970, 1973, DETCHEVA 1972b); Czechoslovakia
(TJRJAKovA 1988, TJRJAKOvA & MATIS 1987a, b); Denmark (BRUNBERG NIELSEN 1968,
STOUT 1968); England (CUTLER 1920, STOUT 1963); soil lichens from the sea shore at
Banyuls-sur-Mer, France (ForssNER 1986c); forest humus in France (PALKA 1991, 1993);
beech forest, limed spruce forests, polluted soils, and other sites in Germany (FUNKE
1986, LEHLE 1989, 1993, 1994, LEHLE et al. 1992, NIEBUHR 1989, VARGA 1935b); Green-
land (DIXON 1939); Scoresby Land, East Greenland (STOUT 1970); Hungary (HORvATH
1950); Italy (COPPA 1921, LUZZATTI 1938, STELLA 1948); Romania (RADu & TOMESCU
380 SYSTEMATIC SECTION

....
..
'

Q.l
II "-
.....

I I

".

\ \
I I I I I \

1J

00
()
I ,

, I I I , •• , , I
, C=>
'C=>

------_
........:... .
.3~~-0SI"'-·
...... J , •••• _ _ - - - - - - - -
' '
\ \ \
I ,
, I t I I , '

00 ,
OJ

'CJ
'0

... .....

-'-
~

I , '
, , ,-..~

'c, •
'c) :

, I I I I
 Gonostomum 381

1972, TOMESCU 1978, 1987); Yugoslavia (VARGA 1962); India (BHATIA 1936, SANDON
1927); USSR (DIXON 1937, MIRZA-ZAnE & rnADOV 1988, MORDKOVICH 1986, MUSAEV &
rnADOV 1979, YAKIMOFF & ZERilN 1924); Himalayan region up to 4500 m above sea level
(FOISSNER 1986a); soil from paddy fields in Japan (HINo & MOMlKI 1931, TAKAHASIll &
SUHAMA 1991); bark of Acacia trees and soil in Costa Rica (FOISSNER 1994c, 1995b); sand
from the Coral Pink Sand Dunes near the Zion National Park, Utah, USA (FOISSNER
1994b; further records from the USA: BAMFORTH 1967, 1968, 1969, BOVEE 1979, MARTIN
& SHARP 1983, SANDON 1928); savannah near Lamto, Ivory Coast (BUITKAMP 1977a,
1979); Australia (BLATfERER & FOISSNER 1988, POMP & WILBERT 1988); New Zealand
(STOUT 1958, 1960, 1961, 1978, 1984); some islands in the southern Atlantic Ocean
(SANDON & CUTLER 1924); Antarctica (SMITH 1973a, 1974a, 1975, 1984, 1985).
BOUTClllNSKY (1895) and BUTSCHINSKY (1897) found Gonostomum affine in inland
salt waters of the USSR, WILBERT (1995) in saline lakes in Australia, and FERNANDEZ-
LEBORANS & NoVILLO (1993) in the sub-littoral of the Bay of Biscay. FERNANDEZ-LEBORANS
& NoVILLO (1994) found the synonym G. algicola in the sea at Castro Urdiales, Spain.
WENINGER (1970) recorded G. affine from percolating filters in Austria, and RICHARDS &
CUTLER (1933) from filters for the purification of waste waters from beet sugar factories.
BUCHAR (1958) found G. affine in the foam of cicadas.
Feeds mainly on bacteria (BtnTKAMP 1977a, DINGFELDER 1962, FOISSNER 1987a), but
also on algae, fungi, heterotrophic flagellates, and detritus (GEILERT 1956, HORVATH
1950). Generation time about 13.4 hat 10°C and 10.6 h at about 21°C when cultured
polyaxenically in lettuce medium with crushed wheat grains to support bacterial growth
(LOFrENEGGER et al. 1985); at 5 °C and 30°C culturing was impossible. Highest individ-
ual densities (3800 indo ml- I ) were obtained at 21°C (Fig. 118r). BUITKAMP (1979), how-
ever, recorded Gonostomum affine from 5 °C to 40°C. STOUT (1956) cultured G. affine
in peat or yeast extract. Biomass of 106 individuals about 39 mg (FOISSNER 1987a), 23 mg
(MUSAEV & IBADOV 1980), 20--45 mg (BUITKAMP 1979), about 19 mg (NESTERENKO &
KOVALCHUK 1991). Some autecological data oflimnetic populations from various running
waters in Bulgaria are summarised in Table 27. Autecological data of terrestrial popula-
tions are given in FOISSNER (1981), FOISSNER & PEER (1985), and FOISSNER et al. (1985).
The latter authors found G. affine at pH 3.1 in an alpine pseudogley of a wooded area.
REINNARTH (1979) observed it in the mud-water interface ofa eutrophic pond in Germany
at following conditions: -0.5 to -2 cm sediment depth, pH 7.1-7.6, -149 to +4 mV redox
potential, 0.73-1.33 mg 1-1 NH/-N, and 0.09-0.99 mg I-I N03--N. Does not tolerate hy-
drogen sulphide (WETZEL 1928b). Effect of heavy metals, see FERNANDEZ-LEBORANS et al.
(1987).

~ Fig. 120a-j Gonostomum affine (from HEMBERGER 1982. a-j, protargol impregnation). a-i) Ventral infracilia-
ture and nuclear apparatus of early, middle, and late morphogenetic stages, a = 110 11m. Gonostomum develops
long cirral streaks during early morphogenesis (e); later, these so-called primary primordia divide transversely
to form secondary primordia (t). j) Dorsal infraciliature of a late morphogenetic stage. Arrows mark new cau-
dal cirri of opisthe. OP = oral primordium; p 369.
382 SYSTEMATIC SECTION

Table 23 Morphometric data of Gonostomum affine (afl, from BuITKAMP & WILBERT 1974; at2, from Burr·
KAMP 1977a; aD, 6 populations from the Glockner area from FOiSSNER 1982; af4, 5 populations from the Tull-

nerfeld from FOiSSNER 1982; af5, population from saline Australian soils; af6, from SHIN & KIM 1995), G.
gonostomoida (gon, from HEMBERGER 1985), G. kuehnelti (\rue, from FOiSSNER 1986b), and G. strenuum (str,
from SONG 1990). All data are based on protargol-impregnated specimens. All measurements in micrometres. ?
= sample size not indicated; if only 1 value is known it is listed in the mean column, if 2 values are available
they are listed as Min and Max. CV = coefficient of variation (in %), Max = maximum value, mean = arithme-
tic mean, Min = minimum value, n = sample size, SO = standard deviation

Character Species mean SD CV Min Max n

Body, length afll 59.0 1
at2 1 105.0 1
aD 69.3 16.0 23.0 43 112 26
af4 65.2 10.0 15.3 43 84 26
af5 131.2 13.5 100 150 15
af6 83.1 6.9 8.3 73 97 8
kue 60.2 6.6 10.9 50 71 15
str 103.8 7.9 7.6 88 119 11
Body, width afll 22.2 1
at2 1 37.5
as 23.7 6.6 27.6 16 46 26
af4 23.5 5.7 24.4 16 45 26
af5 55.2 6.3 43 68 15
af6 31.9 8.9 27.8 24 52 8
kue 23.7 2.7 11.6 20 28 15
str 37.0 5.0 13.4 28 44 11
Adoral zone of membranelles, length afll 31.0
at2 1 48.0
at3 33.4 10.7 31.9 21 59 26
af4 31.5 5.8 18.3 21 44 26
af5 61.4 5.7 50 70 15
af6 40.2 5.6 13.8 33 49 8
kue 28.0 1.5 5.4 25 32 15
str 54.5 7.2 13.2 41 69 11
Adoral membranelles, number afll 22.0 I
at2 21 25 ?
as 27.9 5.9 21.0 22 40 26
af4 28.5 3.2 11.3 23 38 26
af5 28.2 I 26 30 15
af6 25.1 2.5 10.1 22 29 8
gon 33.0 ?
kue 26.0 1.2 4.6 24 28 15
str 29.5 2.2 7.3 26 33 11
Macronuc1ear nodules, number kue 14.2 2.1 14.9 10 17 15
Right marginal row, number of cirri afl 21 23 ?
at2 17.0 15 31 ?
aD 20.4 3.6 17.7 14 28 26
af4 18.9 2.0 10.8 IS 24 26
af5 28.7 1.9 26 31 IS
af6 16.4 2.1 12.6 14 20 8
gon 23.0 18 27 ?
 Gonostomum 383

Table 23 Continued

Character Species mean SD CV Min Max n

Right marginal row, number of cirri kue 21.3 1.9 9.0 18 25 15
str 25.8 2.0 7.6 23 29 13
Left marginal row, number of cirri afl 13 14 ?
af2 14.0 II 20 ?
af3 14.5 2.9 19.9 8 21 26
af4 13.6 2.1 15.8 9 18 26
af5 15.2 2.1 11 19 15
af6 10.1 1.7 17.1 8 14 8
gon 14.0 12 15 ?
kue 16.3 1.0 6.3 15 18 15
str 17.7 5.2 29.6 17 21 13
Ventral cirri on frontal area, afl 6.0 5 6 ?
number (frontal and buccal cirri af2 7.0 6 13 ?
not included) af3 8.1 1.4 17.6 7 14 26
af4 8.8 2.9 32.9 6 15 26
af5 9 13 15
af6 7.0 0 0 7 7 8
kue 7.0 0 0 7 7 15
Transverse cirri, number (pre- afl 2.0 ?
transverse ventral cirri af2 0 6 ?
included) af3 3.5 1.9 55.6 0 7 26
af4 4.6 1.0 21.4 3 6 26
af5 1.0 0 0 15
af6 4.1 0.35 8.6 4 5 8
gon 2 3 ?
kue 4.0 0 0 4 4 15
str 5.8 0.8 13.1 4 7 17
Caudal cirri, number afl 3.0 ?
a12 3.0 0 0 3 3 '1
aO 2.9 0.5 16.2 I 4 26
af4 3.1 0.4 12.5 2 4 26
afS 3.0 0 0 3 3 15
af6 3.0 0 0 3 3 5
gon 3.0 ?
kue 3.0 0 0 3 3 15
str 3.0 0 0 3 3 17
Paroral, number of basal bodies af2' 13.0 I
aO 13.5 3.7 27.7 8 23 26
af4 11.1 1.9 17.2 8 14 26
af6 10.6 1.9 18.1 8 14 8
gon' 11.0 I
kue about 10 ?
str' 17.0 I

, From corresponding figures.

384 SYSTEMATIC SECTION

Gonostomum strenuum (ENGELMANN, 1862) STERK!, 1878 (Fig. 121,

122a-u, Tables 3, 23)

1862 Oxytricha strenua ENGELMANN, Z. wiss. Zool., 11: 387 (Fig. 121).
1878 Oxytricha strenua ENGELMANN - STERK!, Z. wiss. Zool., 31: 57 (see taxonomy).
1882 Plagiotricha strenua, ENG. sp. - KENT, Manual infusoria II, p 772.
1888 Stichochaeta strenua (ENGELMANN) - GoURRET & RoESER, Archs BioI., 8: 187.
1932 Gonostomum (Oxytricha) strenuum (ENGELMANN, 1862) - KAHL, Tierwelt Otl., 25: 597.
1972 Gonostomum strenuum (ENGELMANN, 1862) STERK!, 1878 - BORROR, J. Protozool., 19: 14.
1984 Gonostomum strenua (ENGELMANN, 1862) STERK!, 1878 - MAEDA & CAREY, Bull. Br. Mus. nat. Hist, 47: 11.
1990 Gonostomum strenua (ENGELMANN, 1862) - SONG, J. Protozool., 37: 249. (Fig. 122a-u).
1997 Gonostomum strenua ENGELMANN, 1862 - 0LMO & TELLEz, Arch. Protistenk., 148: 191, Fig. 1-16, Ta-
ble 1 (morphometric characterisation and morphogenesis).

Nom e n c I a t u rea n d t a x 0 nom y: Although STERK! (1878) did not use the bi-
nomen Gonostomum strenuum, he is generally accepted as combining author (BORROR
1972a, MAEDA & CAREY 1984) because he wrote about Gonostomum "...eine sehr einheit-
liche wohlumgrenzte Gruppe, zu der offenbar auch o. strenua ENGELMANN ZU ziehen ist".
Incorrect subsequent spellings: Gonostomum stenuum (ENGELMANN) (MAnRAZO-GARIBAY
& L6PFZ-OCHOTERENA 1982, p 292); Gonostomum sternuum (MCCASHLAND 1956, p 38);
Gonostomum strenum (ENGELMANN) (CAIRNS & YONGUE 1973a, P 32).
There is no doubt that Gonostomum strenuum and G. affine are difficult to separate
(especially in life), indicating that they are synonymous. For instance, it could be possible
that G. andoi (Fig. 117m) and G. affine sensu GOODEY (1911; Fig.
1180), which are (preliminary) classified as synonyms of G. affine, are
conspecific with G. strenuum because they have a rather long frontov-
entral row terminating behind the level of the buccal vertex. This row,
the higher number of migratory cirri in Gonostomum strenuum, and
the different sites where the left marginal row terminates support the
recognition of both species.
Gonostomum strenuum sensu LUNDIN & WEST (1963; Fig. 117u) is
classified as G. affine because it has a very low number of cirri. The
KOFFMAN (1926; Fig. 232p) and MAnRAW-GARlBAY & LoPEZ-
OcHOTERENA (1973; Fig. 241 e) populations are insufficiently redescribed.

M 0 r p hoI 0 g y and b i 0 log y: In life 90-120 x 25-50 Jim

(SONG 1990), 80-130 x 20-55 Jim (OLMO & TELLEZ 1997), after EN-
GELMANN (1862) up to 150 Jim long. Outline rather similar to in G. af-
fine, anterior end somewhat pointed, posterior rounded. Macronuclear
nodules after protargol impregnation 13-20 x 6-13 Ilm (mean =
Fig. 121 Gonosto-
mum strenuum
17.1 x 9.1 Ilm; n = 16), conspicuously separate, lying slightly left of
(from ENGELMANN median (Fig. 117a, e). 2-3 (mean = 2.1; n = 17), after protargol im-
1862). Ventral view pregnation 1.9-2.8 Ilm (mean = 2.3 Ilm; n = 16) long micronuclei.
from life, 135 Ilm.
 Gonostomum 385

.
o
• 0

o •

.
e.

..
• 0

00
• 0
c
'0 •
o· 0

. ...
0 0

· '.
0
Q
..
\
"

):~- Cy
0

·.
••••• {

.. •. ....
.. . ',,.:.fo
.
'. -

·· . .
"

0
o ••

'.. . 00

-,;---- - - sG

Fig. 122a-c Gonostomum strenuum (from SONG 1990. a--c, from life). a, b) Ventral views, 110 Ilm. The corti-
cal granules (mucocysts?) and the contractile vacuole are shown only in (b). c) Cortical granules (mucocysts?)
in lateral view, bar = 3 Ilm. CV = contractile vacuole, sO = cortical granules; p 384.

Cortical granules (mucocysts?) small, colourless and thus difficult to discern, after ejection
spindle-shaped, about 1.0-1.5 flm long, do not impregnate with protargol (Fig. 119c).
Frontal cirri slightly enlarged, in life about 20 flm long. Arrangement and number of
frontal-ventral-transverse cirri rather invariable. Number of cirri in streak III (without right
frontal cirrus; Fig. 122d) 2-3 (mean = 2.1; n = 17); streak IV 3--4 (mean = 3.3; n = 17);
streak V (without pretransverse ventral cirri and transverse cirri) 6--8 (mean = 7.2; n = 17);
streak VI (only migratory cirri) 4-6 (mean = 4.4; n = 17). Usually only 2 transverse cirri in
the narrow sense; remaining cirri near them must be designated as pretransverse ventral
 Vol
00
0\

't>. FC
FTC--

\1 "
\ " ~

\ "
" " \
'f..l 'I
"
" :
~ ~ " <' ~ \/J
'\
/' j
'(
"
"jl -<
\/J
" ....,
/ tTl
,?I ~I "
,I
~'-,"
/ y
i /' "
"
7
#~~ ¢ ~ ~
....,
~ (=)
{ ,/
" ". \/J
, ~ r~ " +.~ tTl
y " (J
, ,/ " ....,
\ p
\
\
,
I
, " ~
\
.I ~ @
t-- '\t
I
I p
\
\
\
I
I
I
t ~ I f"
t \' "
\ I ~ .I '\ "
\ I \ t
, 't
, \t t
" I J- ~----, "" ~
I , '" \ ,~ ,;; ~ "
PRVC - \ - :..~+-~ "
, i- I-' , ' " ,/ '--/
"d '/ e
, "" --=- CC
Fig. 122d-C Gonostomum strenuum (from SONG 1990. Protargol impregnation). d, e) Ventral and dorsal inftaciliature oCa non-dividing cel\, 100 ~m. Cirri originating
from same primordium connected by a broken line. 1) Very early morphogenetic stage. Arrow marks oral primordium. BC = buccal cirrus, e = endoral, FC = left frontal
cirrus, FTC = 4-{j migratory cirri, p = paroral, PRVC = pretransverse ventral cirri, I-VI = frontal-ventral-transverse cirri primordia I-VI, 1-3 = dorsal kineties; p 384.
 ""
"
~ ,
"" "
" jJ \ ,
\ ,
" ~ ,
" \ ,
" --~~ \, , c::J
c
;:s
"
~ " ~
\ 2
"
" \ ~
" " 1:,< ": '. , §
"
" " "
"

"
" I:
'" '" f "
"".. .",,/g.
..
Fig. 122g-j Gonostomum strenuum (from SONG 1990. g-j, morphogenetic stages after protargol impregnation). g, h) Ventral infraciliature of early stages, g = 115 11m,
h= 110 11m. i,j) Ventral and dorsal infraciliature ofa middle stage, 125 11m. Note that primary primordia are formed! Arrow in (i) marks primordium I ofproter, arrows w
00
in (j) denote sites where proliferation in dorsal kineties commences; p 384. -....l
 w
00
00

\
\1 I
<' ,1
" <' I' I I ~
"" ~ ). \'
/ '" '" JV I~I II ~" ;\~ I"
,?,. ,;
rf
X, p
<,ff!
v j \ 1 , \ t ~, l
/
i "
t, 1 VI { . \ ' I
~ !
\ \'
'\
) ~ """
; I j~ 't \ ~
\' I ~ \(
p \ jj~ C/l
,ln 1~ I :I,~ ,
'"" >-<:
N ,
'" 'I I '"
t
,I ~~~
{ 17, \~ ~ /~" C/l
/
}; (! ,:. {:,1p~ @
I l
(! •• '
I (! f #J4~ '" ~ "I~
~ \
(! ,~" .. ,
r
>...
(!
() I
1/\ 1,Ii' ~ . .
\ \ if § ~
1 , T" _' ~
::l
t.}
(J \ \,:, ! ~ " . ("')
r\l ~ I
J ~ " C/l
i
: \ \\pl,1
", i =' \ < f
tTl
("')
~ ~ .....,
I (!
~,
tlA-' \"i.~
\ t " " \' j;:= , ~
\\ ~ '. I "'" , (I

j
~
\ ~
~
"'.\.,. ~ .I --:
~\ )"
\ J
\ it \, If J / \ ,J
p t \ ,J
.. '"
" ., ( '\ r-/
" " <> . k
m ""~n
!.....!...--

Fig. 122k-n Gonostomum strenuum (from SONG 1990, k-n, morphogenetic stages after protargo\ impregnation), k, I) Ventral and dorsal infraciliature of a middle stage,
130 Ilm. Arrows mark primordia of marginal rows. m, n) Ventral and dorsal infraciliature of a late stage, 140 Ilm. I-VI = frontal-ventral-transverse cirri primordia I-VI;
P 384.
 Gonostomum 389

\
\

, ~
, ~
,
,, ~
r
, ,,
I

,
, \
I
\
',-1 '-2 ,
\

,, ,
\

,,
,, \/
! I
"'.. {! .(
I !
\
p

Fig. 122o-q Gonostomum strenuum (from SONG 1990. o-q, morphogenetic stages after protargol impregna-
tion). 0, p) Ventral and dorsal infraciliature of a late stage, liS /lm. q) Ventral infraciliature of a very late
stage, 160 /lm. I-VI = frontal-ventral-transverse cirri primordia I-VI, 1,2 = dorsal kineties 1 and 2; p 384.

cirri. Occasionally one or two cirri in front of these cirri. Right marginal row terminates at
level of transverse cirri, left row usually behind right transverse cirrus, thus posterior-most
left marginal cirri difficult to discern from the about 20 11m long caudal cirri. Cyst spheri-
cal, with smooth wal~ in life about 40 11m across (OLMO & TELLFZ 1997).
M 0 r p hog e n e sis of cell division is described in great detail by SONG (1990;
Fig. 122d-u). As in the type species, primary primordia are formed (Fig. 122i, k). The
number of cirri originating from the frontal-ventral-transverse streaks I-VI are as follows
(see Fig. 122d, 0, q, s, u): 1,2,2-3 (4?), 3-4, 8-11, and 7-10. Morphogenesis of dorsal
infraciliature is in Gonostomum pattern, that is, no fragmentation of kinety 3 and no dor-
somarginal row. Further details, see Figures 122f-u and OLMO & TELLEZ (1997).
o c cur r e n c e and e col 0 g y: According to literature rare in limnetic and
terrestrial habitats; however, it cannot be excluded that G. strenuum was sometimes con-
fused with the rather common G. affine. The locus classicus is near Leipzig, Germany,
 W
\0
o
,-
~
J-
J-
I
I
I
I
I
I
I
"W> I
- "~.' , I
' '~(,l
I
I
I
~ , \J, , \
I
C/1
-J, , I
-<
C/1
I) ~ I >-j
I, I tTl
~ I
,,
.' I
, \
I ~
\ (5
\
,
i·.\",.
"'~ C/1
• I •
... .. ...
I
.'
.'
.:,\,,'
"; ,.~\'
~
\

\
t" ,,
,, tTl
(j
" , ::l
" "t,;"~:: '.
"',' I §2
III
'/~ I
, f; I
,,,,.
~ \ I" "

§,
,~~ '~ I I
'
\ \, I
...
..
,
.., ~":~.' :i\•
" ",',;',.. I,
.., p\ ,
, ...
J
. ---cc
~
." .. .., cc .' u

Fig. 122r-u Gonostomum strenuum (from SONG 1990. Protargol impregnation). r-t) Dorsal and ventral infraciliature of very late morphogenetic stages, r-t =
155-160 11m. u) Postdivider, 75 11m. Cirri originating from the same streak are connected by a broken line. CC = caudal cirri, p = parora!; p. 384.
 Gonostomum 391

where ENGELMANN (1862) discovered G. strenuum in a ditch with Lemna sp. SoNG (1990)
isolated it from the upper soil layer in the Baguan hills (about 50 m above sea-level) in
Qingdao, China. OLMO & TEU.EZ (1997) found it in moss samples from emergent stones at
the bank of the Guadarrama River, Spain (1188 m above sea-level). Records not substanti-
ated by illustrations: mud of Loch Leven, England (BRYANT & LAYBOURN 1974); Indone-
sia 01AN OYE 1922); Cape Fear River near Fayetteville, North Carolina, USA (CAIRNS &
YONGUE 1973a); pond on Mount Desert Island, USA (MCCASHLAND 1956); agricultural
soils in Czechoslovakia (TIRJAKOVA 1988); Gulf of Mexico (BOVEE in BORROR 1962).
Feeds on bacteria and green algae and can be cultured in Eau de Volvic with some
crushed wheat grains (SONG 1990). CAIRNS & YONGUE (1973a) found G. strenuum at fol-
lowing conditions: 23-26°C, pH 6.2-7.1, 5.8-8.5 mg 1'( O2, 0.9-3.8 mg 1-( BODs, 18 to
24 mg 1'( Cl-, 0.44-0.59 mg 1-( NH/-N, 0.08-0.67 mg 1-( N03-·N, 0.08-0.14 mg 1-(
POl--P, 0.16-0.21 mg 1-( Fe2-. Does not tolerate hydrogen sulphide (WETZEL 1928b).

Gonostomum kuehnelti FOISSNER, 1987 (Fig. 123a-f, Table 23)

1987 Gonoslomum kuehnelli FOISSNER, Sber. ost. Akad. Wiss., 195: 263 (1 slide ofholotype specimens and 1
slide of paratype specimens are deposited in the OberOsterreichische Landesmuseum in Linz, Upper
Austria).

T a x 0 nom y: Differs from G. affine unequivocally in the number of macronuclear nod-

ules and the almost invariable number and arrangement of frontal, ventral, and transverse
cirri; however, using only the infraciliature, a clear separation is not always possible be-
cause some populations of G. affine possess an almost identical cirral pattern (cp. Fig.
123e with Fig. 117r). Gonostomum kuehnelti is similar to Kahliella franzi (FOlSSNER,
1982) BERGER & FOISSNER, 1988b as concerns body shape and number of macronuclear
nodules. However, K. franzi is larger, has no cortical granules, and has more ventral cirri,
which are arranged in distinct rows.

M 0 r p hoi 0 g y and b i 0 log y: In life 60-90 x 25-35 Jim. Outline slender to

broad oval, posteriorly more rounded than anteriorly. Macronuclear nodules spherical or
ellipsoidal, lying slightly left of median. Micronuclei do not impregnate with protargol
during interphase, in morphogenetic stages about 7 micronuclei were observed. Contrac-
tile vacuole with two collecting canals. Cortical granules (mucocysts?) rod-shaped,
1.0-1.5 !lm long, loosely arranged, colourless, stain red with methyl green-pyronin, usu-
ally they were ejected under the cover slip, hence they can be easily overlooked. Cyto-
plasm strongly opaque by many about 10!lm sized food vacuoles and 1-3 !lm sized,
shining inclusions. Movement slowly gliding or rotating.
Adoral zone of membranelles displaced slightly more inwards than in Gonostomum
affine, proximal portion covered by a broad protoplasmic lip. Arrangement and number
of frontal, ventral, and transverse cirri almost invariable (Fig. 123e). Left frontal cirrus
392 SYSTEMATIC SECTION

Fig. 123a-£ Gonoslomum kuehne/Ii (from FOISSNER 1987f. a-d, from life; e, t; protargol impregnation). a, b)
Ventral and lateral view, 90 Jim. c) Pellicle with 1.0-1.5 Jim long cortical granules (black rods). d) Dorsal view
showing cortical granules and contractile vacuole during diastole. e, t) Ventral and dorsal infraciliature,
65 Jim. Large arrows in (e) mark migratory cirri VIl3 and VIl4, small arrows denote "postoral" ventral cirri.
Arrow in (f) marks caudal cirri. sG = cortical granules; p 391.

often slightly enlarged. Marginal rows open posteriorly, right row always terminates more
anteriorly than left one, which ends in median of cell. All cirri about 20 Ilm long.

o c cur r e nee and e colo g y: In terrestrial habitats often associated with the very
common G. affine; thus, nuclear apparatus should be checked. Locus classicus is Seekir-
chen, Salzburg, Austria, where FOlSSNER (1987f) discovered G. kuehnelti in the upper soil
layer of an organically farmed field; for detailed description of this site see FOISSNER
(l987f) and FOlSSNER et al. (l987a; "Versuchsflache G"). Further records: leaf litter ofa
beech forest in Germany (BONKOWSKI 1996); soil sample from Kenya (FOlSSNER 1987f).
Feeds on bacteria and heterotrophic flagellates. Biomass of 106 individuals about 30 mg
(FOlSSNER 1987a).

Gonostomumgonostomoida (HEMBERGER, 1985) comb. nov. (Fig. 124a-f,

Table 23)

1982 Trachelochaeta gonoslomoida HEMBERGER, Dissertation, p 45 (Fig. 124a, c-t).

 Gonostomum 393

1985 Trachelochaeta gonostomoida HEMBERGER, Arch. Protistenk., 130: 400 (1 slide of type specimens is
deposited in the Institut fUr landwirtschaftliche Zoologie of the University of Bonn, Germany; Fig.
124a).

T a x 0 nom y: The shape of the adoral zone of membranelles, the short undulating
membranes, and the dorsal infraciliature requires classification in Gonostomum. The
main difference to the other Gonostomum species is the additional cirral row left of the
right marginal row. Morphogenetic stages reveal that this row and its primordium is ho-
mologous to primordium VI of Gonostomum or other typical oxytrichids. This is mainly
indicated by the two anterior-most cirri of this row, which are not involved in primordia
formation during morphogenesis, identically to the migratory cirri (VII3 and VII4) of the
other oxytrichids (Fig. 124e). The origin and variability of the transverse cirri is some-
what unclear. HEMBERGER (1982) wrote that during morphogenesis one or two cirri are
differentiated at the posterior end of the frontoventral row (Fig. 124t). However, the
specimens shown in Figures 124a, d have three enlarged cirri, suggesting that probably at
least one, but possibly even two parental transverse cirri are retained after division. This
hypothesis, however, would require a rather variable number of 1-3 transverse cirri
within a population because no parental transverse cirri are present in the proter. Such a
variability is not described by HEMBERGER (1982), indicating a possibly different mode of
morphogenesis. Redescription necessary.

M 0 r p hoi 0 g y and b i 0 log y: In life (?) 110-200 x 30-50 J.1m (that is, distinctly
larger than the rather similar G. affine!). Lanceolate. Macronuclear nodules ellipsoidal.
Two micronuclei. Adoral zone of membranelIes about 40 % of body length. Bases of the
frontal-ventral-transverse cirri, except that of frontoventral row, slightly enlarged. All
cirri about 25 J.1m long. Usually 9, sometimes 10 or 11 enlarged cirri in frontal area; HEM-
BERGER (1985) wrote, obviously erroneously, that usually 10 cirri are present. Frontoven-
tral row with 17 or 18 cirri, rarely specimens with 2 rows occur (Fig. 124c). Transverse
cirri slightly enlarged. Dorsal cilia 5 J.1m, posteriorly about 9 J.1m long.

o c cur r e n c e and e col 0 g y: As yet found only at the very extraordinary type
location: an infusion of excrement of the common land snail, Deroceras reticulatum
(MOLLER) collected near Bonn, Germany (HEMBERGER 1982, 1985, KERNEY et al. 1983).
Biomass of 106 individuals about 204 mg (ForSSNER 1987a).

Species indeterminata

Gonostomum parvum LEPSI, 1947, Bull. Sect. scient. Acad. roum., 29: 670 (Fig. 228h).
Remarks: Possibly identical with G. affine. In life 60-80 J.1m long, cytoplasm colourless.
Two macronuclear nodules. Transverse cirri lacking. Ponds in Romania. GELLERT (1957)
recorded it from the soil of a woodland in Hungary.
 394 SYSTEMATIC SECTION

o
a o
o
.-

Q o
o o
d

Fig. 124a-f Gonostomum gonostomoida (a, c-f, from HEMBERGER 1982, 1985. a-f, protargol impregnation).
a) Ventral infraciliature and nuclear apparatus, 175 )lm. b) Same as (a), however, cirri originating from same
primordium are connected by a broken line. The cirri between the two arrowheads are lacking in the very simi-
lar, but smaller Gonostomum affine (cp. with Fig. 117r). Arrows mark "postoral" ventral cirri. c) Specimen
with two ventral rows. d-1) Ventral infraciliature of middle and late morphogenetic stages. Old structures
white, new black. CC = caudal cirri, p = paroral; p 392.
 Gonostomum 395

Insufficient redescriptions

Gonostomum affine STEIN - RosA, 1957, Pi'irodov. Sb. ostrav. Kraje, 18: 45 (Fig. 233b).
Remarks: Soil of a woodland in Czechoslovakia.

Gonostomum affine (STEIN) KAHr.. - DEUIEZ & CHARDEl, 1970, AnnIs Speleol., 25: 135
(Fig. 228d). Remarks: Especially the very long caudal cirri indicate a misidentification.
Limnetic in a Belgian cave.

Gonostomum sp. - CHAUDHURI, 1929, AnnIs Protist., 2: 54, 57 (Fig. 231b, c). Remarks:
Soils in India.

Gonostomum sp. - MOTE, 1954, Proc. Iowa Acad. Sci., 61: 578, 588 (Fig. 2250). Re-
marks: Neither the shape of the adoral zone of membranelles nor the arrangement of the
frontal, ventral, and transverse cirri indicate that it is a Gonostomum. Soil of virgin prai-
rie in Iowa, USA.

Gonostomum sp. - WALKER & HOFFMAN, 1985, Cytobios, 44: 153. Remarks: WALKER &
HOFFMAN (1985) examined the encystment of an unidentified species of Gonostomum by
transmission electron microscopy. Unfortunately they gave no drawing or description of
the infraciliature. They found that the cyst of Gonostomum sp. is clearly of the
kinetosome-resorbing (KR) type and stated that the results of their investigation further
reinforces the SMAIL & LYNN (1985) classification. Accordingly, Gonostomum is a sti-
chotrichine and its encystment varies slightly but significantly from that described for
several sporadotrichine hypotrichs. This classification is not in accordance with the re-
sults obtained by the morphogenetic studies on G. affine and G. strenuum, which indicate
a close relationship to typical oxytrichids such as Urosoma and Tachysoma (HEMBERGER
1982, SONG 1990).

Gonostomun stenuum (ENGELMANN, 1862) STERK!, 1878 - MAoRAZo-GARIBAY & LQpEZ-

OCHOTERENA, 1973, Revta Soc. mex. Hist. nat., 34: 66 (Fig. 241e). Remarks: Incorrect
subsequent spellings of genus and species name. The low number of cirri in the anterior
half indicates that it is not a Gonostomum of the strenuum type. Found in the San Anton
waterfall in Morelos State, Mexico.

Gonyostomum semen (EHRBG.) - SCHMIDT, 1916, Verh. naturh. Ver. preuss. Rheinl., 72:
93. Remarks: Very likely erroneously, SCHMIDT (1916) mentioned this phytomastigopho-
rean taxon in the Oxytrichidae of his faunal list from the Rhineland and Westphalia, Ger-
many.

Oxytricha (Plagiotricha, KENr.) strenua ENGELM. - KOFFMAN, 1926, Acta zool., 7: 312
(Fig. 232p). Remarks: Soil near Stockholm, Sweden.
396 SYSTEMATIC SECTION

Urosoma KOWALEWSKIEGO, 1882

1882 Urosoma KOWALEWSKlEGO, Pam. flzyogr., 2: 406 - Type (by monotypy): Urosoma cienkowskii Kow.
ALEWSKlEGO, 1882 (= junior synonym of Oxytricha caudata EHRENBERG, 1833).
1883 Urosoma KOWALEWSKI, BioI. Zbl., 3: 240 (German translation of Polish original description by August
WRZESNIOWSKI; without illustrations).
1889 Urosoma KOWALEWSKY 1882 - BOTsCHLI, Protozoa, p 1748.
1932 Urosoma KOWALEWSKI, 1882 - KAHL, Tierwelt Dtl., 25: 606 (see nomenclature).
1982 Urosoma KOWALEWSKI, 1882 - FOISSNER, Arch. Protistenk., 126: 109.
1986 Urosoma KOWALEWSKI, 1882 - DRAGESCO & DRAGESCO-KERNEIS, Faune tropicale, 26: 457.
1987 Urosoma - FOISSNER, Jber. Haus Nat. Salzburg, 10: 64.
1997 Urosoma KOWALEWSKlEGO, 1882 - BERGER & FolSsNER, Arch. Protistenk., 148: 148.

C h a r act e r i sat ion: Adoral zone of membranelles and undulating membranes in

Gonostomum pattern. Frontoventral cirri arranged in a row with anterior cirrus (liIl2)
usually slightly displaced to left. Postoral ventral cirri in dense cluster behind buccal ver-
tex or absent. Usually 2 pretransverse ventral and 5 transverse cirri. One right and 1 left
row of marginal cirri. Four dorsal kineties. Caudal cirri present. Primary primordia. Dor-
sal morphogenesis in Urosomoida pattern.

Add i t ion a I c h a r act e r s: Body flexible, but not distinctly contractile, flattened
about 2: 1. Anterior end rounded Two, 4 (u. acuminata), or about 8 (u. octonucleata, U.
polynucleata) macronuclear nodules slightly left of median. Contractile vacuole slightly
to distinctly in front of mid-body. Pellicle and cytoplasm colourless. Close underneath
pellicle in most species (u. acuminata, U. caudata, U. emarginata, U. gigantea, U. kar-
ini, U. octonucleata, U. polynucleata) numerous, about 2-3 /lm sized, ellipsoidal, colour-
less structures, very likely mitochondria. Some species with tiny cortical granules. Adoral
zone of membranelies 18 % (u. polynucleata) to 30 % (u. salmastra) of body length and
usually clearly arranged on left body margin, similarly to in Gonostomum (Fig. 125i,
127g, 128t). Buccal area flat and small. Frontal cirri not to slightly enlarged. Cirrus III12
slightly enlarged in some species (u. acuminata, U. karini, U. polynucleata, U. similis).
Buccal cirrus often near anterior end of anterior undulating membrane. Number of right
marginal cirri usually slightly higher than number of left marginal cirri. Distance between
marginal cirri posteriorly often wider than anteriorly. Dorsal cilia 3-4/lm long. Very
likely all species with one caudal cirrus each on dorsal kineties 1,2, and 3. Common, but
usually not very abundant in freshwater and soil.

Nom e n c I a t u rea n d t a x 0 nom y: KARL (1932) classified Urosoma as subge-

nus of Oxytricha; the correct name in his revision is thus Oxytricha (Urosoma). Incorrect
spellings: Urosomas (?; EDDY 1925, P 85); Urozoma cienkowskii KOWALEWSKI, 1882
(DETCHEVA 1992, P 109; RUSSEV et al. 1984).
The characterisation is according to BERGER & ForSSNER (1997); see chapter 2 in the
general section for detailed description of patterns mentioned. KOWALEWSKlEGO (1882)
 Urosoma 397

and KAHL (1932) separated Urosoma from other oxytrichids mainly by the tapered or tail-
like posterior end of the cell. BORROR (1972a) included it in Oxytricha, whereas STILLER
(1974b), CORLISS (1979), CURDS et al. (1983), and TUFFRAU (1987) considered Urosoma
as valid. However, only ForssNER (1982, 1983a, 1987b), DRAGESCO & DRAGESCo-KERNErs
(1986), and BERGER & ForssNER (1987a, 1989a) described some species in detail. Most
species now assigned to Urosoma are in fact rather slender and posteriorly more or less
distinctly narrowed. However, the only autapomorphy of Urosoma is the peculiar ar-
rangement of cirrus 11112, namely in front of the level of cirrus V1I4, that is, the fronto-
ventral cirri form a row, and not a V-shaped pattern as in most other oxytrichids (Fig.
19a, 125i; BERGER & ForSSNER 1997). Such a row also occurs in some species originally
assigned to Perisincirra (P. similis; Fig. 131d, t) and Hemisincirra (H. polynucleata; Fig.
130f, h). This character and the agreements in the oral apparatus and dorsal infraciliature
strongly suggest that these species belong to Urosoma, although the frontal-ventral-
transverse cirri behind the adoral zone are partially or completely reduced. Very likely,
Perisincirra simi/is and Hemisincirra polynucleata form a distinct clade in Urosoma.
Urosoma has several synapomorphies with Gonostomum, supporting the sister group
relationship proposed by FOISSNER (1983a; Fig. 25a). WIRNSBERGER et al. (1986) ignored
the primary primordia (Fig. 127m; character 16 in Fig. 25a) and the distinctive oral appa-
ratus (Fig. 19h; characters 1,3 in Fig. 25a), but used the decreased number of pretrans-
verse ventral and transverse cirri as autapomorphies. Thus, WIRNSBERGER et al. (1986) ar-
rived at Gonostomum and Urosomoida as sister groups.
According to JANKOWSKI (1979, P 68), Tricholeptus FROMENTEL, 1875 (p 177) is pos-
sible a synonym of Urosoma. However, since the type species, Tricholeptus aculeatus, is
insufficiently described (FROMENTEL 1876, P 294; Planche XX, Fig. 7) and since this
taxon was never accepted by later authors (for example, KAHL 1932) it is considered as
nomen oblitum (CoRLISS 1979, p 208).

Key to species

1 2 macronuclear nodules ............................................... 4

More than 2 macronuclear nodules (Fig. 128b, 129a, 130a) .................. 2
2 Usually 4 macronuclear nodules (Fig. 128a-o) .............. U. acuminata (p 414)
Usually 8 (6-9) macronuclear nodules ................................... 3
3 3 postoral ventral cirri; transverse cirri subterminal (Fig. 129a-t) ................ .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. U. octonucleata (p 417)
Only 1 postoral ventral cirrus; transverse cirri terminal and inconspicuous (Fig.
130a-i) .............................................. U. polynucleata (p 419)
4 (1) Posterior end rounded, tapering, or tail-like ............................ 5
Right margin posteriorly distinctly emarginated (Fig. 127a-c) ................... .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . U. emarginata (p 409)
5 Posterior end rounded (for example, Fig. 132a) ............................ 6
398 SYSTEMATIC SECTION

Posterior end tapering or tail-like (for example, Fig. 125g, 126a, c, 131a) ....... 8
6 Length in life 170-230 ~m (Fig. 134a-w) ................... U. gigantea (p 425)
Length in life 50-160 ~m ............................................. 7
7 Cortical granules lacking; undulating membranes arranged one after another, endoral
much shorter than paroral (Fig. 133a-c) .................... U. salmastra (p 424)
Cortical granules present; undulating membranes overlapping, of about equal length
(Fig. 132a-e) ............................................... U. karini (p 422)
8 (5) Posterior body part tail-like (Fig. 125a-q) ................. U. caudata (p 398)
Posterior body part tapering (for example, Fig. 126c, 131a) .................. 9
9 3 postoral ventral cirri; 5 subterminaIIy arranged, distinctly enlarged transverse cirri
(Fig. 126a, c-e) .................................... U. macrostyla (p 404)
Postoral ventral cirri absent; 2-3 inconspicuous, terminal transverse cirri (Fig.
131a-g) .............................................. U. similis (p 419)

Urosoma caudata (EHRENBERG, 1833) comb. nov. (Fig. 125a-u, Tables

24,41)

1833 Oxytricha caudata EHRENBERG, Abh. preuss. Akad. Wiss., year 1833: 278, 279.
1838 Oxytricha caudata - EHRENBERG, Infusionsthierchen, p 365, in part: Tafel XL, Fig. XIt- 3; not Tafel XL,
Fig. X4.6 (Fig. 125a-c).
1882 Urosoma cienkowskii KOWALEWSKIEGO, Pam. fizyogr., 2: 406 (Fig. 125r; new synonym).
1883 Urosoma cienkowskii KOWALEWSKI, BioI. Zbl., 3: 240 (Gennan translation of the Polish original de-
scription by August WRZESNIOWSKI).
1887 Oxytricha caudata STOKES, Ann. Mag. nat. Hist., 20: 112 (Fig. 125f; new synonym; homonym).
1888 Oxytricha caudata, STOKES - STOKES, 1. Trenton nat. Hist. Soc., I: 291 (Fig. 125t).
1930 Oxytricha caudata STOKES - WANG, Contr. bioI. Lab. Sci. Soc. China, 6: 14 (Fig. 125d).
1932 Urosoma (Oxytricha) caudata STOKES, 1887 - KAlIL, Tierwelt Dtl., 25: 607.
1932 Urosoma cienkowskii KOWALEWSKI, 1882 -KAHL, Tierwelt Dtl., 25: 607 (Fig. 125e).
1935 Urosoma cinekowski KOWALEWSKI 1882 - WANG & NIE, Sinensia, Shanghai, 6: 500 (Fig. 125s; incor-
rect subsequent spelling).
1956 Urosoma cienkowskyi KOWALEWSKI - GRAAF & MEUER, Micro-organismen, pili, 112 (Fig. 125t; in-
correct subsequent spelling).
1963 Urosoma caudata (STOKES) - LuNDIN & WEST, Free-living protozoa, p 67 (Fig. 125p).
1967 Urosoma caudata (STOKES) 1887 - BIERNAcKA, Wiss. Z. Ernst Moritz Arndt-Univ. Greifswald, 16: 247
(Fig. 125u).
1972 Urosoma cienkowskii KOWALEWSKI - BlcK, Ciliated protozoa, p 19 (Fig. 125q).
1972 Trichotaxis caudata (EHRENBERG, 1838) - BORROR, 1. Protozool., 19: 11 (incorrect subsequent spelling
of Trichototaxis).

Fig. 125a-k Urosoma caudata (a-c, after EHRENBERG 1838; d, from WANG 1930; e, from KAlIL 1932; f, from ~
STOKES 1887b; g-k, from FOISSNER 1982. a-h, k, from life; i, j, protargol impregnation). a-c) Various views,
170-210 )lm. d-g) Ventral views, d = 240 )lm, e = 250 )lm, f = 220 )lm, g = 225 )lm. h) Mitochondria(?) close
beneath the pellicle. i, j) Ventral and dorsal infraciliature of same specimen, 105 )lm. Arrow marks frontoven-
traI cirri, which are arranged in a row and not in a V-shaped pattern as in most other oxytrichids. k) Two speci-
mens creeping on a soil particle.
Urosoma 399
 400 SYSTEMATIC SECTION

I
. \

.~ !'
~

.. ,. .
: t'r;

.'i'
,
r: /

.'.
>...:~
ot! ) \
I
I \
I \.
, I
I \.
\.
~
tJ
\.
~ \.

•• / "\.
II.
/
-'<!II.'
,I
~
j \.
...
!t~~; \.
- / \
/
~
J)

m
q

Fig. 1251-q Urosoma caudata (h>, from FOISSNER 1984; p, from LUNDIN & WEST 1963; q, from BICK 1972a. I,
m, p, q, from life; n, 0, protargol impregnation). I, p, q) Ventral views, I =200 J-lm, p = size not indicated, q =
270 J-lm. m) Dorsal view ofa very slender, worm-like specimen, 200 J-lm. n, 0) Ventral and dorsal infraciliature
of same specimen, 195 J-lm. Arrow marks inconspicuous caudal cirri at tip of tail; p 398.

1982 Urosoma cienkowskii KOWALEWSKI, 1882 - FOISSNER, Arch. Protistenk., 126: 112 (Fig. 125g--k; authori-
tative redescription; slides of neotype material are deposited in the OberOsterreichische Landesmuseum
in Linz, Upper Austria).
1984 Urosoma cienkowskii KOWALEWSKI, 1882 - FOISSNER, Stapfia, 12: . 121 (Fig. 1251-0; 1 slide of protargoI
impregnated specimens is deposited in the Oberosterreichische Landesmuseum in Linz, Upper Austria).

Nom e n cIa t u rea n d t a x 0 nom y: KAliL (1932) classified Urosoma as subge-

nus of Oxytricha; the correct names in his revision are thus Oxytricha (Urosoma) cau-
data STOKES and Oxytricha (Urosoma) cienkowskii. Incorrect subsequent spellings: Uro-
 Urosoma 401

soma caudatum STOKES (WEBB 1961; P 140); Urosoma cienckowski Kow. (COPPA 1921,
P 188); Urosoma cienkowsky KOWALEWSKI, 1882 (LUZZATTI 1938, P 102).
EHRENBERG (1838) emphasised that Oxytricha caudata is very similar to O. rubra
(now Pseudokeronopsis rubra), except that 0. caudata is more pointed posteriorly, and
in that he observed cirri not in the median of the cell (that is, he observed no mid-ventral
cirri!), but near the oral apparatus and also 5 cirri near the posterior end. This strongly in-
dicates that EHRENBERG observed an oxytrichid and not an urostylid as suggested by KENT
(1882) and BOTsCHLI (1889). Thus, I classify O. caudata EHRENBERG as senior synonym of
Urosoma cienkowskii KOWALEWSKIEGO and Oxytricha caudata STOKES. The latter is ajun-
ior primary homonym, which, however, need not be replaced because of synonymy (Arti-
cle 60a OfICZN 1985).
KAHL (1932) separated U. caudata (STOKES) and U. cienkowskii by the body shape
and the number of postoral and pretransverse ventral cirri: U. caudata anteriorly broad
and 5 cirri (Fig. 125t), U. cienkowskii parallel sided and 8 cirri (Fig. 125r). WANG & NIE
(1935) confIrmed the increased number of cirri between buccal vertex and transverse cirri
in U. cienkowskii (Fig. 125s), a phenomenon which can also be observed in other oxytri-
chid species. Thus, I preliminarily agree with FOISSNER (1982), who considered these dif-
ferences to be a mainly intraspecifIc variability. However, there is no doubt that U. cien-
kowskii has to be separated from U. caudata if the increased number of cirri is substanti-
ated by protargol impregnation and morphometric analysis.
Without foundation, BORROR (1972a) transferred Oxytricha caudata EHRENBERG to
Trichototaxis and synonymised Oxytricha caudata STOKES with Urosoma acuminata,
which has, however, 4 macronuclear nodules. Some illustrations of the present species
are rather poor; in spite of this, the identifications can be accepted (for example, Fig.
125p). The redescriptions by CONN (1905; Fig. 225i), DUJARDIN (1841), DUMAS (1929,
1930, Fig. 236a, b, r, 239g), and FROMENTEL (1876) are insufficient. "Oxytricha caudata
EHR." sensu CLAPAREDE & LACHMANN (1859, Planche V, Fig. 7) is an Uroleptus because
mid-ventral cirri are present.

M 0 r p hoI 0 g y and b i 0 log y: In life 150-300 x 20-50 ~m; 170-210 ~m (EH-

RENBERG 1838), 240 ~m (KoWALEWSKIEGO 1882), 210-255 ~m (STOKES 1887b), 150 to
300 ~m (KAHL 1932), 250-300 x 40-50 ~m (WANG & NIE 1935), 180-270 times
20-35 ~m (FOISSNER 1982; terrestrial population), 150-250 ~m (FOISSNER 1984; freshwa-
ter population); ratio oflength to width usually 7: I, that is, slender-elongate, anterior end
rounded, posterior end tail-like; some populations almost vermiform (Fig. 125m). Macro-
nuclear nodules in life about 14-17 x 6-1 0 ~m, after protargol impregnation 8-14 times
4-7 ~m (n = 17; FOISSNER 1982, 1984).2-4, in life about 4 ~m, after protargol impregna-
tion 2.5-4.0 x 2-3 ~m sized micronuclei. Contractile vacuole with short and inconspicu-
ous collecting canals. Mitochondria give cells a slightly brownish shimmer at low magni-
fIcation (Fig. 125h; FOISSNER 1982). Around bases of cirri and dorsal cilia small (about
0.5 ~m), colourless cortical granules (Fig. 125h). Cytoplasm of anterior cell part some-
times with reddish crystals (FOISSNER 1982), according to KOWALEWSKIEGO (1882) whole
 402 SYSTEMATIC SECTION

cytoplasm appears reddish. Ac-

cording to KAliL (1932; sapro-
pelic populations), WANG & NIE
(1935), and RECK (1987) symbi-
otic algae present (confused
with ingested algae? other spe-
cies?). Moves moderately rap-
idly,jerks hastily to and fro.
Adoral zone of membra-
nelles 15-20 % of body length.
Usually 3 postoral ventral cirri
and 1 pretransverse ventral cir-
rus (Fig. 125i, n); according to
KOWALEWSKIEGO (1882) and
WANG & NIE (1935), however,
6 (5 plus 1) postoral and 2 pre-
transverse ventral cirri (Fig.
u 125r, s; see taxonomy). Trans-
verse cirri about 25 J.lm long, at
anterior end of tail-like process.
Left marginal row terminates
about 20 J.lm in front of tip of
tail; cirri in life 10-15 J.lm long.
Dorsal kinety 1 distinctly short-
ened anteriorly. Distance be-
tween basal body pairs widens
posteriad. Caudal cirri difficult
to discern even after protargol
impregnation; very likely three
Fig.12Sr-u Urosoma caudata (r, after KOWALEWSKlEGO 1882; s,
rather fine, about 15 J.lm long
after WANG & NIE 1935; t, from GRAAF & MEUER 1956; U, from
BIERNACKA 1967. r-u, from life). r-u) Ventral views, r = 240 Jlm,
cirri. Morphogenesis of Ura-
S = 285 Jlm, t = size not indicated, u = >200 Jlm; p 398. soma caudata commences with
formation of an oral primor-
dium left of and beside postoral
ventral cirri (FOISSNER 1984), identically to in Urosoma emarginata and Urosoma poly-
nucleata.

o c cur r e n c e and e colo g y: Common, but usually with low abundance in detri-
tus and aufwuchs (rarely in the pelagial) of moderately to distinctly polluted lentic and
lotic waters; also common in terrestrial habitats. Recorded from Eurasia, South America,
and Africa. After KAliL (1932) one of the most common oxytrichids in saprobic habitats;
 Urosoma 403

according to my experience, however, much more seldom than, for example, Tachysoma
pellionellum, Stylonychia pustu/ata, or Oxytricha spp.
Locus classicus is a freshwater habitat near Berlin, Germany (EHRENBERG 1833,
1838). Locus classicus of the synonym, Urosoma cienkowskii, is the Weichsel River near
Warsaw, Poland (KOWALEWSKlEGO 1882, KOWALEWSKI 1883). Locus classicus of the syno-
nym, Oxytricha caudata STOKES, is a Lemna pond in the USA, probably near Trenton,
New Jersey, where STOKES lived and worked. Further records substantiated by illustra-
tions: arable land and bottom-land in the Tullnerfeld, Lower Austria (FOISSNER 1982; see
FOISSNER et al. 1985 for detailed description of sites); very abundant in cyanobacteria
slimes (Spirolina jennen) in a hypertrophic pond in Salzburg, Austria (FOISSNER 1984);
the Netherlands (GRAAF 1956, 1957, GRAAF & MEllER 1956, GROOT & GRAAF 1960); in
great abundance together with species of the Sty/onychia mytilus-complex in standing
water collected more than a month previously from Lake Ho Hu in Nanking, China
(WANG 1930, WANG&NIE 1935).
Records from freshwater habitats not substantlated by illustrations: brook in Upper
Austria (BLATTERER 1994); Maritza River, Bulgaria (DETCHEVA 1981); submerged and wet
mosses and Turlek River in Czechoslovakia (SCHMARDA 1846, TIRJAKovA 1993, TIRJAK-
ovA & MAns 1987a); sediment of Esthwaite, one of the most eutrophic lakes in the Eng-
lish Lake District (FINLAY 1982, WEBB 1961); pronounced summer maximum in a
eutrophic water body in England (FINLAY et al. 1979); peat-bogs in France (GROLIERE
1977,ORMANCY 1852); betamesosaprobic rivers in Germany (BUCK 1959, 1961, MAUCH
1976); mesotrophic lake in Germany (MOCKE 1979); about 20 indo cm·2 in the pseudoperi-
phyton of a eutrophic pond in Bonn, Germany (WILBERT 1969); eutrophic lake in Ger-
many with a maximum of 2937 ind.l- I in the autumn plankton (RECK 1987; with symbi-
otic algae, other species?); sediment of Hamburg Harbour, Germany (BARTSCH &
HARTWIG 1984); together with alphamesosaprobic species in the sediment of sewage oxi-
dation ponds of Munich, Gennany (KAUFMANN 1958); detritus drifts in Lake Balaton,
Hungary (GELLERT & TAMAS 1958); 4 indo cm-2 in an alphamesosaprobic section of the Po
River, Italy (MAooNI 1979); well in Italy (GRISPINI 1938; further records from Italy: CAN-
ELLA 1954, COPPA 1921); sediment of Latvian rivers (LIEPA 1986, 1990, VEYLANDE & LI-
YEPA 1985); Spain (FINLAY et al. 1993, MARGALEF LOPEZ 1945); in bogs near Berne (Swit-
zerland) from April to December (PERTY 1852b); eutrophic ponds in Yugoslavia
(SCHMARDA 1847); rivers, reservoirs, and other habitats in USSR (ALIEv 1982a,
KRAvCHENKO 1969, MOVCHAN & PROTASOV 1986, WEISSE 1848a, b, c); Lake Dong Hu in
Wuhan, China (SHEN 1980); Tibetan Plateau (WANG 1977); slightly polluted third order
stream in a mid-Atlantic State, USA (NIEDERLEHNER et al. 1990); Colorado, USA (BEARD-
SLEY 1902, HAMILTON 1943); in bogs with up to 14.2 mg I-I H2S in Iowa, USA (HEMP-
STEAD & JAHN 1940, SHAWHAN et al. 1947); at 15-17 °C in a slow-moving stream in Ken-
tucky, USA, the bed littered with leaves, rocks, and fallen tree limbs (GITILESON & FER-
GUSON 1971); Louisiana, USA (BAMFORTH 1963); Massachusetts, USA (COLE 1853); at
20°C and pH 6.0-6.5 in a bog and in a river in Michigan, USA (CAIRNS & YONGUE
1966); Cape Fear River in North Carolina, USA (CAIRNS & YONGUE 1973a); among
404 SYSTEMATIC SECTION

algae-covered poplar roots and once in black sediment of Mirror Lake in Ohio, USA
(STEHLE 1920); Conestoga drainage basin in Pennsylvania, USA (CAIRNS 1965a); Savan-
nah River, USA (PATRICK et al. 1967); Mexico (L6PEZ-OCHOTERENA & ROURE-CANE 1970,
MAnRAZO-GARIBAY & LOPEZ-OCHOTERENO 1973, PEREZ REYES & SALAS GOMEZ 1961); Peru
(EscoMEL 1929, only 70 pm long; SARMIENTO & GUERRA 1960); irrigation ditches with
diatoms and cyanobacteria in Egypt (ScHMARDA 1854).
Records from terrestrial habitats not substantiated by illustrations: sub-alpine grass-
land in Austria (FOISSNER et al. 1990); Czechoslovakia (ROSA 1957a); Italy (LUZZATIl
1938); Abaco Island, Bahamas (CAIRNS & RUTIfVEN 1972); cedar glade in Tennessee
(MARTIN & SHARP 1983); soil near Lake Baringo, Kenya (W. FOISSNER, pers. comm.).
Records from saline habitats: littoral detritus of the Hiddensee Island and saline res-
ervoir near the Baltic (BIERNACKA 1967, Fig. 125u; CZAPIK 1962); saline lake in Romania
(TucoLESco 1962b); low salinity lagoons (about 1 %) in the Caspian Sea (AGAMALIYEV
1974); sea water (EICHWALD 1849; uncertain).
Feeds on bacteria, cyanobacteria (Oscillatoria), diatoms, ciliates, heterotrophic flag-
ellates, hyphae and/or spores of fungi and yeast (FINLAyet al. 1993, FOISSNER 1987a,
WANG 1930, WANG & NIE 1935). Biomass of 106 individuals according to FOISSNER
(l987a) 34 mg, according to NESTERENKO & KovALCHUK (1991; incorrectly spelled as
"Urozoma cienkowekii KOWAL.") 180 mg (other species?). Autecological data, see Table
41. RUTIfVEN (1972; station 6) and CAIRNS & DICKSON (1972) found U. caudata at follow-
ing heavy metal concentrations: 0.046 mg 1-1 Cu, 0.067 mg 1-1 Zn, 0.036 mg 1-1 Cr,
0.017 mg I-I Pb.

Urosoma macrostyia (WRZESNlOWSKIEGO, 1866) comb. nov. (Fig. 126a-g,

Table 24)

1866 Oxytricha macrostyla WRZESNJOWSKlEGO, Wykaz Szkoty Glownej Warszawskiej, NO.5: 17 (Latin diag-
nosis, without illustration).
1867 Oxytricha macrostyla WRZE5NIOWSKlEGO, Przyczynek do historyi naturalnej wymoczk6w, p 56 (Fig.
126a, b).
1867 Oxytricha macrostyla WRZE5NIOWSKlEGO, Rocmik ces. kr6!. Towarzystwa Naukowego Krakowskiego,
12: 284 (Fig. 126a, b).
1870 Oxytricha macrostyla WRZESNIOWSKI, Z. wiss. Zoo!., 20: 474.
1932 Urosoma (Oxytricha) macrostyla WRZESNIOWSKI, 1870 - KAHL, Tierwelt Ot1., 25: 607, in part: Fig.
11324; not Fig. 11322• 23•
1972 Urosoma acuta ORAGESCO, Annis Fac. Sci. Univ. fed. Cameroun, 11: 86 (Fig. 126f, g; new synonym).
1986 Urosoma acuta ORAGESCO, 1972 - ORAGESCO & ORAGESCO-KERNEIS, Faune tropica1e, 26: 457.
1986 Urosoma ambigua ORAGESCO & ORAGESco-KERNEIS, Faune tropicale, 26: 459 (Fig. 126c-e; new syno-
nym).

Nom e n cia t u rea n d t a x 0 nom y: Incorrect subsequent spelling: Urosoma

macrostya (HoRvArn 1950, p 149). KARL (1932) classified Urosoma as subgenus ofOxy-
tricha; the correct name in his revision is thus Oxytricha (Urosoma) macrostyla, that is,
 Urosoma 405

Table 24 Morphometric data of Urosoma acuminata (acl, from FOISSNER 1982; ac2, from BERGER & FOISSNER
1987a), U. caudata (cal, from FOISSNER 1982; ca2, from FOISSNER (984), U. emarginata (eml, from FOISSNER
1982; em2, from FOISSNER 1983a), U. gigantea (gig, from BERGER & FOISSNER 1987a), U. karini (kar, from
FOISSNER 1987b), U. macrostyla (mac, from DRAGESCO & DRAGESCO-KERNEIS (986), U. octonucleata (oct, from
BERGER & FOISSNER (989), U. polynucleata (pol, from FOISSNER (984), U. salmastra (sal, from DRAGESCO &
DRAGESCO-KERNEIS (986), and U. similis (sim, from FOISSNER (982). All data are based on protargol-
impregnated specimens. All measurements in micrometres. ? = sample size not indicated; if only I value is
known it is listed in the column mean, if 2 values are available they are listed as Min and Max. CV = coeffi-
cient of variation (in %), Max = maximum value, mean = arithmetic mean, Min = minimum value, n = sample
size, SD = standard deviation

Character Species mean SD CV Min Max n

Body, length acl 113.2 20.4 18.0 80 152 11
ac2 154.5 14.2 9.2 125 180 13
cal 111.3 16.1 14.5 80 132 6
ca2 167.1 19.2 11.5 136 190 11
eml 87.0 6.8 7.8 73 95 10
em2 109.4 13.7 12.5 83 140 25
gig 157.7 12.3 7.8 140 182 25
kar 119.2 18.1 15.2 92 154 10
mac 150.0 130 168 5
oct 141.2 13.5 9.6 105 155 11
pol 143.3 19.8 13.8 112 182 15
sal 73.0 48 101 29
sim 113.3 15.6 13.8 83 140 17
Body, width acl 29.4 3.4 11.5 23 36 '11
ac2 43.8 3.3 7.5 38 49 13
cal 21.3 4.8 22.5 15 29 6
ca2 20.6 1.4 6.9 17 22 11
eml 21.5 1.7 8.1 20 25 10
em2 26.8 1.9 6.9 22 31 25
gig 47.2 3.7 7.8 42 58 25
kar 36.1 4.6 12.7 29 42 10
mac 33.0 24 39 5
oct 25.8 1.9 7.5 22 28 11
pol 21.4 2.1 9.6 18 25 15
sal 21.0 15 30 27
sim 17.1 1.7 9.9 14 20 17
Adoral membrane lies, number acl 23.6 1.1 4.9 22 26 11
ac2 36.0 1.4 3.9 34 38 13
cal 26.5 1.0 3.6 25 28 6
ca2 23.6 1.4 5.8 21 25 II
eml 25.0 0.9 3.6 24 27 10
em2 31.9 1.6 5.1 26 34 25
gig 46.8 2.7 5.7 42 58 25
kar 22.0 1.0 4.5 21 24 10
mac 43.0 38 45 5
oct 25.1 0.7 2.8 24 26 11
pol 21.5 1.4 6.6 19 24 IS
sal 23.0 21 26 17
sim 21.6 0.7 3.1 20 23 17
406 SYSTEMATIC SECTION

Table 24 Continued

Character Species mean SD CV Min Max n

Macronuclear nodules, number' aci 4.0 0 0 4 4 II

ac2 3.9 0.4 9.7 2 4 34
oct 7.6 0.8 10.9 6 8 II
pol 7.7 0.7 9.1 6 9 15
Right marginal row, number of cirri acl 35.3 3.0 8.6 29 40 II
ac2 43.2 2.4 5.4 37 46 13
cal 45.2 6.0 13.4 36 53 6
ca2 54.6 7.3 13.3 3.8 64 II
emi 33.3 1.3 4.0 32 36 10
em2 37.9 2.9 7.6 31 43 25
gig 42.9 2.2 5.1 38 46 25
kar 29.2 2.3 8.0 26 33 10
mac 50.0 40 59 5
oct 37.3 1.8 4.8 35 40 II
pol 33.0 2.0 6.2 30 38 15
sal 26.0 22 30 24
sim 33.6 2.8 8.4 27 37 17
Left marginal row, number of cirri acl 32.4 3.2 9.9 28 38 II
ac2 40.4 1.4 3.4 38 43 13
cal 41.2 4.9 11.8 33 48 6
ca2 51.3 7.4 14.4 38 60 II
emi 29.9 1.3 4.3 28 32 10
em2 36.3 3.2 8.9 27 40 25
gig 34.6 2.2 6.3 30 38 25
kar 30.4 3.4 11.3 26 36 10
mar 50.0 45 55 5
oct 31.1 1.7 5.5 28 34 II
pol 26.9 1.8 6.7 24 30 15
sal 21.0 16 25 20
sim 27.6 2.1 7.8 24 32 17
Pretransverse ventral cirri, number acl 0.5 0.5 109.5 0 I II
ac2 1.9 0.3 14.4 2 13
cal 0.7 0.5 70.7 0 I 6
ca2 0.9 0.3 33.1 0 1 II
eml 1.9 0.5 28.3 3 10
em2 2.0 0 0 2 2 25
gig 1.9 0.3 14.4 1 2 25
kar l.l 0.3 28.7 2 10
mac 2.0 ?
pol 0 0 0 0 0 15
sal 1.0 ?
sim probably absent
Transverse cirri, number acl 4.0 0 0 4 4 11
ac2 4.9 0.3 5.6 4 5 13
cal 4.3 0.5 11.0 4 5 6
ca2 4.9 0.3 6.1 4 5 II
eml 4.1 0.3 7.3 4 5 10
em2 4.9 OJ 5.6 4 5 25
 Urosoma 407

Table 24 Continued

Character Species mean SO CV Min Max n

Transverse cirri, number gig 5.0 0 0 5 5 25
kar 5.0 0 0 5 5 10
mac 5.0 ?
oct2 3.5 0.5 15.1 3 4 11
pol 2.0 0 0 2 2 15
sal 4.0 3 5 ?
sim 2.1 0.2 11.4 2 3 17

I All other species have two macronuclear nodules.

1 Pretransverse ventral cirri included.

Oxytricha macrostyla was never fonnally transferred to Urosoma, which hereby under-
taken.
KAHL (1932) and FOISSNER (1982) synonymised Opisthotricha emarginata STOKES
with Urosoma macrostyla, assuming that WRZESNIOWSKI (1870) had overlooked the con-
spicuous notch at the right posterior margin described in O. emarginata. BORROR (1972a)
synonymised both species with Urosomoida agilis, which has, however, cortical
granules, fewer pretransverse ventral and transverse cirri, and a different arrangement of
frontoventral cirri. I consider both Urosoma macrostyla and U. emarginata as valid spe-
cies. By contrast, U. ambigua is obviously a junior synonym of U. macrostyla, which is
indicated especially by the distinctly enlarged and anteriorly displaced transverse cirri
(cp. Fig. 126a with 126c, e). Urosoma acuta is rather superficially described and only af-
ter protargol impregnation; the body shape and size, the subtenninal transverse cirri, and
many morphometric data agree rather well with U. ambigua and/or U. macrostyla. I have
some doubt that the arrangement and the number of frontoventral and postoral ventral
cirri is correctly illustrated (Fig. 126g). Redescription of U. macrostyla necessary.

M 0 r ph 010 g Y and b i 0 log y: In life about 120 x 30 11m (WRZESNIOWSKI 1870),

according to DRAGESCO & DRAGESCo-KEIlliEIS (1986) 130-170 x 25-40 11m (mean =
150 x 33 11m; n = 5). Elongate, anterior broad, posterior narrowly rounded (Fig. 126a, c,
g). Not distinctly contractile. Macronuclear nodules after protargol impregnation
13 x 16 11m. 1-3 globular micronuclei. Contractile vacuole slightly behind buccal vertex.
Cortical granules lacking (not mentioned by WRZESNIOWSKI 1870 and DRAGESCO &
DRAGESCo-KEllliEIS 1986).
Adoral zone of membranelles about 27 % of body length. Transverse cirri do not
protrude beyond posterior end of cell, distinctly enlarged (species name! Fig. 126a, C, e,
g). Marginal rows tenninate very close to posterior tip (Fig. 126a, d). Four dorsal kineties
(DRAGESCO & DRAGESCo-KEIlliEIS 1986). According to WRZESNIOWSKI (1870), 4-7 dis-
tinctly elongated, rather fme cirri at posterior end (caudal cirri?).
408 SYSTEMATIC SECTION

.--- :.·fJM,..
.
1
,,~ '~~.,

-•
~

........
~
,"
~
~·.·.:~;;i\
~
,, ,•

•
.
.-.-
,,,
....
~
1(.~
~
~
®
.-
,.
,.,.
~

... ,
,,, .
..
~ "
~

,
,. •
f

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,,
.-
.-.-
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.....
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a b c d e 9
Fig. 126a-g Urosoma macrostyla (a, b, after WRZESNIOWSKlEGO 1867a; c--e, from DRAGESCO & DRAGESCo-
KERNElS 1986; f, g, from DRAGESCO 1972b. a, b, f, from life; c--e, g, protargol impregnation). a, c, g) Ventral
views, a = 120 ~m, c = 165 ~m, g = 145 ~m. Note the distinctly enlarged transverse cirri. b) Lateral view of
anterior end. d) Ventral infraciliature, 145 ~m. e) Ventral infraciliature of anterior (above) and posterior (be-
low) portion of cell, bar = 10 ~m. Note prominent transverse cirri. t) Outline in ventral view; p 404.

The following data are from the synonym U. acuta (Fig. 126f, g): after protargol im-
pregnation 120-170 !lm long. Posteriorly tapered. 43-53 adoral membranelles. Six cirri
on frontal area, 5 postoral ventral cirri, 6 transverse cirri (probably 5 transverse cirri and
1 pretransverse ventral cirrus), 42-63 right and 39-62 left marginal cirri.

o c cur r e n c e and e colo g y: Not very common in freshwater habitats; records

from soil not substantiated by illustrations. Locus classicus is a small water body with
Sphagnum and Lemna trisulca in Gr6jec near Warsaw (poland), where WRZESNIOWSKIEGO
(1866) discovered Urosoma macrostyla very abundantly in the surface film ("Kahm-
 Urosoma 409

haut") of a several day old sample together with numerous other species (WRZESNlOWSKI
1870). Locus classicus of the synonym Urosoma acuta is Lake Chad (Chad), where
DRAGESCO (1972) found it very abundantly. Locus classicus of the second synonym, U.
ambigua, is a saline pool near Cotonou, Benin (DRAGESCO & DRAGESco-KERNEIS 1986).
Records not substantiated by illustrations: wells in Czechoslovakia and Italy (VEJD()-
VSKY 1882; GRISPINl 1938, detennination uncertain because 290 /.lm long); agricultural
soils and freshwater in Czechoslovakia (TIRJAKovA 1988, 1992b; possible U. emargi-
nata); caves in France (MoNlEZ 1889); soil infusion with pH 7 in Gennany (DINGFELDER
1962); sandy soil near Tisza River, Hungary (HORVATH 1950); benthal ofa freshwater la-
goon of the Caspian Sea (AGAMALIEV 1973, 1986); periphyton of Lake Dong Hu in China
(SHEN 1980); soil in Costa Rica and Brazil (FOISSNER 1995b, 1997b). Feeds on bacteria
and algae (HORvATH 1950).

Urosoma emarginata (STOKES, 1885) comb. nov. (Fig. 127a-y, Tables 3, 24)

1885 Opisthotricha emarginata STOKES, Ann. Mag. nat. Hist., 15: 445 (Fig. 127a).
1888 Opisthotricha emarginata, STOKES - STOKES, J. Trenton nat. Hist. Soc., 1: 289 (Fig. 127a).
1932 Urosoma (Oxytricha) macrostyla WRZESNIOWSKI, 1870 - KAHL, Tierwelt Ott., 25: 607, in part: Fig.
11322,23, not Fig. 11324 (Fig. 127b; see taxonomy).
1982 Urosoma macrostyla (WRZESNIOWSKI, 1870) - FOISSNER, Arch. Protistenk., 126: 114 (Fig. 127c-h; see
taxonomy; 1 slide of protargol-impregnated specimens is deposited in the OberOsterreichische Landes-
museum in Linz, Upper Austria).
1983 Urosoma macrostyla (WRZESNIOWSKI, 1870) - FOISSNER, Arch. Protistenk., 127: 413 (Fig. 127i-y; see
taxonomy).

T a x 0 nom y: KAlIL (1932) and FOISSNER (1982) synonymised Urosoma emarginata

(STOKES) with U. macrostyla (WRZESNlOWSKIEGO), assuming that WRZESNlOWSKIEGO (1866)
had overlooked the conspicuous notch at the right posterior margin. I consider both spe-
cies as valid. Furthennore, U. macrostyla has rather prominent, distinctly anteriorly dis-
placed transverse cirri. BORROR (1972a) synonymised both species with Urosomoida agi-
lis (ENGELMANN), which has, however, cortical granules, fewer pretransverse ventral and
transverse cirri, and a different arrangement offrontoventral cirri (Fig. 110c).

M 0 r p hoI 0 g y and b i 0 log y: In life 125-145 /.lm long (STOKES 1885b), accord-
ing to FOISSNER (1982) 150-190 x 40-60 /.lm. Elongate, anterior end rounded, posterior
end narrowly rounded or even pointed, right posterior margin conspicuously emargi-
nated; not distinctly contractile. Usually 2 (rather seldom 4), after protargol impregnation
8-24 x 5-7 /.lm sized macronuclear nodules. 2-4, in life about 9 x 3 /.lm, refractile micro-
nuclei. Contractile vacuole with long collecting canals, anterior one often with a small
vesicle at level of cytostome. Cortical granules absent. Posterior third of cell usually
packed with 1-2 /.lm sized, colourless globules; cells thus appear dark at low magnifica-
tion and bright field illumination. Food vacuoles about 7 /.lm across. Movement rapid and
erratic.
410 SYSTEMATIC SECTION

e f 9 h

· 'iJ
~
o "" ~ ~
• • ""
·••
• o d
"
"" nfi
•, " H ,
"" •,

I
• "" ••, ",
•• " ,
•,
()

-OP"¥;:' ""
-~:~; ", "" ",
" ",, "
"
()

"" "" ""

,, ,,
'!;:;:

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(J

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,
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p
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.-
 Urosoma 411

Adoral zone of membranelles about 24 % of body length. Right frontal cirrus often
slightly thicker than other frontal cirri. Transverse cirri do not protrude beyond posterior
end of cell and are not distinctly larger than marginal cirri. Marginal cirri in life about
13 J.lm long, right row terminates at level of transverse cirri, left at tip of posterior end.
Dorsal kinety 1 anteriorly, kinety 4 posteriorly distinctly shortened.
Resting cysts smooth, colourless, in life about 45 J.lm across, surrounded by an about
20 J.lm thick mucous layer.
M 0 r p hog e n e sis is described in great detail by ForSSNER (1983a; Fig. 127i-y).
It commences left of the postoral ventral cirri (Fig. 127j), identically to in U. polynu-
cleata and U. caudata. Streaks II-VI of proter and opisthe originate by division of five
long primary primordia (Fig. 127m, 0, p). According to FOISSNER (1983a), 1,2,3,3 (4),
and 5 (4) cirri originate from the primordia I-VI. However, if streak II produced only 2
cirri, U. emarginata could have only four transverse cirri. Thus, BERGER et al. (1985a)
changed it to the common pattern, namely 1,3,3,3,4, and 4.

o c cur r e n c e and e colo g y: Not very common in freshwater and terrestrial

habitats. Locus classicus is central New Jersey, USA, where STOKES (1885b) discovered
Urosoma emarginata in lentic water with aquatic plants. KAHL (1932) found it ''not sel-

~ Fig. 127a-l Urosoma emarginata (a, from STOKES 1885b; b, from KAHL 1932; c-h, from FOISSNER 1982; i-I,
from FOISSNER 1983a a-f, from life; g-I, protargol impregnation). a-c) Ventral views, a = 130-145 11m, b =
140 11m, c = 165 11m. Note conspicuous notch at posterior end of right margin. d) Layer of cortical structures
(mitochondria?) in surface (above) and lateral view (below). e, 1) Dorsal and left lateral view showing contrac-
tile vacuole with collecting canals and defecation (arrowhead), e = 185 11m, f = 175 11m. g-i) Ventral and dor-
sal infraciliature, g, h = 90 11m, i = 125 11m. j) Ventral infraciliature and nuclear apparatus of a very early mor-
phogenetic stage, 125 11m. k, I) Ventral infraciliature of early morphogenetic stages, Ie, 1= 110 11m. Arrows in
(I) mark 4 streaks in frontal area, including the modified anterior-most postoral ventral cirrus. OP = oral pri-
mordium; p 409.

Fig. 127m-t Urosoma emarginata (from FOISSNER 1983a m-t, protargol impregnation). m-q) Ventral and ~
dorsal infraciliature of middle morphogenetic stages, m, n = 125 11m, 0 = 115 11m, p, q = 130 11m. The
posterior-most frontoventral cirrus (lV/3) and the two posterior postoral ventral cirri (V/3 and V/5) are modi-
fied to primordia Five long primary primordia are recognisable (m, n). The primary primordia are divided and
the buccal cirrus and the undulating membranes are modified to primordia (0). Some cirri of the right marginal
row are also modified to primordia (p, q). r-t) Ventral and dorsal infraciliature oflate morphogenetic stages, r
= 130 11m, t = 115 11m. Differentiation of new frontal-ventral-transverse cirri. Parental structures white, new
black. AD = anlage of dorsal kineties; p 409. Figures on p 412.

Fig. 127u-y Urosoma emarginata (from FOISSNER 1983a u-y, protargol impregnation). u-w) Ventral and ~
dorsal infraciliature of late morphogenetic stages, u = 130 11m, v, w = 100 11m. Cirri originating from one
streak are connected by a broken line; supernumerary cirri will be resorbed later. The dorsomarginal rows (=
dorsal kineties 4) originate from parental marginal cirri (u). New caudal cirri are formed at posterior end of
new dorsal kineties 1-3 (w). x, y) Ventral and dorsal infraciliature of a very late morphogenetic stage, 180 11m.
The new cirri migrate to their final positions. Parental dorsal kinety 4 is still recognisable (y). Parental struc-
tures white, new black. AD. = anlage of new dorsal kinety 4; P 409. Figures on p 413.
412 SYSTEMATIC SECTION

,•
",
••
••
,,
••• ••
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127m
 Urosoma 413

....
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~~
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<:1" 1<:1"
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414 SYSTEMATIC SECTION

dom", mainly among Lemna, probably somewhere in Germany. FOISSNER (1982, 1983a)
isolated U. emarginata from various soils (water meadow, field) in the Tullnerfeld,
Lower Austria (Profile 2,5,6 in FOISSNER et al. 1985). I found few specimens in a leaf lit-
tered pond near the village of Bad Ischl, Austria, and in a six day old, saprobic sample
from a eutrophic pond in Salzburg city, Austria.
FOISSNER (1983a) cultured U. emarginata in salad medium diluted 1:1 with table wa-
ter (Volvic, France) and with some squeezed wheat grains to support bacterial growth;
high abundances occurred after 5-10 days. Biomass of 106 individuals 41 mg (FOISSNER
1987a).

Urosoma acuminata (STOKES, 1887) BOTsCHLI, 1889 (Fig. 128a-o, Table 24)

1887 Oxytricha acuminala STOKES, Ann. Mag. nat Hist, 20: III (Fig. 128a).
1888 Oxytricha acuminata, STOKES - STOKES, J. Trenton nat Hist Soc., 1: 290 (Fig. l28a).
1889 Oxytricha acuminata STOKES - BilTscHLI, Protowa, p 1748, 1749 (see nomenclature).
1932 Urosoma (Oxylricha) acuminata STOKES, 1887 - KAHL, Tierwelt Dt!., 25: 606.
1982 Urosoma acuminata (STOKES, 1887) - FOISSNER, Arch. Protistenk., 126: 109 (Fig. 128b-g; authoritative
redescription; I slide of neotype specimens is deposited in the OberOsterreichische Landesmuseum in
Linz, Upper Austria).
1987 Urosoma acuminata (STOKES, 1887) KAHr., 1932 - BERGER & FOISSNER, Zoo!. lb. Syst., 114: 231 (Fig.
128h-Q; 1 slide of protargoI impregnated specimens is deposited in the Oberosterreichische Landesmu-
seum in Linz, Upper Austria).

Nom e n c I at u rea n d t a x 0 nom y: Already BOTsCHLI (1889) proposed a classi-

fication in Urosoma; thus, he and not KAHL is the combining author. Furthermore, KAHL
(1932) classified Urosoma only as subgenus of Oxytricha, so that the correct name in his
revision is Oxytricha (Urosoma) acuminata. The morphometric characterisations by FOI-
SSNER (1982) and BERGER & FOISSNER (l987a) showed that U. acuminata rather constantly
has four macronuclear nodules. Therefore, synonymy with the binuclear U. caudata, as
proposed by BORROR (1972a), can be excluded.

M 0 r ph 0 log y and b i 0 log y: In life 110-220 x 20-35 /lm: 110-170/lm

(STOKES 1887b), 160-220 x 23-35 /lm (FOisSNER 1982), after BERGER & FOISSNER (l987a)
up to 50 /lm broad; maximum width usually at level of buccal vertex. Slightly contractile.
Posterior end tapered or tailed, often curved to the right, not, however, emarginated as in
Urosoma emarginata! Macronuclear nodules in life about 16 x 9 /lm, after protargol im-
pregnation 7-21 x 4-10 /lm. 2-8, in life 4 x 3 /lm sized micronuclei. Contractile vacuole
with collecting canals; anterior one sometimes with a small vesicle at level of cytostome.
After protargol impregnation many argentophilic globules especially around bases of cirri
(BERGER & FOISSNER 1987a). Endoplasm colourless, often opaque by many, about l/lm
sized ellipsoidal structures (bacteria?). Movement moderately rapid, often stands still for
short time and adheres close to substrate particles.
 Urosoma 415

c d e 9
Fig. 128a-g Urosoma acuminata (a, from STOKES 1887b; b-g, from FOISSNER 1982. a-e, from life; f, g, protar-
gol impregnation). a, b) Ventral views, a = 170 j.lm, b = 200 j.lm. c, d) Dorsal and lateral view showing con-
tractile vacuole with collecting canals (c), c, d = 175 j.lm. e) Pellicle with mitochondria(?). C, g) Ventral and
dorsal infraciliature of same specimen, 145 j.lm. P = ingested fungal spore; p 414.

Adoral zone ofmembranelles about 20-26 % of body length. Usually 3, rarely only 2
postoral ventral cirri. Transverse cirri about 15 flm long, distinctly subterminal. Marginal
and caudal cirri 10-15 flm long. Dorsal kinety 1 anteriorly, kinety 4 anteriorly and poste-
riorly slightly shortened.
Cyst spherical, in life 40-50 flm (43 flm on average; n = 17) across (BERGER & FOISS-
NER 1987a). Cyst formation takes a long time in cultures. Initially, the outer layer is
smooth and about 2 flm thick. Cyst densely filled with 1-3 flm sized, clod-like inclusions.
Sometimes contractile vacuole recognisable and movement of cytoplasm can be clearly
seen. A second type of cysts occurred rather frequently in older cultures. It had an about
3 flm thick, rough wall and a rather clear zone which sometimes contained fast moving
particles, probably some sort of parasite. There were many transitions between these two
types (Fig. 1281, m).

o c cur r e n c e and e colo g y: Not very common in the aufwuchs of freshwater

and in terrestrial habitats. Eurasia and Africa. Locus classicus is a pond with algae in
 ./:>.
0'1
-
X
-,{. \" ,,-
• r
-.,:"j'. ",,/' (" f
" ~ j < ',
,-'
",
/
k "~.. ,-'
,'1/ , \ .~Ii.
~./
Y-',.fI( ,........
,('
I : """: ,:.~' , ' :/ r
,, ~r;
f' , ,- ,-"
-.I' ,,~.
;
~ ~:...
,, , ~
, ~, ," /

., '
·t '-, ,-
,,, (/).
, , -,'~.. ; .j'-
~"©':."'.' -
. 'o;:"~ ,/ , -<
(/).
.,, ...,
,, ~" ~t~: :: :1 tTl
v~ ~.
T "o'~''': -7 (' ~
...,
I ,-,'" (' '1
L:: .: l n
(/).
tTl
(1
\" - ! ,/ /" ...,
T' '-,
.. r -, ..-/ ~
'\ -.J'
~,~~, ,j(
\.-, .
~
\'\
\10
Fig. 128h~ Urosoma acuminata (from BERGER & FOISSNER 1987a. h-m, from life; n, 0, protargol impregnation). h-j) Ventral, dorsal, and lateral view, h = 205 J.lm, i =
200 J.lm,j = 180 J.lm. k) Cross-section about in mid-body, bar = 30 J.lm. I, m) Resting cysts, 43 J.lm in diameter on average. n, 0) Ventral and dorsal infraciliature of same
specimen, 165 J.lm; p 414.
 Urosoma 417

USA, probably near Trenton, New Jersey, where STOKES lived and worked. FOISSNER
(1982) found U acuminata in the soil of a flood plain in the Tullnerfeld, Lower Austria
(see FOISSNER et al. 1985, "Profil2", for detailed description of site). BERGER & FOISSNER
(1987a) isolated it from the upper soil layer (1-5 cm) of an uncultivated grassland (domi-
nated by Poa sp.; about 1500 m above sea-level) in the Golan Hills, Israel.
Records not substantiated by illustrations: brook in Upper Austria (BLATTERER 1994);
soil from the Fuscher Valley, Austria (FOISSNER 1987b); conventionally farmed, sub-
alpine grassland in Styria, Austria (FOISSNER et al. 1990); sediment of unpolluted foothill
stream (Breitenbach) near the village of Schlitz, Germany (PACKROFF & ZWICK 1996); auf-
wuchs of the littoral from Lake Balaton, Hungary (TAMAs & GELLERT 1959); well in Italy
(GRISPINI 1938); Turiec River in Slovakia (TIRJAKovA 1993); Donor water reservoir,
USSR (ALEKPEROV 1989); soil from paddy field in Japan (TAKAHASID & SUHAMA 1991);
freshwater habitats in south-eastem Louisiana and Conestoga drainage basin in Pennsyl-
vania, USA (BAMFORTH 1963, CAIRNS 1965a); soil near Lake Baringo, Kenya (W. FOISS-
NER, pers. comm.).
Feeds on autotrophic flagellates, diatoms (Hantzschia sp.), and fungal spores (BER-
GER & FOISSNER 1987a, FOISSNER 1982). Biomass of 106 individuals 73 mg (FOISSNER
1987a). PATRICK et al. (1967) found U acuminata with low abundance in the Savannah
River, USA, at 12-17 °c, 9-<11 mg}-I O2, 0.5-1 mg I-I BOD5, 5-10 mg 1-1 CO2 free,
0.009-0.03 mg 1-1 NH3-N, 0.001 mg I-I NOi-N, 0.07-0.2 mg I-I N03--N, 1-3 mg I-I Cl".

Urosoma octonucleata BERGER & FOISSNER, 1989 (Fig. 129a-f, Table 24)

1989 Urosoma octonucleata BERGER & FOISSNER, Bull. Br. Mus. nat Hist. (Zool.), 55: 41 (reference numbers
of types slides deposited in British Museum: 1988:2:1:26 holotype, and 1988:2:1:27 paratype).

T a x 0 nom y: In life, U octonucleata can be easily confused with U. polynucleata be-

cause the body shape and the nuclear apparatus are very similar. Furthermore, the infra-
ciliature of the frontal area and the dorsal surface and many morphometric characters are
almost identical. However, they can be separated by the number of postoral ventral cirri
(3 in U octonucleata against 1 in U polynucleata) and the number and position of the
transverse cirri including the pretransverse ventral cirri (three to four distinctly subtermi-
nal in U octonucleata against two terminal in U polynucleata).

M 0 r p hoi 0 g y and b i 010 g y: In life 140-180 x 24-40 !lm. Vermiform, poste-

rior end tapered and usually curved to the right. Macronuclear nodules in life about
7 x 5 !lm, arranged in a line slightly left of median. 2-3 (usually 2) in life about 4!lm
sized, spherical micronuclei. Contractile vacuole distinctly in front of mid-body, during
diastole with two collecting canals. Cortical granules lacking. Cytoplasm with numerous
1 !lm sized, colourless granules and some about 7 !lm sized food vacuoles; posterior por-
tion with many cytoplasmic crystals.
 418 SYSTEMATIC SECTION

".
\

· :t

t \,
\
· '1
I' () l
\
· 'I
,
\
·1
,
,,'l~~
,
, I
'\
, ,
, , II
, ,
, P, \,

, , "- . 'I
"1\ ~\
}'
, ,
,
I \ , '\
' ~,
, ,
,
\
\
,.- , ~aJ \
, y
,
, , ($' "-, '\
'. '\
,,
,,
"•
')
'\

\\j"
...
•, f

Fig. 129a-f Urosoma octonucleata (from BERGER & FOISSNER 1989a. a-d, from life; e, f, protargol impregna-
tion). a--c) Ventral, lateral, and dorsal view, a = 170 J.1m, b, c = 160 J.1m. d) Colourless, about 2 J.1m sized struc-
tures (mitochondria?) close beneath the pellicle. e, 1) Ventral and dorsal infraciliature, 155 J.1m; p 417.

Adoral zone of membranelles about 23 % of body length; distal 4 membranelles

slightly separated from proximal, nearly perpendicularly arranged portion of adoral zone.
Bases of largest membranelles in life about 5 Jlm wide. Frontal cirri in life about 15 Jlm,
marginal cirri about 10 Jlm long. Postoral ventral cirri arranged in a line. Usually 2
slightly enlarged transverse cirri. Dorsal kinety 1 anteriorly, kinety 4 posteriorly short-
ened.

o c cur r e n c e and e col 0 g y: Locus classicus is a slightly reddish-brown soil

grown with Opuntia jicusindica and tufts of grass (0-5 cm, pH 4.8, 150 m above sea-
 Urosoma 419

level) in Garajan Kap, Madeira, Portugal. Feeds on bacteria. Biomass of 106 individuals
about 60 mg.

Urosoma polynucleata (FOISSNER, 1984) comb. nov. (Fig. l30a-i,

Tables 3, 24)

1984 Hemisincirra polynucleata FOISSNER, Stapfia, 12: 119 (l slide ofholotype and 1 slide ofparatype speci-
mens are deposited in the Oberllsterreichische Landesmuseum in Linz, Upper Austria).

T a x 0 nom y: The arrangement of the frontoventral cirri and undulating membranes,

the dorsal infraciliature, and the mitochondria (?) close beneath the pellicle indicate that
this species belongs to Urosoma. Should not be confused with U. octonucleata (more
postoral ventral and transverse cirri, Fig. 129a-f) and U. similis (2 macronuclear nodules,
Fig. l3la-g).

M 0 r p hoi 0 g y and b i 0 log y: In life 140-200 x 20-30 Jim. Vermiform, anterior

three quarters parallel sided, posterior quarter tapered and usually somewhat curved to
the left, sometimes slightly twisted. Macronuclear nodules in life 8 x 4 Jim. Two (rarely
up to 4), in life about 4 x 3 Jim sized micronuclei, usually one each near anterior and pos-
terior macronuclear nodules (Fig. l30a, f). Contractile vacuole with two long collecting
canals. Close beneath pellicle many tiny granules (mucocysts?) and ellipsoidal structures
(mitochondria?; Fig. 130c). Movement slow.
Adoral zone of membranelies about 18 % of body length. Invariably only 1 postoral
ventral cirrus (cirrus IVI2?). Transverse cirri at tip of tail (or lacking and possibly con-
fused with caudal cirri?). All cirri rather fine, 10-13 Jim long. Dorsal kinety 1 anteriorly,
kinety 4 posteriorly shortened (Fig. BOg). Morphogenesis commences with the forma-
tion of a small oral primordium just behind the buccal vertex (Fig. 130h).

o c cur r e n c e and e colo g y: Locus classicus of this autochthonous soil species

is the upper soil layer ofaxerothermic site ("Heisslande Althann") in the Tullnerfeld,
Lower Austria ("Profil 1" in ForssNER et al. 1985). Biomass of 106 individuals about
30 mg (ForssNER 1987a).

Urosoma simi/is (FOISSNER, 1982) comb. nov. (Fig. l31a-g, Table 24)

1982 Perisincirra similis FOISSNER, Arch. Protistenk., 126: 94 (Fig. 131a-g; 1 slide ofholotype and 1 slide of
paratype specimens are deposited in the Oberllsterreichische Landesmuseum in Linz, Upper Austria).
1984 Hemisincirra similis (FOISSNER, 1982) - FOISSNER, Stapfia, 12: 119.

T a x 0 nom y: The arrangement of the frontoventral cirri, the oral apparatus in Gono-
stomum pattern, and the possession of caudal cirri and 4 dorsal kineties indicate a class i-
420 SYSTEMATIC SECTION

Cl.
o
I
.. ..c

~ ~" ~ ~ _~_' '.:r~~ ~ I

.;;.~~SL.~ @~©~~
.... '"
" " " " " " "_" • '\. '\ ... .. '\
a:
" - ~
a:
"""""""""" ,,,, "

.0
 Urosoma 421

Fig.131a-g Urosoma similis (from FOISSNER 1982. a-c, from life; d-g, protargol impregnation). a-c) Ventral,
dorsal, and lateral view, a = 165 ~m, b, c = 185 ~m. d, e) Ventral and dorsal infraciliature of same specimen,
120 ~m. C, g) Ventral infraciliature of anterior and posterior portion of cell, bar = 20 ~m. This specimen has
only three frontoventral cirri. CC = caudal cirri, e = endoral (possibly the paroral), p = paroral, TC = transverse
cirri, III/2 = frontoventral cirrus III/2; p 419.

fication in Urosoma. In contrast, the type species of Hemisincirra HEMBERGER, 1985 (ba-
sionym: Uro/eptus kahli BUITKAMP, 1977) has no caudal cirri and only 3 dorsal kineties.

M 0 r p hoI 0 g y and b i 0 log y: In life 120-180 x 18-28 ~m. Lanceolate, often

slightly sigmoid, anteriorly often truncated, margins continuously converging posteriorly.
Macronuclear nodules in life 17 x 7 ~m, with many small and large nucleoli. Rather in-
variably two, in life about 4 ~m sized ellipsoidal or spindle-shaped micronuclei. Contrac-
tile vacuole slightly in front of mid-body, without distinct collecting canals; however,

~ Fig. 130a-i Urosoma polynucleata (from FOISSNER 1984. a-e, from life; f-i, protargol impregnation). a, b)
Ventral and dorsal view, 155 ~m. c) About 2 ~m sized structures (mitochondria?) close beneath the pellicle. d)
Cross-section in mid-body. e) Right lateral view, 155 ~m. C, g) Ventral and dorsal infraciliature, 145 ~m. h)
Ventral infraciliature of a very early morphogenetic stage, 140 ~m. i) Ventral infraciliature of anterior portion
of cell. CC = caudal cirri, OP = oral primordium, RMR = posterior-most cirrus of right marginal row, just be-
hind two transverse cirri; p 419.
422 SYSTEMATIC SECTION

sometimes a small additional vacuole in front of it; immediately before systole the two
vacuoles fuse. Cortical granules and typical subpellicular "mitochondria" lacking. Cyto-
plasm packed with many colourless, about 0.5 Ilm sized granules and tiny yellowish crys-
tals. Food vacuoles 4-8 Ilm across. Movement rapid.
Adoral zone of membranelles about 20 % of body length. Adoral zone and undulat-
ing membranes in Gonostomum pattern. Frontal cirri in life about 15 Ilm long, right cirrus
slightly enlarged, left arranged very close to adoral zone of membrane lIes. Some speci-
mens with less than 4 frontoventral cirri (Fig. 131t). Transverse cirri in life about 18 Ilm
long, protrude distinctly beyond posterior end of cell. Distance between basal body pairs
in dorsal kineties 1 and 2 posteriorly about 2 times as large as anteriorly. Three caudal
cirri, one each on dorsal kineties 1-3.

o c cur r e n c e and e colo g y: Autochthonous soil species (FOISSNER 1987a). L0-

cus classicus is the village of Zwentendorf, Lower Austria, where FOISSNER (1982) dis-
covered U. similis in the upper soil layer of an intensively fanned field. Records not sub-
stantiated by illustrations: rarely in organically and conventionally fanned sub-alpine
grassland in Styria, Austria (FOISSNER et al. 1990); agricultural soils in Czechoslovakia
(TIRJAKovA 1988); soil in Costa Rica (FOIssNER 1995b). Feeds on bacteria. Biomass of
106 individuals about 32 mg (FOIssNER 1987a).

Urosoma karini FOISSNER, 1987 (Fig. 132a-e, Table 24)

1987 Urosoma karini FOISSNER, Jber. Haus Nat. Salzburg, 10: 62 (I slide ofholotype and I slide of paratype
specimens are deposited in the OberOsterreichische Landesmuseurn in Linz, Upper Austria).

T a x 0 nom y: Urosoma karini is very similar, possibly even synonymous with U. sa/-
mastra from Africa (Fig. 133a-c). Concerning the infraciliature there are only minor but
possibly significant differences in the arrangement of the undulating membranes and the
number of marginal cirri. However, live observation (cortical granules present or not?) on
the African species should be awaited.

M 0 r ph 0 log y and b i 0 log y: In life 100-160 x 25-35 Ilm. Long elliptical,

sometimes almost parallel sided, posterior end rounded; not contractile. Two (rarely 3),
after protargol impregnation 18-31 x 7-11 Ilm sized macronuclear nodules. 1-3 (usually
2), after protargol impregnation 6 x 4 Ilm sized micronuclei. Contractile vacuole without
distinct collecting canals. Cortical granules tiny, colourless, arranged in rows, well distin-
guishable from ellipsoid mitochondria (Fig. 132b). Posterior cell part with many crystals,
which appear dark at low magnification and bright field illumination. Slow movement.
Adoral zone of membranelles about 20 % of body length. Row of frontoventral cirri
slightly curved. Postoral ventral cirri distinctly behind level of buccal vertex, form an ir-
regular row with frontoventral cirri; some specimens with 2-3 supernumerary postoral
 Urosoma 423

Fig. 132a-e Urosoma karini (from FOISSNER 1987b. a-<:, from life; d, e, protargol impregnation). a) Ventral
view, 135 Jim. b) Pellicle with mitochondria and colourless cortical granules. c) Right lateral view, 135 Jim. d,
e) Ventral and dorsal infraciliature of same specimen, 155 Jim. G = cortical granules, M = mitochondria;
p422.

ventral cirri. Transverse cirri fine, about 20 Jlm long, protrude distinctly beyond posterior
end of cell. Rather invariably 6 pretransverse ventral and transverse cirri (very likely 2
pretransverse ventral cirri and 4 transverse cirri; see, however, Table 24). Marginal cirri
in life about 12 Jlm long. Dorsal kinety 1 (and 3?) anteriorly slightly shortened, kinety 4
about 1/3 of body length.
Cyst spherical, about 32-40 Jlm in diameter (n = 14), colourless to slightly yellowish,
covered by a very inconspicuous mucous layer. Surface of cyst with many fibrils, similar
to in Engelmanniella (data kindly supplied by W. FOISSNER, from a terrestrial population
from st. Vincent, an island in the Caribbean Sea).
424 SYSTEMATIC SECTION

Fig. 133a-c Urosoma sal-

mastra (from DRAGESCO &
DRAGESCO-KERNEIS 1986. a-c,
protargol impregnation). a, b)
Ventral infraciliature, a =
75 11m, b = 65 11m. c) Dorsal
infraciliature, 70 11m.

Occurrence and
e colo g y: Locus clas-
sicus is the upper soil
layer of an alpine pas-
ture (PH 7.5, 1,270 m
above sea-level) in the
Fuscher Valley in Salz-
burg, Austria. Accord-
ing to ForSSNER (l987a),
U. karini is an auto-
chthonous soil species.
W. ForSSNER (pers.
comm.) found it in a
soil sample from St
Vincent, an island in the
Caribbean Sea. Feeds
on heterotrophic flagel-
lates and bacteria.

Urosoma salmastra (DRAGESCO & DRAGESCO-KERNEIS, 1986) comb. nov.

(Fig. 133a~, Table 24)

1986 Oxytricha salmastra DRAGESCO & DRAGESCo-KERNEIS, Faune tropicale, 26: 467.

T a x 0 nom y: The anteriorly displaced frontoventral cirrus IIV2, the arrangement of the
undulating membranes, and the dorsal infraciliature indicate a classification in Urosoma.
Possibly the senior synonym of U. karini (see there). Reinvestigation (especially live ob-
servation) necessary to check whether cortical granules are present or not.

M 0 r ph 0 log y: After protargol impregnation 50-100 x 15-23 Jim. Long elliptical,

posteriorly rounded. Macronuclear nodules about 12 ~m long. Two about 2 Jim sized mi-
cronuclei. Cortical granules not mentioned. Adoral zone of membranelles about 30 % of
body length. Endoral very short, arranged in front of paroral (if correct, an important dif-
 Urosoma 425

ference to U. karim). Some specimens with 7 or 9 frontoventral cirri. Dorsal kinety 4

about 113 of body length.

o c cur r e n c e: Locus classicus is a salt marsh at the lagoon of Cotonou, Benin (DRA-
GESCO & DRAGESCO-KERNEIS, 1986). CHAOUlTE et al. (1990) found U. sa/mastra in mineral
and thermo-mineral waters in France.

Incertae sedis

Urosomagigantea (HoRvATH, 1933) comb. nov. (Fig.134a-w, Tables 3, 24)

1933 Oxytricha gigantea HORVA11l, Arch. Protistenk., 80: 298 (Fig. 134a, b).
1935 Urosoma (filr Oxytricha) gigantea J. HORVA11l, 1933 - KAHL, Tierwelt Dtl., 30: 841.
1987 Urosoma gigantea (HoRvA11l, 1933) KAHL, 1935 - BERGER & FOISSNER, Zoot. lb. Syst., 114: 233 (Fig.
134c--g; 1 slide of protargol-impregnated specimeus is deposited in the OberOsterreichische Landesmu-
seum in Linz, Upper Austria).
1988 Oxytricha gigantea HORVA11l, 1933 - BERGER & FOISSNER, Arch. Protistenk., 136: 66 (Fig. 134h-w).

Nom e n c I a t u rea n d t a x 0 nom y: KAHL (1932, 1935) classified Urosoma as

subgenus of Oxytricha, so that the correct name in KAHL (1935) is Oxytricha (Urosoma)
gigantea. Consequently, KAHL (1932) cannot be the combining author as erroneously as-
sumed by BERGER & FOISSNER (1987a). Thus, I formally transfer it to Urosoma, although
its systematic position is uncertain (see list of synonyms). The Gonostomum-like oral ap-
paratus, the distinct mitochondria close beneath the pellicle, and the reduced number (4)
of dorsal kineties (despite the large size) indicate a classification in Urosoma. By
contrast, the arrangement of the frontoventral cirri, the lack of primary primordia, and the
long adoral zone of membranelIes (37 %) do not match the characterisation of Urosoma.
The type population and the BERGER & FOisSNER (1987a, 1988b, c) population agree very
well in body shape and size, ventral infraciliature, arrangement of the undulating mem-
branes, possession of short rows of cortical granules, and habitat; thus, I have no doubt
that the identification is correct, although HORvAm (1933) counted 5 dorsal kineties.

M 0 r p hoI 0 g y and b i 0 log y: In life 170-230 x 65-90 ~m, according to

HORvAm (1933) 200-230 ~m, according to BERGER & FOISSNER (1987a; n = 3)
170-210 ~m long. Long ovoid, usually distinctly narrowed posteriorly, sometimes body
margins almost in parallel; both ends rounded. Macronuclear nodules in life brightly shin-
ing, about 32 x 14 ~m, after protargol impregnation 18-28 x 7-11 ~m (mean =
21 x 9 ~m; n = 25); nucleoli large. 2-7 (usually 3), in life about 3 ~m sized globular mi-
cronuclei. Cortical granules spherical, less than 1 ~m across, colourless, arranged in short
rows (Fig. 134t). Cytoplasm greyish, packed with cytoplasmic crystals, especially in pos-
terior part of cell. Movement moderately rapid.
 426 SYSTEMATIC SECTION

, ,
1 2 3

a 9
Fig. 134a-g Urosoma gigantea (a, b, after HORvArn 1933; c-g, from BERGER & FOISSNER 1987a a, opal blue
staining after BRESSLAU; b, after live and stained material; c-g, from life). a--e) Ventral, lateral, and dorsal
views, a, b = 265 11m, c = 220 11m, d, e = 175 11m. f) Pellicle with ellipsoid structures (mitochondria?) and tiny
«111m), colourless cortical granules. g) Resting cyst, diameter 3 = 145 11m. M = mitochondria?, sG = cortical
granules, 1,2,3 = diameters of cyst (see text); p 425.

Adoral zone of membranelles about 37 % of body length. Bases of largest mem-

branelles in life about 8 11m wide. Buccal area largely covered by an arched hyaline
plasma lip. Paroral after protargol impregnation 20-27 11m (mean = 24 11m; n = 25), en-
doral 17-25 11m (mean = 20 11m; n = 25) long. Frontal cirri in life about 25 11m long. Ar-
rangement of frontal, ventral, and transverse cirri rather invariable (Fig. 134a, h). Trans-
verse cirri in life about 30 11m long, distinctly protruding beyond posterior end of cell.
Marginal cirri in life about 21 11m long; right marginal row begins at level of right frontal

Fig. 134h-m Urosoma gigantea (from BERGER & FOISSNER 1988b. h-m, protargol impregnation). h, i) Ventral ~
and dorsal infraciliature of a non-dividing specimen, 160 11m. The cirri of a primordium are connected by a
broken line. j, k) Ventral infraciliature of very early morphogenetic stages,j = 180 11m, k = 170 11m. I, m) Ven-
 Urosoma 427

3
2 I 1- nO

.;;'-
\ 'J
'J
"
~
"0
'/
0
1 '/
'.
'-, '. " !
"
,I
,

~ A.·
!
"
°6
R

",
'. /

" ?:
J
IV-
,I 'J ,,
"~
'. 11
,/

r',
'-
\
'\
\ ,~ ''1J ,
,
" ,
f' \ /

T ,
,- , 0
I'c...
" '.
:1

~
f
'.'

'~
,
-.. ~>
VI- '-
, .\ >;1
, '
'-, \,
" \
"'. '-,
"
" '.
'.-., ,

·r; \"
o , ~1. ~
A 'A
o Q j
A""
..........
u \
~':~ (~
.. ~
0'
.

~
•. 0 ,, .
.'

,, :,

.-
,.-
,,
, \\~J'I/
(:" .

.- ,

,,
1
;
Q
,
,
I

traI infraciliature of early morphogenetic stages, I = 180 pm, m = 165 pm, Arrowhead marks frontoventral cir-
rus IIII2. J-VI = frontal-ventral-transverse cirri primordia J-VI, 1--4 = cirri within a primordium and dorsal ki-
neties 1--4; p 425.
428 CTION
SY ST EM A TIC SE

f...)'.•..'
.
• • ft,
" ,

-
- ?
I)
Q 0
, ,/l
I)
,

13 4n ' t} Q 0 P

f o

( /
!o?
1.:-
I: ;.:-
;"
\~,
,, f:
;?'

,
,
"
h.·.~O
(j f; {
'I
I,
•
i
\:"
\,
\1
o " ,,
, I

o
o
,
°0o I
,
(/ " '.
q r
 Urosoma 429

,
,
,
. ....
'
~fJ
,.'
.-:-
,~
r.:.
.. · ..

Fig. 134t-w Urosoma g;gantea (from BERGER & FOISSNER 1988b. t-w, protargol impregnation). t, u) Dorsal
and ventral infraciliature of very late morphogenetic stages, t = 180 11m, u = 210 11m. v, w) Ventral and dorsal
infraciliature of an opisthe, 110 11m. Parental structures white, new black; p 425.

cirrus, terminates slightly subterminally, left row J-shaped. Dorsal kinety 1 slightly short-
ened anteriorly, kinety 4 tenninates in mid-body (Fig. 134i).
In cultures U gigantea encysted after conjugation. Cysts yellowish, consist of (i) an
outer hyaline mucous layer with adhering bacteria, (ii) an inner, relatively finn mucous
layer with 2 J!m sized ellipsoid yellow inclusions, (iii) an about 3.5 J!m thick cyst wall,

~ Fig. 134n-s Urosoma g;gantea (from BERGER & FOISSNER 1988b. n-s, protargol impregnation). n-p) Ventral
and dorsal infraciliature of middle morphogenetic stages, n = 195 11m, 0, p = 185 11m. q-s) Ventral infracilia-
tore oflate morphogenetic stages, q = 165 11m, r = 180 11m, s = 170 11m. Arrow in (s) marks a supernumerary
cirrus within primordium VI. Parental structures white, new black. P = primordium of new dorsal kinety 4;
p425.
430 SYSTEMATIC SECTION

and (iv) the cytoplasm with 4-8 /lm sized globules. Diameters of cysts (see Fig. 134g; n
= 14): 1 = 52-70/lm (mean = 65/lm), 2 = 76-105/lm (mean = 90/lm), and 3 =
111-150 /lm (mean = 132 /lm).
M 0 r p hog e n e sis of cell division commences apokinetally with the prolifera-
tion of basal bodies in the area between the postoral ventral cirri and transverse cirri (Fig.
134j). A large oral primordium is formed, its narrowed anterior portion elongates to the
level of the buccal vertex and splits in the middle region to form streaks I and II. The pos-
toral ventral cirri V/3 and V/4 are modified to primordia (Fig. 134k, 1). Next, the postoral
ventral cirrus IVl2 is disorganised. Five streaks of the opisthe are recognisable at this
stage (Fig. 134m). Simultaneously the buccal cirrus (11/2) and the cirrus 11112 organise
primordia for the proter. Three other primordia are present at the level of the cytostome.
At least the middle and right primordium are derived from a streak of the opisthe. Six
frontal-ventral-transverse primordia are discernible in the next stage in both the proter
and the opisthe. Cirrus IV13 has been reabsorbed (Fig. 134n). Some loosely arranged ba-
sal bodies are present between the right streak (primordium VI) of the proter and primor-
dium IV of the opisthe. Morphogenesis continues with elongation of the frontal-ventral-
transverse streaks and the formation of marginal primordia in the anterior and middle
portion of the right marginal row (Fig. 1340). The proliferation of new basal body pairs
occurs at two levels in dorsal kineties 1, 2, and 3 (Fig. 134p). The parental undulating
membranes commence with reorganisation when the undulating membranes of the opis-
the are formed. The anterior cirri are already segregated from the frontal-ventral-
transverse streaks. Both the right and left marginal primordia are elongated (Fig. 134q).
The primordia of dorsal kinety 4 originate at the anterior end of the right marginal pri-
mordia at about the time the macronuclear nodules fuse (Fig. 134r). Morphogenesis then
continues very similarly to, for instance, in Oxytricha granulifera. Unfortunately, we
could not fmd the exact origin of all primordia in either the proter or the opisthe (BERGER
& FOlSSNER 1988b). Thus, the data of Urosoma gigantea are not used in Table 4.

o c cur r e n c e and e colo g y: Locus classicus is a periodically occurring, sodium

hydroxide rich pond north ofSzeged (Hungary). HORVATH (1933) found U. gigantea also
in infusions ofdty soil from this pond. BERGER & FOlSSNER (1987a) isolated it from a very
similar habitat, namely from soil of a saline grassland with halophytes near a lake (Neu-
siedlersee) in eastern Austria. Feeds on bacteria, fungi, heterotrophic flagellates, and cili-
ates like Vorticella sp. Biomass of 106 individuals 270 mg (FOlSSNER 1987a).

Insufficient redescriptions

Oxytricha caudata - DUJARDIN, 1841, Zoophytes, p 420, Planche XIII, Fig. 6.

Oxytricha caudata, EHR. - DUMAS, 1929, Microzoaires, p 70 (Fig. 236a, b) and DUMAS,
1930, Microzoaires, p 53 (Fig. 236r, 239g). Remarks: Very likely an Uroleptus because
4-5 cirral rows are present. Pond in France.
 Tachysorna 431

Oxytricha caudata - FROMENTEL, 1876, Microzoaires, p 266, Planche XIII, Fig. 4. Re-
marks: Freshwater in France.

Urosorna cienkowski Kow. - CONN, 1905, Bull. Conn. St. geol. nat. Hist. Surv., 2: 59
(Fig. 225i). Remarks: Freshwater in Connecticut, USA.

Urosorna sp. - LEPSI, 1927, Studii Cerc. Acad. RPR, 12: 126 (Fig. 226j). Remarks: Per-
haps U. caudata. In life 146 x 56 ).1m. Between Oscillatoria. Poly- or mesosaprobic. Ro-
mania.

Urosorna sp. - LEPSI, 1965, Protozoologie, p 35 (Fig. 225h). Remarks: Macronuclear

nodules arranged biserially. Romania?

Tachysoma STOKES, 1887

1887 Tachysoma STOKES, Ann. Mag. nat. Hist., 20: 108 - Type (not fixed by STOKES 1887b; subsequent des-
ignation by BORROR 19728, piS): Trichoda pellione/la MOLLER, 1773.
1888 Tachysoma, STOKES - STOKES, J. Trenton nat. Hist. Soc., 1: 293.
1932 Tachysoma STOKES, 1887 - KAm., Tierwelt Otl., 25: 604 (see nomenclature).
1972 Tachysoma STOKES, 1887 - BoRROR, J. Protozool., 19: IS.
1974 Tachysoma STOKES - STILLER, Fauna Hung., 115: 138.
1997 Taclrysoma STOKES, 1887 - BERGER & FOISSNER, Arch. Protistenk., 148: 148.

C h a r act e r i sat ion: Adoral zone of membranelles formed like a question mark.
Undulating membranes in Oxytricha pattern. Frontoventral cirri in V-shaped pattern or
arranged almost in a row. Postoral ventral cirri in dense cluster behind buccal vertex.
Two pretransverse ventral and (4 to) 5 transverse cirri. One right and 1 left row of mar-
ginal cirri. Four to 6 dorsal kineties. Caudal cirri absent. Frontal-ventral-transverse cirri
originate from primary primordia or separate anlagen. Dorsal morphogenesis in Tachy-
sorna, Oxytricha, or Urosornoida pattern.

Add i t ion a I c h a r act e r s: Dorso-ventrally flattened about 2:1. Very flexible.

Two macronuclear nodules slightly left of median. Contractile vacuole about in mid-body
or slightly in front of it. Pellicle and cytoplasm colourless. Buccal area small to moder-
ately broad. Transverse cirri protrude distinctly beyond posterior end of cell. Marginal
rows distinctly separated posteriorly.

Nom e n c I at u rea n d t a x 0 nom y: KENT (1882, p 783) proposed the name

Mesotricha for Sty/onychia echinata, a generally accepted junior synonym of T. pe/-
lionel/urn, to separate it from other oxytrichids because of the long dorsal cilia. However,
this character is inappropriate for the distinction of supra-specific taxa. Furthermore,
Mesotricha is a nomen oblitum, because neither KAHL (1932), CORLISS (1979), JANKOWSKI
(1979), nor anyone else mentioned it. KAHL (1932) classified Tachysorna as subgenus of
432 SYSTEMATIC SECTION

Oxytricha; the correct name in his revision is thus Oxytricha (J'achysoma). Some incor-
rect subsequent spellings of Tachysoma: Tachiosoma pellionella (BICzOK 1979, P 127;
HAMAR 1971, P 120); Tachyosoma pellionella (SCHONBORN 1984, P 22); Taxysoma (Oxy-
tricha) pel/ionel/a KAlIL (LEPINIS et al. 1973, p 169).
The characterisation is according to BERGER & FOISSNER (1997). See chapter 2 in the
general section for a detailed description of patterns mentioned. Cladistic analysis re-
vealed that Tachysoma is closely related to Gonostomum and Urosoma because of pri-
mary primordia formation (Fig. 25a). However, in Tachysoma this type of primordia for-
mation occurs only in the type species, T. pellionellum, and not in T. terricola, the second
species so far morphogenetically analysed (HEMBERGER 1982). Different strategies not
only in the ventral but also in the dorsal morphogenesis suggest that Tachysoma is an ar-
tificial assemblage of flexible oxytrichids unified by the loss of caudal cirri, a character
which very likely evolved independently in several oxytrichids, namely Tachysoma, His-
trieulus, Pleurotricha, and Parastylonychia.
Tachysoma hyalina BERGER, FOISSNER & ADAM, 1984, T. perisincirra HEMBERGER,
1985 (both without postoral ventral cirri), and T. longa HEMBERGER, 1985 (postoral ven-
tral cirri left of frontoventral cirri) were transferred to Lamtostyla BUITKAMP, 1977b by
BERGER & FOISSNER (1987a, 1988a). Tachysoma raptans HEMBERGER, 1985 (Arch. Protis-
tenk., 130: 414) differs from Lamtostyla longa and L. granuliferum FOISSNER, 1997b only
in some morphometric characters (number of adoral membranelles and marginal cirri);
FOISSNER (1997b) thus transferred it to Lamtostyla: L. raptans. However, detailed data on
ventral and dorsal morphogenesis (and likely molecularbiological characteristics) are
needed for final classification of these species. Their ventral infraciliature is reminiscent
of Gonostomum, especially as concerns the "postoral" ventral cirri, which are arranged
right (and not behind!) of the proximal portion of the adoral zone of membranelies.
Tachysoma spec. in WILBERT & KAHAN (1981, P 91) has several frontoventral cirri
arranged in zigzag, indicating that it is a holostichid hypotrich. The marine T. parvulum
CORLISS & SNYDER, 1986 is superficially described; according to PETZ et al. (1995, p 170)
it is also a holostichid, and they thus transferred it to Notocephalus, N. parvulus (CORLISS
& SNYDER, 1986) PETZ, SONG & WILBERT, 1995.
Tachysoma should not be confused with Oxytricha and Urosomoida, which, howev-
er, have inconspicuous caudal cirri (use differential interference contrast optics or protar-
gol impregnation). Five species (T. pellionellum, T. granuliferum, T. humicola, T. dra-
gescoi, T. ovata) are described in detail, that is, both by live observation and after protar-
gol impregnation.

Key to species

1 Linmetic or terrestrial ................................................ 2

Marine ................................................................... 8
2 Length in life about 200 ~m (Fig. 138a) ...................... T. chilensis (p 451)
 Tachysoma 433

Length in life less than 150 11m .......................................•• 3

3 Dorsal cilia about 10-15 11m long ....................................... 4
Dorsal cilia 2-5 11m long .............................................. 6
4 Each macronuclear nodule with an adjacent micronucleus; terrestrial (Fig. 139i) .. .
· . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. T. terricola (p 458)
Only one micronucleus between the two macronuclear nodules; freshwater (for ex-
ample, Fig. 135g) .................................................... 5
5 Posterior-most left and right marginal cirrus distinctly enlarged and elongated (rare
species; Fig. 139h) ....................................... T. bicirratum (p 457)
All marginal cirri of about same length (very common species; Fig. 136a)......... .
· . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. T. pellionellum (p 433)
6 (3) Cortical granules yellowish to orange-yellow (Fig. 140c) ..................... .
· ............................................. " T. granuliferum (p 459)
Cortical granules absent .............................................. 7
7 Length in life about 40-60 11m; terrestrial (Fig. 138b-e, 139a-t) ................. .
. . . . . . . . . . . . . . .. T. humicola (p 452; identification of subspecies, see couplet 10)
Length in life about 130-140 11m; freshwater (Fig. 139g) .... T. balatonica (p 456)
8 (1) Length in life 150-260 11m .......................... T. rigescens (p 461)
Length in life about 35-50 11m ......................................... 9
9 Single micronucleus between macronuclear nodules; 5 transverse cirri (Fig. 141j, p,
q) .................................................... T. ovata (p 464)
Two very small micronuclei, that is, both macronuclear nodule with each one micro-
nucleus; 4 transverse cirri (Fig. 141d, e, t) ................. T. dragescoi (p 463)
10 Posterior-most dorsal cilia 2-3 11m long (Fig. 139a, e) T. humicola humicola (p 456)
Posterior-most dorsal cilia about 10 11m long (Fig. 138b, c, e) ................. .
· . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. T. humicola longisetum (p 456)

Tachysoma pellionellum (MOLLER, 1773) BORROR, 1972 (Fig. 135a-s,

136a-x, 137, Tables 3, 25-28)

1773 Trichoda pellionella MOLLER, Vennium Terrestrium et F1uviatilium, p 80.

1786 Trichoda pel/ionella - MOLLER, Animalcula Infusoria, p 222 (Fig. 135a').
1824 Oxitricha pelionella - BORY DE SAINT-VINCENT in LAMOUROux, BORY DE SAINT-VINCENT & DESLONG-
CHAMPS, Encyclopedie methodique, p 595 (incorrect subsequent spelling ofpe/lione/Ia).
1824 Oxitrichajob/otii BORY DE SAINT-VINCENT in LAMOUROux, BORY DE SAINT-VINCENT & DESLONGCHAMPS,
Encyclopedie methodique, p 596 (according to EHRENBERG 1838, a synonym).
1826 Oxilricha pelionella - BORY DE SAINT-VINCENT, Essai d'une classification, p 56 (incorrect subsequent
spelling ofpellionella).
1831 Oxytricha pel/ionel/a - EHRENBERG, Abh. preuss. Akad. Wiss., year 1831: 118.
1833 Oxytricha pellionella - EHRENBERG, Abh. preuss. Akad. Wiss., year 1833: 174, 325, Tafel III, Fig. VI
(on p 325 it is designated as "Das hautige Hecheltierchen").
1838 Oxytricha pel/ionella - EHRENBERG, Infusionsthierchen, p 364 (Fig. 135a).
1850 Oxytricha pellioneUa BORY - DIESING, Systema Helminthum I, p 157.
434 SYSTEMATIC SECTION

~
,6
I 5'
3 4 I.
I

(J
I, I
, ,

If'iI'A~ ~,

'U'
" y .......
.
,
\

\
\

, J

Fig. 13Sa-g Tachysoma pel/ionel/um (a, after EHRENBERG 1838; a', from MOLLER 1786; b--d, from STEIN
1859b; e, from PATSCH 1974; 1: g, originals of a population from the Fuscher-Ache, a small, oligosaprobic to
betamesosaprobic alpine river in Salzburg, Austria a-d, from life; e-g, protargol impregnation). a-c) Ventral
views, sizes not indicated. d) Dorsal view. The two micronuclei are probably a misobservation because the
general appearance indicates unambiguously that the identification is correct e, f) Ventral infraciliature, e =
70I.lln, f= SO /lm. g) Dorsal infraciliature, 60 /lm. Arrow marks the single micronucleus between the two mac-
ronuclear nodules. FC = right frontal cirrus (lIII3), FVC = anterior-most frontoventral cirrus (VII4), 1--6 = dor-
sal kineties 1--6; p 433.

1858 Stylonychia echinata CLAPAREDE & LACHMANN, Mem. lost. nato. genev., 5: 165 (Fig. 136r).
1859 Pleurotricha echinata - STEIN, Organismus der Infusioosthiere I, p 171.
1859 Oxytricha pel/ionel/a. EHRBG. - STEIN, Organismus der Infusionsthiere I, p 185 (Fig. 135b-d).
1878 Oxytricha pelionella - STERKI, Z. wiss. Zool., 31: 55, 56 (Fig. 1360, q; incorrect subsequent spelling).
1882 Oxytricha pellionella, MOLL. sp. - KENT, Manual infusoria II, p 786.
1887 Tachysoma agile STOKES, Ann. Mag. nat. Hist., 20: 108 (Fig. 136k).
1887 Tachysoma mirabile STOKES, Ann. Mag. nat. Hist., 20: 109 (Fig. 1361).
1888 Tachysoma agilis, STOKES - STOKES, J. Trenton nat. Hist. Soc., 1: 294.
1888 Tachysoma mirabilis, STOKES - STOKES, J. Trenton nat. Hist Soc., 1: 295.
1901 Oxytrichapel/ionel/a (O.-F. MOLL.)- Roux, Mem. lost. nato. genev., 19: 101 (Fig. 1365).
1908 Oxytrichapellionella O. F. MOLL. - BORGER, An. Univ. Chile, 122: 187 (Fig. 135q).
1932 Tachysoma (Oxytricha) pellionella (MOLLER - STEIN, 1859) - KAHL, Tierwelt Dtl., 25: 606 (Fig. 136i).
1932 Tachysoma echinata (CLAP. u. L.)- KAHL, Tierwelt Dtl., 25: 617.
1932 Tachysoma mirabilis STOKES, 1887 - KAHL, Tierwelt Dtl., 25: 606.
1933 Oxytricha pellionel/a O. F. M. - GAJEWSKAJA, Zoologica, 32: 147 (Fig. 136t).
1935 Oxytricha pellionella (MOLLER) KAHL, 1932 - WANG & NIE, Sinensia, 6: 497 (Fig. 136m).
 Tachysoma 435

1953 Tachysoma pellionella - JIROVEK, WENIG, FOTI, BARTOS, WEISER & SRAMEK-HUSEK, Protozoologie,
p 512, 514 (Fig. 136p).
1963 Tachysoma pel/ionella - SLAoECEK, Sb. vys. Sk. chem.-technol. Praze, 7: 590 (Fig. 136p).
1968 Oxytricha (l'achysoma) pellionella (0. F. MOLLER) STEIN, 1859 - CHORlK, Free-living ciliates, p 135
(Fig. 136u).
1969 Tachysoma pellionella (MOLLER-STEIN, 1859) - CURDS, Wat. Poll. Res., No. 12: 68 (Fig. 136n).
1972 Tachysoma pel/ioneUa (MOLLER, 1786) KAHL, 1932 - BORROR, J. Protozool., 19: 15.
1972 Tachysoma pellionella (0. F. MOLLER) - BlcK, Ciliated protozoa, p 182 (Fig. 136h).
1974 Tachysoma pellionella (0. F. MOLLER) - PATSCH, Arb. lnst. landw. Zool. Bienenk., I: 65 (Fig. 135e).
1975 Tachysoma pel/ionella (MOLLER-STEIN, 1859) KAlIL, 1932 - GROLIERE, Protistologica, 11: 491 (Fig.
1361: g).
1977 Oxytricha (l'achysoma) pellionella (0. F. M.) STEIN 1859 - BERECZKY, Annis Univ. Scient. bpest. Rola-
ndo EOtvOs, 18-19: 169 (Fig. 135r).
1979 Oxytricha pe/ionel/a MOLL.-STEIN, 1859 - MAMAEvA, Infusoria of the Volga basin, p 71 (Fig. 135s; in-
correct subsequent spelling).
1981 Tachysoma pellionella (MOLLER, 1786) - FOISSNER & DIDIER, Annis Stn Iimnol. Besse, IS: 259 (Fig.
136a--e).
1981 Tachysoma pellionella (MOLLER) STEIN 1859 - CHARDEZ, Revue verviet. Hist. nat., 38: 53 (Fig. 136j).
1982 Tachysoma pel/ionella (MOLLER, 1786) KAHL, 1932 - HEMBERGER, Dissertation, p 243 (Fig. 135h-o).
1988 Tachysoma pel/ionel/um (0. F. MoLLER, 1773) - FOISSNER, Hydrobiologia, 166: 30, 43 (emendation).
1991 Tachysoma pellionellum (MUELLER, 1773) BORROR, 1972 - FOISSNER, BLATIERER, BERGER & KOHMANN,
lnformationsberichte Bayer. Landesamtes fUr Wasserwirtschaft, 1/91: 304.
1993 Tachysomapel/ionel/um (MOLLER, 1773)- SHIN & KIM, Korean J. Zool., 36: 228 (Fig. 136v-x).
1997 Tachysoma pellionellum (MOLLER, 1773) BORROR, 1972 - BERGER, FOISSNER & KOHMANN, Bestimmung
und Okologie der Mikrosaprobien, p 167 (Fig. 135p).

Nom e n c I at u rea n d t a x 0 nom y: KAliL (1932) classified Tachysoma as sub-

genus of Oxytricha; the correct names in his revision are thus Oxytricha (Fachysoma)
pellionella, O. (Fachysoma) echinata, and 0. (Fachysoma) mirabilis. This implies that
KAliL (1932) is not the combining author as generally, but erroneously, assumed (for ex-
ample, BORROR 1972a, JANKOWSKI 1979). In my opinion, BORROR (1972a) is the combin-
ing author in that he fIXed it as the type species.
Incorrect subsequent spellings: Histrio pelionella (LACKEY 1936, P 269); Oxytricha
(Fachysoma) pelionella (LIEPA 1973, p 34); Oxytricha pelionella (for example, BALDENS-
PERGER 1927, P 243; ClENKOWSKY 1855, P 305; GlTILESON & HOOVER 1969, P 750; MA-
MAEVA 1979c, P 233; MYLNIKOVA 1993, P 193; SCHIEDERMAYR 1887, P 13; ZoLOTAREV
1987, P 114); Oxytricha pellionela (NIKOLYUK 1965, P 46); Oxytricha pellionne//a O. F.
M. (lLoWAISKY 1926b, P 97; Roux 1900, p 464); Oxytricha perione//a MOLL (SHIBUYA
1927, P 385); Oxytrichia pelionel/a (LACKEY 1936, P 269); Oxytrichia pellionella
(BALDENSPERGER 1927, P 295); Tachysoma pelionella (HORVATH 1950, P 159; TIRJAKovA
1992b, p 294); Tachysoma pelionellum (LARSEN 1992, P 301; TIRJAKovA 1991, p 868);
Tachysoma petlionella (ALEKPEROV 1986, P 57).
Tachysoma pellionellum is one of the most common oxytrichid hypotrichs in fresh-
water, which should, however, not be confused with the also very common and thus often
associated holostichid, Holosticha pullaster (MDLLER, 1773) FOISSNER, BLATTERER,
BERGER & KOHMANN, 1991 (contractile vacuole distinctly behind mid-body; midventral
 436 SYSTEMATIC SECTION

o
•
o
OP

h k

Fig. 13Sh-o Tachysoma pellionellum (from HEMBERGER 1982. h-o, protargol impregnation). Morphogenesis
in ventral view, h = 90 ~m. The most important event is the formation of so-called primary primordia (I). Ar-
rows in (0) mark primordia of dorsal kinety 6. See also text. OP = oral primordium; p 433.

row, that is, many more cirri on the ventral surface; more transverse cirri; short dorsal
cilia).
BERECZKY (1977a; Fig. 135r) illustrated rather short dorsal cilia, so that the identifica-
tion is questionable. The HORVAru (1933; Fig. 45a, b) population is classified as Oxytri-
cha tonga and that OfWRZESNIOWSKI (1861; Fig. 57d) as Oxytricha simi/is. The redescrip-
tions of the following authors are insufficient: ALAoRo-LUBEL et al. (1990; Fig. 241d),
BALDENSPERGER (1927), BELTRAN (1928; Fig. 228c), BIERNACKA (1959; Fig. 225t), BROD-
 Tachysoma 437

Fig. 135p-s Tachysoma pellionellum (p, from BERGER et aI. 1997; q, from BORGER 1908; r, from BERECZKY
1977a; s, from MAMAEvA 1979b. p-s, from life). p) Ventral view of a freely motile (not squeezed) specimen
from a pond (Leopoldskroner Weiher) in the city of Salzburg, Austria; note the curved outline, documenting
that T. pel/ionellum is very flexible. Arrow marks right transverse cirrus. q-s) Ventral views, q = 7S lun, r = 90
11m, s = 70 11m. Arrows in (q) mark the "Fettkom" in anterior and posterior portion of cell, respectively. CV =
contractile vacuole in mid-body, Ma = anterior macronuclear nodule; p 433.

SKY & YANKOWSKAYA (1929; Fig. 226t), BUISAN (1944; Fig. 233d), CONN (1905; Fig.
225f, 1), DUJARDIN (1841), EDMONDSON (1906; Fig. 234i), FEllERS & AlliSON (1920; Fig.
225r), FERNANDEZ-LEBORANS & CASTRO DE ZALDUMBIDE (1985), KOFFMAN (1926), LEPINIS
et al. 1973 (Fig. 225j), LEPSI (1957; Fig. 225e), QUENNERSTEDT (1865; Fig. 225n), ROSA
(1957a, b; Fig. 233a, c), SMITH (1978; Fig. 225m), SMITH (1914; Fig. 233g), SAMANO &
SoKOLOFF (1931; Fig. 225g), SOKOLOFF & ANCONA (1937; Fig. 234j), TCHANG et al. (1984;
Fig. 232j, k), WANG (1925; Fig. 242d). The illustration in MAnoNI (1981) is identical with
that by BICK (1972a).

M 0 r p hoi 0 g y and b i 0 log y: In life 55-100 x 15-30 Jlm, usually about

65-85 Jlm long. Long elliptical or even with parallel margins. Both ends rounded. Anteri-
orly somewhat more flattened than posteriorly (Fig. 135b-d, 135p, 136a-<:). Slightly con-
tractile. Macronuclear nodules ellipsoidal, in life about 10-12 x 5--7 Jlm, after protargol
impregnation 8-12 x 4-7 Jlm, with many small and large nucleoli. Invariably 1, in life
about 3-4 Jlm, after protargol impregnation 2.2-2.7 x 1.8-2.6 Jlm sized, globular micro-
nucleus between the two closely spaced macronuclear nodules; micronucleus in life con-
spicuously shining, thus it can sometimes be seen already at middle magnification. STEIN
(1859b) and Roux (1901) draw two micronuclei, which must, however, be interpreted as
438 SYSTEMATIC SECTION

:#f,I
;

"",

•
...
c
a d

Fig.136a-e Tachysomapellionellum (from FOISSNER & DIDIER 1981. a-c, from life; d, e, protargol impregna-
tion). a-c) Ventral, dorsal, and lateral view, a = 70 ~m, b, c = 85 ~m. d, e) Ventral and dorsal infraciliature, d
= 50 ~m, e = 40 ~m. The specimen in (d) has 3 pretransverse ventral cirri; p 433.

misobservation. Especially the illustrations by STEIN (1859b) unambiguously show T. pel-

lionel/urn (Fig. 135b-d). Contractile vacuole without collecting canals, situated at body
margin so that pellicle sometimes arched. Cortical granules absent (see also ScHNEIDER
1930). Cytoplasm with some small granules and 2-5 Jl.m sized, yellowishly shining
spherical inclusions. Usually a very conspicuous, about 5 Jl.m sized, greasily shining glob-
ule that looks like a ring with a thick (about I Jlm) wall (in German, "Fettkom"), in both
ends of the cell (Fig. 135b-d, q, 136i, m); these globules often surrounded by numerous
small granules which appear dark at bright field illumination; thus, middle portion of cell
usually looks bright! Moves hastily to and fro, body distinctly curved (Fig. 135p); for-
ward motion in numerous short, jerky movements.
Adoral zone of membranelles in life 114-1/3 of body length. Buccal area flat and
rather narrow. According to FOISSNER & DIDIER (1981) both undulating membranes con-
sist of two rows of basal bodies each; according to FERNANDEZ-LEBORANS (1983, 1985) the
paroral is made up of one row only. Frontal cirri slightly, transverse cirri strongly en-
larged, bases of remaining cirri consist of2 rows of basal bodies only. Base of buccal cir-
rus rather small. Anterior-most frontoventral cirrus (VI/4) somewhat separated from the
three other frontoventral cirri, which are invariably arranged as shown in Figures 135f,
 Tachysoma 439

Fig. 136C-j Tachysomapellionellum (f, g, from GROLIERE 1975; h, from BICK 1972a; i, from KAHL 1932;j, af-
ter OWUlEZ 1981. f, g, protargol impregnation; h-j, from life). C, g) Ventral and dorsal infraciliature, 65 ~m.
b-j) Ventral views, h = 70 ~m, i = 80 /lm, j = 62 /lm. Only drawing (i) shows the typical "FettkOmer", con-
spicuous ring-like structures in anterior and posterior portion of cell; p 433.

136d, w. Postoral ventral cirri arranged nearly in line, base of anterior-most cirrus orien-
tated longitudinally. Usually 2, sometimes 3 (mean = 2.4; n = 12) pretransverse ventral
cirri. Transverse cirri very prominent, in life about 15-20 J.lm long, posteriorly often
fringed and curved to the left (Fig. 135b-d, i, t). Marginal rows with roughly same num-
ber of cirri. Right one begins at about level of posterior-most frontoventral cirrus (Fig.
135f, 136d); marginal cirri in life 8-10 J.lm long. Dorsal cilia anteriorly 8-10 J.lm, posteri-
orly up to 15 J.lm long, always spread and thus easily recognisable even at low magnifica-
tion (lOOx; use differential interference contrast optics). Dorsal kineties 1 and 2 as long
as body, kineties 3 and 4 slightly shortened anteriorly, kinety 6 terminates in about mid-
body (Fig. 135g, 136e); arrangement of kineties especially in anterior and posterior por-
tion of cell difficult to discern.
 440 SYSTEMATIC SECTION

Fig. 136k-u Tachysoma pellionellum (k, I, after STOKES 1887b; m, from WANG & NIE 1935; n, from CURDS
1969; 0, after STERKI 1878; p, after JIRovEc et at. 1953 from SLAoECEK 1963; q, after STERKI 1878; r, after Cu-
PAR/mE & LACHMANN 1858; s, after Roux 1901; t, afterGAlEWSIWA 1933; U, from CHORIK 1968. k-u, from life).
k-n, p-u) Ventral views, k = 106 11m, I = 55 11m, m = 75 11m, n = 62 11m, p-r = size not indicated, s = 100 11m,
t = 90 11m, u = 85 11m. 0) Left lateral view; p 433.
 Tachysoma 441

,
\
, 'f\/"
\(/ /' ,
j'j
--... /""
./
.~,
--'
/

w x
Fig. 136v-x Tachysoma pellionellum (from SHIN & KIM 1993b. v, from life?; w, x, protargol impregnation).
v) Ventral view, 85 11m. w, x) Ventral and dorsal infraciliature, 90 Ilm. Note the prominent transverse cirri and
the single micronucleus between the two macronuclear nodules; p 433.

AUERBACH (1854) and PROWAZEK (1899a) described a cyst about 23 J.l.m in diameter.
Cysts subjected to very low oxygen tensions were found to excyst (STOUT 1954a). How-
ever, Tachysoma pellionel/um was never reliably recorded from terrestrial habitats,
strongly indicating that it does not make cysts. Accordingly, these old observations have
to be verified on reliably determined populations.
Tachysoma pe/lionel/um has 1.2 x 0.6 J.1m sized mitochondria with several cords,
each consisting of four tubules (WILBERT & HELLER 1971).
M 0 r p hog e n e sis of cell division is described by FERNANDEZ-LEBORANS & CAS-
TRO DE ZALDUMBIDE (1983) and, in more detail, by HEMBERGER (1982, Fig. 135h-o). It
commences with the proliferation of basal bodies near the left transverse cirrus (Fig.
135h). The oral primordium moves anteriorly into the area of the postoral ventral cirri
(Fig. 135i). These cirri are involved in primordia formation and a narrow streak origi-
nates from the oral primordium in the direction of the buccal vertex. This streak divides
into 3 long primordia, which are arranged in parallel to 2 further streaks right of them.
Thus, in the middle stage of morphogenesis 5 long, so-called primary primordia are pre-
sent (Fig. 135j, k). Posteriorly they are connected by an oblique, loosely arranged group
442 SYSTEMATIC SECTION

Table 25 Morphometric data of Tachysoma dragescoi (dra, from SONG & WILBERT 1997), T. granuliferum
(gra, from BERGER & FOISSNER 1987a), T. humicola humicola (hum, from FOISSNER 1984), T. humicola longise-
tum (Ion, from FOISSNER 1998), T. ovala (ova, from SONG & WILBERT 1997), T. pellionellum (pel, from FOIss..
NER & DIDIER 1981; pe2, from SHIN & KIM 1993b), and T. lerricola (ter, from HEMBERGER 1985). All data are
based on protargol-impregnated specimens. All measurements in micrometres. ? = sample size not indicated; it
only I value is known it is listed in the mean column, if2 values are available they are listed as Min and Max.
CV = coefficient of variation (in %), Max = maximum value, mean = arithmetic mean, Min = minimum value,
n = sample size, SD = standard deviation

Character Species mean SD CV Min Max n

Body, length dra 35.8 3.8 10.6 30 42 16
gra 62.6 7.0 11.2 55 76 II
hum 42.5 4.3 10.0 38 50 II
Ion 44.0 3.9 8.8 37 50 13
ova 38.3 3.8 9.9 32 45 16
pel 52.4 5.1 9.7 45 63 12
pe2 87.5 6.9 7.9 79 100 8
ter 90.0 70 100 ?
Body, width dra 21.1 2.8 13.3 16 26 16
gra 23.6 2.3 9.8 21 27 II
hum 21.0 2.6 12.4 18 27 II
Ion 21.2 2.4 11.2 18 27 13
ova 28.6 3.0 10.5 24 34 16
pel 17.1 2.2 8.8 15 19 12
pe2 38.9 3.4 8.8 35 45 7
ter 30.0 28 40 ?
Macronuclear nodule, length dra 5 6 5
gra 12.4 1.6 13.2 10 14 11
hum 7.8 1.8 22.9 6 10 II
Ion 7.2 1.3 18.8 6 10 13
ova 6.9 0.9 13.4 6 9 16
pel 9.2 1.5 16.4 8 12 12
pe2 13.1 1.8 13.5 II 15 7
Macronuclear nodule, width gra 5.4 1.0 19.1 4 7 II
hum 5.7 0.4 6.3 6 6 II
Ion 5.4 0.7 14.3 4 7 13
pel 5.3 0.9 17.6 4 7 12
pe2 5.9 0.7 II.8 5 7 7
Adoral zone of membranelIes, length dra 13.4 l.l 8.1 12 15 16
gra 21.7 1.4 6.2 20 24 II
hum 14.0 0.9 6.4 13 16 11
Ion 14.6 0.7 5.3 13 15 13
ova 15.6 l.S 9.6 13 18 16
pel 19.5 2.1 11.0 17 25 12
pe2 26.8 3.6 13.4 20 32 8
Adoral membranelIes, number dra 13.7 1.0 6.9 12 15 16
gra 22.7 1.4 6.0 21 25 II
hum 14.8 0.6 4.1 14 16 II
Ion 15.0 0 0 15 15 13
ova 15.6 1.0 6.2 14 17 10
 Tachysoma 443

Table 25 Continued

Character Species mean SO CV Min Max n

Adoral membrane\les, number pel 20.5 1.0 5.1 18 22 12
pe2 20.6 1.1 5.1 19 22 8
ter 16.0 ?
Left marginal row, number of cirri dra 7.6 0.7 9.6 7 9 16
gra 14.7 0.8 5.3 13 16 II
hum 7.2 0.4 5.6 7 8 II
Ion 7.1 7 8 13
ova 15.1 0.8 5.1 14 16 16
pel 13.7 1.8 23.6 9 16 12
pe2 16.0 0.5 3.3 15 17 8
ter 12 14 ?
Right marginal row, number of cirri dra 6.3 1.0 16.1 5 8 16
gra 16.3 1.1 6.8 15 18 II
hum 6.5 0.7 10.7 6 8 11
Ion 6.0 0 0 6 6 13
ova 15.4 1.3 8.5 13 17 16
pel 14.0 1.1 7.7 12 16 12
pe2 15.5 0.9 6.0 14 17 8
ter 11 14 ?
Dorsal kineties, number dra 3.0 0 0 3 3 16
gra 4.0 0 0 4 4 11
hum 4.0 0 0 4 4 11
Ion 4.0 0 0 4 4 13
ova 3.0 0 0 3 3 16
pel 6.0 0 0 6 6 12
pe2 6.0 0.5 8.3 5 7 9
ter 4.0 ?

of basal bodies which fonn the primordium of the undulating membranes of the opisthe
(Fig. 1351). In the next stage, these primary primordia divide at about the level of the buc-
cal vertex, similarly to in Urosoma emarginata (Fig. 1270). Now, 6 secondary primordia
(including the primordium of the undulating membranes and the left frontal cirrus) are
present in both the proter and the opisthe. Dorsal morphogenesis is not figured by HEM-
BERGER (1982). However, he mentioned that an anterior and posterior primordium oc-
curred in dorsal kineties 1-3. Unlike in other typical oxytrichids, not only the primordia
of dorsal kinety 3 divided, but also those of dorsal kinety 2 (Fig. 24d). Thus, the parental
dorsal kineties 1-3 fonn the new dorsal kineties 1-5 of the proter and opisthe. Kinety 6 is
a dorsomarginal row (Fig. 1350, arrows).

o c cur r e n c e and e colo g y: One of the most common hypotrichs in the auf-
wuchs and detritus of freshwater habitats such as running waters, lakes, ponds, peat-bogs,
springs, and infusions; sometimes abundant (EHRENBERG 1838, FINLAY et al. 1993, KAHL
1932, STEIN 1859b, own observations). Very likely oligo-euryhaline (up to 4 %0 salinity)
444 SYSTEMATIC SECTION

with freshwater-optimum (ALBRECHT 1983, 1984). According to WILBERT (1969) and

NUSCH (1970) more abundant during spring and autumn than in the remaining seasons.
However, SCHONBORN (1982, 1984) found highest abundances (110 indo cm-2) in February
and observed a correlation with high population densities of diatoms; BICK (l972a) found
T. pellione/lum frequently also in layers ofnaviculid diatoms, and REuss (1976) observed
maximal abundances during spring and summer. These deviating observations indicate
that no season is preferred (KALMus 1928). Often associated with the likewise very com-
mon Holosticha pu/laster with which it may be confused at superficial observation (see
taxonomy). Cosmopolitan.
Locus classicus is Denmark, where MOLLER (1773) discovered T. pellionellum in a
plant infusion. Records largely substantiated by illustrations: very common in lotic and
lentic waters in Austria (own observations); submerged Sphagnum (PH 4) in Belgium
(CHARoEZ 1981); Sava River, Croatia (PruMC-HABDUA et al. 1996); Czechoslovakia
(JiROVEC et al. 1953); peat-pit in France (GROLrERE 1975); pond-like section of moderately
organically polluted brook with filamentous algae near Besse-en-Chandesse, France
(ForSSNER & DIDIER 1981); about 20 indo cm-2 in the aufwuchs ofmesosaprobic brooks in
Germany (PATSCH 1974); common in infusions and stagnant waters in Berlin, Germany
(EHRENBERG 1838); peat-pit and ponds near Berlin, Germany (locus classicus of the syno-
nym Stylonychia echinata; CLAPAREDE & LACHMANN 1858); plankton of the mesosaprobic
Danube River in Hungary (BERECZKY 1977a, b, 1979, NOSEK & BERECZKY 1981); littoral
and profundal of Lake Geneva, and common in lentic waters in Switzerland (ANDRE
1914, 1916, FOREL 1904, Roux 1900, 1901); Volga River (MAMAEVA 1979a-c); at 3-19
°C among algae (Tetraspora, Draparnaldia) in the littoral of Lake Baikal (GAJEVSKAJA
1927, GAJEWSKAJA 1933, ROSSOLIMO 1923); USSR (CHORIK 1968); pond in China (WANG
& NIE 1935); Han River in Seoul, Korea (SHIN & KIM 1993a, b); freshwater near
Trenton?, New Jersey, USA (locus classicus of the synonyms T. agile and T. mirabile;
STOKES 1887b); Chile (BORGER 1908).
Records from freshwater habitats of Europe and Asia not documented by illustra-
tions: small stagnant and running waters in the Austrian Alps (ForSSNER 1979a, 1980c,
FOISSNER & ADAM 1979); Enns River, Austria (MEISRIEMLER & RrnoL 1985); beta- to al-
phamesosaprobic brook in Salzburg, Austria (HAsLAUER & PICHLER 1979); mesosapro-
bic Traun River and Danube River, Austria (ForSSNER & MOOG 1992, KALTENBACH 1960,
SCHIEDERMAYR 1887); very abundant in ponds in western Austria (DALLA TORRE 1891);
pelagial of groundwater ponds in Styria, Austria (KRAlNER 1988); hypertrophic pond at
Salzburg University, Austria (BLATTERER 1989); betamesosaprobic rivers in Upper Aus-
tria (for example, AOOLR 1994b, own observations); drinking water of the town ofLinz,
Upper Austria (SCHIEDERMAYR 1882; further records from Austria: ForSSNER & ForSSNER
1988, HELD 1937, SCHMARDA 1846, STORCH 1869); abundant in the aufwuchs of oli-
gosaprobic to betamesosaprobic running waters in Belgium (AELTERMAN 1952; further
records from Belgium: BERVOETS 1940, CHARDEZ 1987, GILLARD 1950); running waters in
Bulgaria (DETCHEVA 1979b, 1981, 1986, 1991, 1993, DETCHEWA 1972, RussEv et al.
1984); springs in Prague, Czechoslovakia (VEJDOVSKY 1882); in the plankton of a fast
 Tachysoma 445

flowing (0.9 m sol) river in Czechoslovakia (PROWAZEK 1899a); abundant in the Mold-
avia River, Czechoslovakia, throughout the year (KALMus 1928); in 5 betamesosaprobic
and 2 alphamesosaprobic sites in waters of Czechoslovakia (SRAMEK-Hu~EK 1956b,
1957); saprobic brook and ponds in Czechoslovakia (BUCHAR 1957, MATIS 1973, SVEC
1897); submerged and wet mosses in Czechoslovakia (TIRJAKovA 1991, TIRJAKovA &
MATIS 1987a); spring water with suspended organic matter in Bratislava, Czechoslova-
kia (COHN 1875; further records from Czechoslovakia: HANu~KA 1962, HAsSDENTEUFE-
LOvA- MORAvcovA 1955, SLAoEl:EK& SLAoEl:KOvA 1974, SRAMEK-Hu~EK 1953, TIRJAKovA
1992b, 1993); Denmark (NIELSEN 1960); sand and mud flats from rivers near Plymouth,
England? (LACKEY & LACKEY 1963); near Perth, England (CRAIGIE 1921); slow sand fil-
ters at the Metropolitan Water Board, London, England (LLOYD 1973; see also CURDS
1979); rivers, ponds, ditches, and bogs in Estonia (JACOBSON 1928); near Lyon, France
(ORMANCY 1852); alphamesosaprobic to polysaprobic section of a French river (GRO-
LIERE et al. 1990; further record from France: MONIEZ 1889); backwash sludge from oxi-
dators and filters of waterworks in Germany (FOISSNER 1996a, GIERIG 1993); oligosapro-
bic to betamesosaprobic and betamesosaprobic to alphamesosaprobic areas of brooks
near Bonn, Germany (JUTRCZENKI 1982); abundant in the aufwuchs of oligosaprobic, oli-
gosaprobic to betamesosaprobic, and alphamesosaprobic reservoirs in Germany (NuSCH
1970); highly polluted Salzach River at the village of Laufen, Germany (BAUER 1983);
benthal of mesosaprobic brooks and rivers in Germany (BAUER 1987, FOISSNER et aI.
1992a, b, SCHONBORN 1981, 1982, 1996, ZIEMANN 1985); highly abundant, especially
during winter, in the aufwuchs of the betamesosaprobic Elbe River and in the aufwuchs
of Hamburg harbour, a highly stressed habitat in the freshwater section of the Elbe Estu-
ary, Germany (BARTSCH & HARTWIG 1984, GRIMM 1968, HECKMAN et al. 1990, TENT
1981); caves in Germany (GITILESON & HOOVER 1969, GRIEPENBURG 1934, 1935); cool-
ing system of a conventional power station at the beta- to alphamesosaprobic Main
River, Germany (BERNERTH 1982); clean brooks in Bavaria, Germany (FOISSNER 1997a);
Schussen River, Germany (WETZEL 1928a); very abundant in the aufwuchs of a
eutrophic lake and pond in Germany throughout the year (MOCKE 1979, SONG & WIL-
BERT 1989, WILBERT 1969); aufwuchs of oligotrophic, mesotrophic, and eu- to polytro-
phic EifeImaar lakes, Germany (PACKROFF 1992, PACKROFF & WILBERT 1991); pond with
leaf litter in Germany (BICK 1958); in the sludge of the influent ofa wastewater fishpond
in Munich, Germany (KAUFMANN 1958; further records from Germany: BAIER 1934, GRE-
ISER 1974, HENDERSON 1905, REuss et al. 1972, PACKROFF & ZWICK 1996, SCHMIDT 1916);
Greece (STEPHANIDES 1948); radioactive (200-450 cps) spring in Greenland (LARSEN
1992); Lake Balaton, Hungary (ENTZ 1897, FRANCE 1897); pond with iron bacteria in a
cave in Hungary (DUOICH 1932; further records from Hungary: BERECZKY 1995, BICWK
1955a, HAMAR 1971, STILLER 1942); Lake Como, Italy (CATTANEO 1882); alpine lakes
(1267 m and 1920 m above sea level) in Italy (MONTI 1904, 1905); well and spring in it-
aly (GRISPINI 1938, STELLA 1947); up to 24 indo cm-2 in mesosaprobic brooks in Italy
(MAnoNI 1980, 1983, 1984, MAnoNI & GHETTI 1977, 1981a; further records from Italy:
CALLONI 1890, CUNEO 1891, ENRIQUEs 1913); Latvian rivers (LIEPA 1990); various habi-
446 SYSTEMATIC SECTION

tats in the Netherlands (VERSCHAFFELT 1930); sporadically with low abundance in the
Sphaerotilus community of a Polish river (HUL 1986); pond near Warsaw, Poland, to-
gether with Stylonychia pustulata, Cyclidium glaucoma, and species of the Parame-
cium aurelia complex (WRZESNIOWSKIEGO 1866; further record from Poland: JAWOROWSKI
1893); Danube River in Romania (SPANDL 1926b); Henares River, Spain (FERNANDEZ-
LEBoRANs & NovILW 1996, FERNANDEZ-LEBORANS et al. 1985, FINLAyet al. 1993, MAR-
GALEF L6pEZ 1945); Danube River in Slovakia (SZENTIVANY & TIRJAKovA 1994); bogs
near Berne, Switzerland (SAKOWSKY-CAMPIONI 1906); alpine lakes in Switzerland (MEs-
SlKOMMER 1954, ZSCHOKKE 1900); profundal (35 m) of Lake Neuchatel, Switzerland
(MONARD 1920; further records from Switzerland: ANDRE 1912, 1915, 1926, BOURQUlN-
LINDT 1919, CDANN 1907, MERMoD 1914, MEsSlKOMMER 1948, PERTY 1852b); aufwuchs
from alphamesosaprobic Save River, Yugoslavia (pRIMC 1981, 1984); in the epiphyton
of more than 75 % of collected samples from karstic running waters, Yugoslavia (PRIMc-
HABDUA & HABDUA 1991); 30 indo cm-2 in the littoral of Lake Kinneret, Israel (MAnoNI
1990); stagnant water in India (BHATIA 1936); Lake Dzhandar, Azerbaijan? (ALlEV
1988); Araks River (ALlEV 1982a), Oka River (NEISWESTNOWA-SHADINA 1935, SASSUCHlN
1930, SASSUCHlN et al. 1927), and Volga River in the USSR (ZYKOFF 1903); plankton of
reservoirs in the USSR (ALEKPEROV 1980, MYLNIKOVA 1992; further records from the
USSR: ALEKPEROV 1984c, BELOVA 1988, DADAY E. 1904, DADAY J. 1903, EHRENBERG
1830, EICHWALD 1844, 1849, GORIDCHENKO 1984, LIEPA 1973, 1983, MERESCHKOWSKY
1877, MiNKEwrrSCH 1898, MOVCHAN & PROTASOV 1986, MYLNlKOVA 1981, 1993, OLEKSIV
1985, SHADlN et al. 1931, VEYLANDE & LlYEPA 1985, WEISSE 1845, 1848a, ZHARlKOV
1982, 1992, ZoLOTAREV 1987); sediment of a Chinese river (Gu et al. 1988; further re-
cords from China: MA 1994, as T. pe/linnel/a; MA & DANG 1994, NING et al. 1993, SHEN
& GONG 1989); Japan (EDMONDSON & KINGMAN 1913); Mongolia (DADAY 1906); stag-
nant waters in Tibetan plateau (WANG 1977).
America: Ice Lake (about 3700 m above sea level) and habitats near Greeley, Colo-
rado (EDMONDSON 1912; further records from Colorado: BEARDSLEY 1902, CoclcERELL
1907, HAMILTON 1943); Iowa (SHAWHAN et al. 1947); south-eastern Louisiana (BAMFORTH
1963); Ocqueoc River, Michigan (CAIRNS & YONGUE 1966); epilimnion of Douglas Lake,
Michigan (YONGUE & CAIRNS 1976); North Carolina (BEVEL 1938); Devil's Lake
complex, North Dakota (EDMONDSON 1920); among algae on submerged rock in clear wa-
ter in Mirror Lake, Ohio (STEHLE 1920); Oklahoma (GABEL 1927); Conestoga drainage
basin and pond of University of Pennsylvania (CAIRNS 1965a, WANG 1928); Mountain
Lake region, Virginia (BOVEE 1960); abundant in clear flowing water with abundant plant
life, USA? (HAUSMAN 1917); Fourth Lake, Canada (FANTHAM & PORTER 1948); in the
moss Catharinea undulata from the bank of a brook near Montreal, Canada (FANTHAM &
PORTER 1946); Mexico (L6PEZ-OCHOTERENA & ROURE-CANE 1970); Argentina (BUSTOS
1933, SECKT 1924); Brazil (CUNHA 1913, PINTO 1925); Venezuela (SCORZA & NuNEz
MONTIEL 1954).
Africa: pond in the bank of the Nile River, Egypt, together with two Euglena species
(SCHMARDA 1854).
 Tachysoma 447

Records from marine and brackish habitats and inland salt waters are not substanti-
ated by illustrations (for example, ANnRUSSOWA 1886, BORROR 1962, BOUTCffiNSKY 1895,
BUTSCffiNSKY 1897, CAREY 1992, DAGAJEVA 1930, FOWELL 1944, GALLIFORD & WILLIAMS
1948, LACKEY 1936, PARONA 1883, PATIERSON et al. 1989, REES 1884, SCHMARDA 1846,
1847, SMITH 1904, VERSCHAFFELT 1930, WILBERT 1995, ZACHARIAS 1888). FINLEY (1930)
noted that under experimental conditions a maximum of 1.4 % salt is tolerated. HAMMER
(1986) found an upper limit of7 % salinity. ALBRECHT (1986) and MrnAiLOWITSCH (1989)
recorded Tachysoma pellionellum from salt polluted running waters in Germany with
36-12763 mg 1-1 Cl- and 51-3590 mS m- I specific conductivity. KARL (1928a, b) did not
find it in saline freshwater habitats (Oldesloer SalzwasserstelIen) in areas with either low
or with high salt content.
Tachysoma pellionellum was never reliably recorded from terrestrial habitats (FOISS-
NER 1987a) indicating that it cannot make cysts; the numerous terrestrial records from all
over the world are thus very likely misidentifications. Probably it was confused with Oxy-
tricha setigera, which is common in terrestrial habitats and also has long dorsal cilia and
a single micronucleus (for example, BAMFORTH 1967, 1968, 1969, 1976, BICZOK 1955b,
1956, 1979, BRUNETTI & CARLETTI 1932, CHARDEZ 1967, CHAUDHURI 1929, CUNNINGHAM
& LOHNIS 1914, DECHEVA 1968, 1970, 1973, DETCHEVA 1972b, DIXON 1939, EHRENBERG
1849,1869, FANTHAM & PATERSON 1926, GEL'CER & GEPTNER 1976, GELLERT 1957, GRAN-
DORl & GRANDORl 1934, HEINIS 1937, HORVATH 1950, RINo & MOMIKI 1931, LoUSIER &
BAMFORTH 1990, MARTIN & SHARP 1983, NIKOlJUK & GELTZER 1972, NIKOLYUK 1956,
1965, ROSA 1962, ROSA & LHOTSKY 1971, SANDON 1927, SHEN et al. 1992, SHIBUYA 1927,
SMITH 1972, 1973a, 1974a-c, 1975, 1982, 1984, 1985, STOUT 1955, 1958, 1960, 1961,
1963, 1968, 1970, 1984, TIRJAKovA & MATIS 1987a, VARGA 1935a, b, 1936, 1953a, 1958,
1960). SMITH (1981) recorded Tachysoma pellionellum from a moss peat (PH 4.1, water
content about 400 % dry mass) on South Georgia, South Atlantic.
Tachysoma pellionellum is occasionally found in activated sludge (BANINA 1983,
1990, CURDS 1966b, 1969, 1975, CURDS & COCKBURN 1970a, ETTL 1996, HAMBURG-
EISENBERG 1933, KUTlKOWA 1984, MAnoNI 1988a, MAnoNI & GHETTI 1981b, MARco et al.
1991, SIMAKOV 1986), percolating filters (BEGER 1935, CURDs 1975, CURDs & CocKBURN
1970a, RICHARDS & CUTLER 1933) and the film of rotating biological contactors (MAnoNI
& GHETTI 1981b, MAnoNI et al. 1979). In activated sludge plants an effluent-BODs range
of 11 to 20 mg I-I is indicated (CURDS & COCKBURN 1970b). SCHERB (1968a) counted up to
600 indo ml- I activated sludge and stated that Tachysoma pellionellum occurs more fre-
quently in plants treating dairy wastes than in plants treating municipal wastes. STOUT
(1954b) found it in an oxidation pond of an experimental meat digestion plant in New
Zealand; the record from the anaerobic digester of the same plant, however, appears
doubtful. SMITH (1982) recorded Tachysoma pellionellum from the guano of a king pen-
guin colony from the sub-Antarctic island of South Georgia; the identification is, how-
ever, not substantiated by morphological data!
Feeds on bacteria, cyanobacteria, diatoms, autotrophic flagellates, green algae, fila-
mentous growth, detritus (BICK 1980, CURTIS & CURDS 1971, FINLAY et al. 1993, FOISSNER
448 SYSTEMATIC SECTION

Table 26 Autecological data of Tachysoma pe/lionellum. References: column 1, from BICK & KUNZE (1971;
summary of literature data); column 2, from BERECKY (1977a; Danube River in Hungary, sample size not indi-
cated); column 3, from DETCHEVA (1972a, 1975a-c, 1976b, 1978, 1979a, c, 1982b, 1983a, b, c; many analyses
from Bulgarian running waters); column 4, from MADaNI & GHETIl (198Ia; Stirone River in Northern Italy,
sample size not indicated); column 5, from FOISSNER et al. (1982a; 31 analyses from stagnate alpine water bod-
ies in Austria); columns 6, 7, from FOISSNER et al. (1991; 6 = total range from various running waters in
Austria, n = 18-35; 7 = only high abundances considered, n = 3)

Parameter' Reference
2 3 4 5 6 7
Saprobity o-a b,a o-blb-a bib-a
Frequency (%) 2.7-10.7 5 26 2.2
Temperature eC) 0-43 7-12.5 7-26 1.8-25 5-20 O.l-II 3-5
pH 6.4-9.1 7.7-8.1 6.3-8.2 7.6-8.4 4.8-5.8 7.0-8.6 7.1-8.2
02(mg I") 0-22.5 9.2-12.8 3.4-14.5 8.1-12.7 2.8-11.3 7.3-14.2 12-13
O2(% saturation) 40-100 33-131 62-117 98-103
BODs (mg I") 10.9-24.5 2-12.85 1.0-22 0.8-4.7 1.7-2.3
Total hardness (OdH) 1-7 3.2-13.7 9.4
KMnO.-consumption 0.6-69 13-244 12-54 2.5-24 11-19
NH/-N (mg 1-') 0-3.9 0.52-0.75 0.008-155 0-0.8 <0.01-1.5 0.01-0.05
N02--N (mg )") 0.003-0.055 0-0.6 0.02-0.04 0 0-0.02 0-0.02
NO)--N (mg I-i) 0.47-1.65 0-9 0.6-3.2 0-0.2 0.2-3 0.4-1.2
PO/'-P (mg I") 0.16-0.38 0-0.26 0-0.25 0.02-0.05
Bacteria mI" (x Hf) 0.03-J02 <0.0045 3 <0.0043

, The figure "0" denotes only that the parameter in question could not be detected with the analytical method
used.
2 Direct counting.
3 Plate method, 22°C, 48 h.

• COD.
S BERECZKY (1977a) designated it as "02(Verbrauch)". Probably she meant the BODs.

1980c, FOISSNER & DIDIER 1981, MAnoNI 1981, NOLAND 1925, RIGGENBACH 1922). One
specimen consumes 48-84 diatoms d- I and about 7200 bacteria d- I , corresponding to a
food volume of 48 900-63100 !lm3 ind: 1 d- I (SCHONBORN 1981, 1982). Tachysorna pe/-
lionel/urn itself is only sometimes consumed by the oligochaete Chaetogaster diastro-
phus (SCHONBORN 1984). Generation time and production data of Tachysorna pel/ione/-
/urn under natural conditions in running waters are summarised in Table 28. SCHONBORN
(1981, 1982, 1992) estimated 50-82 generations per year in running waters in Germany.
Biomass of 106 individuals 12-15 mg (FOISSNER et al. 1991) to 34 mg (SCHONBORN 1981,
1982).
Some autecological data are shown in Tables 26, 27 and Figure 137. Further aut-
ecological data: scattered in a betamesosaprobic brook in Czechoslovakia at 8.5 °C,
pH 7.6, 9.8 mg I-I O2, 80 % O2 saturation, 5.5 mg 1-1 BODs (VASICEK 1964); 7-20°C,
pH 6.tH>.9, 1.2-2.6 mg 1-1 O2 (MATIS 1975); 7-46.5 °C, pH 5.5-7.8 (MATIS & STRAKovA-
-STRIESKovA 1991); 0.3-7.2 mg I-I DOC (STOSSEL 1979); 71-1097!lS cm- I (20°C; n =
 Tachysoma 449

Table 27 Autecological data of Tachysoma pellionellum (columns 1-3), Gonostomum affine (column 4), and
Pleurotricha lanceolata (column 5). References: columns 1,2, from SCHMERENBECK (1975; experimental run-
ning waters; I = total range, 2 = optimal range); column 3, from MIHAILOWITSCH (1989; 68-76 analyses from
salt polluted ditches in Germany); column 4, from DETCHEVA (19723, 1975, 1983c; Bulgarian running waters,
sample sizes not indicated); column 5, from PATRICK et al. (1967; 4 records from the Savannah River, USA)

Parameterl Reference
2 3 4 5
Saprobity 2.3-3.7 2.3-3.0 ola
Frequency (%) 0.87-2.1
Temperature (0C) 1.2-20.5 19.8-21 >8-20
pH 4.9-8.5 6.7-8.3 6.7-8 7.5-7.8
O2 (mg I-I) 1.2-9.5 6.5-9.5 1.2-17.6 5.6-9 7.0-<11
O2 (% saturation) 62-92
BODs (mg I-I) 0.4-41 0.4-2.6 I-IS 0.5-<3
KMn04-consumption 7.2-38
NH/-N (mg 1. 1) 0-27 0-5.4 0.04-7.3 0.08-1 <0.0093
NOi-N (mg 1. 1) 0-67 0-3 0.01-0.8 0.94-2 <0.007
NOi-N (mg 1.1) 10-99 0.38-13.4 0.11-3.3 >O.05-{).2
PO/'-P (mg 1"1) >0.003-0.017
Bacteria ml-I (x 1(6) 1.2-2042 1.2-902

I The figure "0" denotes only that the parameter in question could not be detected with the analytical method
used.
2 Direct counting.
) NHrN.

32),0.9-91 mg I-I CI- (n = 16), 5-45 mg I-I SOl (n = 16), <0.01-0.25 mg 1"1 Ptotal (n =
32),5-19 mg I-I COD (n = 12), <10-13 000 endotypical Coli per 100 ml (44°C; n = 26;
FOISSNER et al. 1991); pH 7.4, 8.2 mg 1"1 O2 (80 % saturation), 45-57 mg I-I BODs,
5.5 mg 1-1 NH3, 92 mg 1"1 nitrate and nitrite (JUSTIN 1960). STOUT (1956) found a survival
time of 113 h at low oxygen levels (identification uncertain because "Chlorella" are pre-
sent). REINNARTH (1979) recorded T. pellionellum in the sediment from -0.5 to -4 cm at a
redox potential of +153 to -229 mY, pH 6.8-7.7,3.2-15.0 mg 1"1 NH/-N, 0.9-2.5 mg 1-1
N03--N. SCHMERENBECK (1975) found, under experimental conditions, that T. pellionellum
is highly abundant at flowing velocities of 0.3-0.6 m S-I, above all when mineralisation is
completed; dominant in the aufwuchs community especially at high current velocities and
high abundances of diatoms. Eurythermic (Tables 26, 27; BICK & BERTRAM 1973, IssEL
1901, NISBET 1984, PRIMC 1983), however, according to WANG (1928) and MONCH (1970)
low temperatures (up to 15°C) were preferred Tachysoma pellionellum survived a direct
transfer from 6 to 43°C and from 43 to 25 °C (SCHARF 1961). Lowest pH = 4.0 (BICK &
DREWS 1973). BICK (1957) and REuss (1976) found it in polluted water bodies with up to
1.6 mg I-I hydrogen sulphide; however, the specimens were only 2.7 times as long as
broad (usually about 3.5:1), indicating that these conditions are not favourable. Accord-
450 SYSTEMA TIC SECTION

Table 28 Annual values of the abundance, the generation time, and the production of some populations ofhy-
potrichous ciliates from two running waters in Germany (from SCHONBORN 1981, 1982). Species: column 1 =
Eup/otes affinis, column 2 = E. charon, column 3 = E. patella, column 4 = Stylonychia mytilus complex, col-
umn 5 = Tachysoma pellionellum, column 6 = Uroleptus piscis

Parameter Species
2 3 4 5 6
Mean abundance 0.3 l.l 1.2 9.9 0.2
(ind. cm·2), 2.4 0.08 0.1 13.9
Mean abundance 134 1058 1648 31811 419
(ind. m·2)' 24800 800 1700 139000
Mean generation time (h)' 84.0 97.5 118.5 95.4 72.0
88.0 72.0 258.0 106.2
Minimum generation 24 24 24 24 24
time (h)' 72 72 120 48
Maximum generation 120 120 360 240 120
time (h)' 88 72 258 106
Annual production 80 274 126 132 23
(ind. cm-2 y')' 386 23 10 3700
Annual production 9909 81521 476500 8200000 136400
(ind. m-2 y')' 3800000 200000 99000 37000000
Mean biomass 0_0 0.52 0.71 1.06 0_19
(mgm-2)' 0.79 0_06 0.70 4.84
Annual production 0.3 40_9 191.0 278 66.3
(mgm-2y')' 121.6 17.7 39.8 1321
PIB ratio (a)' 75 79 269 262 348
153 295 57 273
Food consumption 32 842 736 123
(mgm-2 y') 377 34 1021 10302

, Upper line: Ammerbach, a brook near the village of Jena, Germany. Mean width = 163 em, mean depth =
7.5 cm; current velocity at mean water = 0.55 m s·'; mean water (MQ) = 34 I s-" annual mean of water tem-
perature = +8.5 °C (minimum temperature about 4°C, maximum temperature about 14°C), 02-saturation =
80-90 %, saprobic index according to ciliate community = 2.6. Lower line: Saale River near the village of
Rothenstein, Germany. Width about 55 m, mean depth = 0.75 m, current velocity at MQ = 0.70 m sol, MQ
about 29 m3 s-', minimum water temperature about 0 °C, maximum water temperature about 17°C, O2 =
8-10 mg 1-', BODs <6 mg 1-', saprobic index = 2_5.

ing to WETZEL (l928b) H2S is not tolerated. BIcz6K (1955a) studied the effect of root ex-
tracts of various plants on T. pellionellum. For the effect of carbonic acid, see NIKlTINSKY
& MUDREZOWA-WYSS (1930).
The saprobic classification of T. pellionellum is vel)' diverse; it varies from beta-
mesosaprobic (CURDS 1966b, FOISSNER 1979b, SRAMEK-HusEK 1958) over alphameso-
saprobic (for example, KOLKWITZ 1950, KOLKWITZ & MARSSON 1902, 1909, MARSSON
1903, MAUCH 1976) to alphamesosaprobic to polysaprobic (REuss & FRIEDRICH 1978,
SLAoECEK 1969, SLAoECEK et al. 1981). The latter authors proposed the following sapro-
bic valencies: a-p; 0 = 1, b = 2, a = 3, p = 4, I = 1, SI = 3.0. This classification, especially
 Tachysoma 451

Fig. 137 Tachysoma pel/ionel/urn

to.
'~ l-T"""'!''"''I-~-,II'''~•••
(from HEUSS 1976). Distribution ex-
pressed as percentage of n records in
the spectra of some physical and
chemical factors. I = temperature, 2
= pH, 3 = specific conductivity, 4 = I'

O2, 5 = O2 saturation, 6 = BODs, 7 = "

KMn04-consumption, 8 = CI", 9 =
POl-,lO=NH':.

the strong emphasis of the

polysaprobity, does not match
the autecological data (Tables
26, 27, Fig. 137) and most of
the numerous records. How-
ever, there is no doubt that T.
pellionellum has a rather n. "7
6
wide saprobic range, which is
indicated by records such as
those by CURTIS (1969) and
".1811
CURTIS & CURDS (1971), who 7
found it rather frequently in
the slime of sewage fungus,
and KOlIN (1940), who re-
corded it from springs. REuss
(1976) found it predomi-
nantly at BODs values of
<9 mg I-I (Fig_ 137). I found
T. pellionellum mainly in run-
ning waters with saprobic in-
dices from 1.7-2.6 (mean =
2.2; n = 34), and 2.2-2.5 (n =
3) if only records with high abundances are considered_ Thus, ForssNER et al. (1991) sug-
gested the following distribution: b-a; 0 = 1, b = 4, a = 4, P = 1, I =1, SI = 2.5. This
agrees largely with the classification by FRIEDRICH (1990; see also DIN 1990 and BERGER
et al. 1997): b = 3, b-a = 12, a = 3, I = 8, SI = 2.1. The distributions proposed by BICK
(l972a; b = 5, a = 5, i = 3) and MORAvcovA (1977; b = 3, a = 7, 1=4, SI = 2.6) seem to
stenosaprobic.

Tachysoma chilensis (BORGER, 1905) comb. nov. (Fig. 138a)

1905 Oxytricha pellionella O. F. MOLL (var. chilensis) BORGER, An. Univ. Chile, 117: 436.
452 SYSTEMATIC SECTION

Nom e n c I a t u rea n d t a x 0 nom y: The rank of the name

chilensis is sub-specific because it was published before 1961 (Article
45 g, IcZN 1985). Since a change in rank of the species group does not
affect the authorship of the name of the taxon, BORGER (1905) is the
author of this species (Article 50 c[i]). BORGER (1905) expressly men-
tioned that this large and flexible form has no caudal cirri; I thus trans-
fer it to Tachysoma. Although some characters are not described (num-
ber of micronuclei, probably two or more because the macronuclear
nodules are widely separated; dorsal cilia, probably short since BORGER
did not draw them) it should be possible to identity this rather large
species. Detailed redescription necessary.

M 0 r p hoI 0 g y: In life about 200 x 64 Ilm. Left body margin con-

vex, right concave, both ends rounded. Contractile. Cytoplasm anteri-
Fig. 138a Tachy- orly and posteriorly densely filled with small, greasily shining
soma chilensis granules, right part with greasy globules. Adoral zone of membranelles
(after BORGER about 35 % of body length. Dorsal cilia very likely short (see taxon-
1905). Ventral omy).
view from life,
200 11m; p 451.
o c cur r e n c e: Locus classicus is an irrigation canal in Santiago,
Chile. Also recorded by CARBONELL (1935; as Oxytricha chilensis BUR-
GER) in the River plate, Argentina.

Tachysoma humicola GELLERT, 1957 (Fig. 138b-e, 139a-f, Table 25)

1957 Tachysoma humicola GELLERT, AnnIs Inst. bioI. Tihany, 24: 20 (Fig. 1391).
1984 Tachysoma humicola GELLERT, 1957 - FOISSNER, Stapfia, 12: 121 (Fig. 139a-e; authoritative redescrip-
tion; slides of neotype specimens are deposited in the OberOsterreichische Landesmuseum in Linz, Up-
per Austria).
1998 Tachysoma humicola longisetum FOISSNER, Europ. 1. Protisto!., 34: 226 (slides of type specimens are
deposited in the Oberosterreichische Landesmuseum in Linz, Upper Austria).

T a x 0 nom y: The synonymisation of this terrestrial form with T. bicirratum as sug-

gested by BORROR (1972a) is unjustified because T. bicirratum lives in the sapropel of
freshwater and the posterior-most marginal cirri are enlarged and elongated. The rather
superficially described limnetic Oxytricha saltans VUXANOVICI is somewhat reminiscent
of the present species (Fig. 2320).
ForSSNER (1998) described the subspecies Tachysoma humicola longisetum, which
differs from T. humicola humicola almost exclusively in that the posterior-most dorsal
cilia are distinctly elongated (cp Fig. 138c, e with Fig. 13ge). Thus, the following de-
scription matches both taxa unless otherwise indicated.
 \. /
/'" \ : ,/

/(/>\/ /
~
/'" '\,
(\ \ (/\ / , \ ,. /
.~ ,
~\ ,x
# ,/
'0 \
/1-~
1--.\ / \
I # ---.,
~I \ / :/
y / #
/\ /
~
(':>
~ :- -[
i:\ -"J c
~ # ::!
~ ~ I:l
~ / \! .# •
~ ."
c~ "
• ,I Y. " .;(7
c
-<tl\ • d
><~\ e
Fig. 138b-e Tachysoma humicola longisetum (from FOlsSNER 1998. b, c, in life; d, e, protargol impregnation). b) Ventral view, about 40 /-lm. c) Dorsal view. Large ar-
row marks elongated, posterior-most dorsal cilia; small arrows denote an inconspicuous but rather invariably (see Fig. 138e) discontinuity in dorsal kinety 3. d, e) Ven-
tral and dorsal infraciliature and nuclear apparatus, 44 /-lm. Arrowhead in (d) denotes posterior-most cirrus of right marginal row. Note single micronucleus and promi-
nent transverse cirri; p 452.
.j:>,
v-
I.»
 454 SYSTEMATIC SECTION

--..
- --
.* ......
f/~
))
/
• #::/ S I I
If ~ ;::
)
• !', S )
f'S

,'~
~
,/!'
i I

~ • I )
~
~

I I
,•
~

~,
)
//

Fig. 139a-( Tachysoma humicola humicola (a-e, from FOISSNER 1984; t: after GELLERT 1957. IH:, from life;
d, e, protargol impregnation; f, sublimate fIXation and opaiblue staining after BRESSLAU). a) Ventral view, 52
11m. b) Dorsal view of a backward moving specimen. c) Right lateral view. d-f) Ventral and dorsal infracilia-
ture, d, e = 40 11m, f= 60 11m. Arrow in (d) marks right frontal cirrus, arrowhead points at posterior-most cirrus
of right marginal row; p 452. g Tachysoma balatonica (after GELLERT & TAMAs 1958). Ventral infraciliature,
opalblue staining after BRESSLAU, 150 11m; p 456. h Tachysoma bicirratum (from KMn.. 1932). Ventral view
from life, 60--90 11m; p 457. i Tachysoma terricola (from HEMBERGER 1985). Ventral infraciliature and nuclear
apparatus after protargol impregnation, 90 11m; p 458.
 Tachysoma 455

M 0 r p hoi 0 g y and b i 0 log y: Subspecies T. humicola humicola in life about

45--60 x 15-20 ~m (FOISSNER 1984), according to GELLERT (1957) 50--60 ~m long, sub-
species longiseatum in life about 40 x 20 ~m (FOISSNER 1998). Body margins in life usu-
ally parallel, after protargol impregnation, however, always broadly oval, that is, both
ends broadly rounded. Rather flexible, but not conspicuously contractile. Macronuclear
nodules usually slightly obliquely arranged about in mid-body, in life about 7 ~m across,
with many nuclei of various size. Usually 1, rarely 2 spherical micronuclei between mac-
ronuclear nodules, after protargol impregnation 1.6-2.8 ~m across. Contractile vacuole
during diastole with 2 collecting canals. Cortical granules absent. In posterior half of cell
usually many "saturn-shaped", yellowish, 1.5-4.0 ~m sized crystals. Movement charac-
teristic: crawls a short distance then suddenly jerks back, whereby it is conspicuously
curved (Fig. 139b).
Adoral zone of membrane lIes about 113 of body length. Bases of largest mem-
braneHes in life only 3 ~m broad. Buccal area flat. Paroral only slightly curved, endoral
short, straight, arranged beside paroral. Frontal and frontoventral cirri not enlarged. In-
variably 5 thick, in life about 15 ~m long transverse cirri; GELLERT (1957) drew only 4
transverse cirri; possibly he took the right transverse cirrus as right marginal cirrus. Re-
maining cirri about 8 ~m long. Left transverse cirrus in life conspicuously spread to left,
transverse cirrus VIII in life often slightly thicker than other transverse cirri. Two pre-
transverse ventral cirri, right cirrus (V1/2) almost in gap between right transverse cirrus
and posterior-most cirrus of right marginal row (Fig. 138d, 139d, arrowhead), which be-
gins just at level of cytostome. Left marginal row terminates almost at posterior end. Dor-
sal kineties 1 and 4 slightly shortened anteriorly, kinety 4 only 113 of body length. Dorsal
cilia in life 2-3 ~m long, posterior-most cilia of subspecies humicola also 2-3 ~m long,
posterior-most cilia of subspecies longisetum about 10 ~m long.

o c cur r e n c e and e colo g y: Widely distributed, but rather more rare than com-
mon in terrestrial habitats. Locus classicus (of T. humicola and T. humicola humicola) is
the litter ofa deciduous forest in Hungary (GELLERT 1957). FOISSNER (1984, 1987b) found
Tachysoma humicola in the upper soil layer of a salt-steppe in the so-called "Hl>lle" in
Seewinkel, Burgenland, Austria, and in the soil of a flood plain in the Fuscher valley,
Salzburg, Austria. Further records: spruce forest in Upper Austria (AEsCHT & FOISSNER
1993); bark of Acacia trees and soil in Costa Rica (FOISSNER 1994c, 1995b); soil from
flood plain primary (?) rain forest near Manaus, Brazil (FOISSNER 1997b); soil of the bush
in the Brisbane Water National Park near Sydney and in the litter and root layer and un-
der moss of a pine forest near Adelaide, Australia (BLATTERER & FOISSNER 1988). Locus
classicus of T. humicola longisetum is the upper soil layer from the Shetani volcano in
the Tsavo National Park, Kenya(?).
Feeds on bacteria, testate amoebas (Trinema lineare), and detritus (FOISSNER 1984,
GELLERT 1957). Biomass of 106 individuals about 9 mg (FOISSNER 1987a).
456 SYSTEMATIC SECTION

Tachysoma humicola humicola GELLERT, 1957 (Fig. 139a-f, Table 25)

1957 Tachysoma humicola GELLERT, AnnIs Inst. bioI. Tihany, 24: 20 (Fig. 139t).
1984 Tachysoma humicola GELLERT, 1957 - FOISSNER, Stapfia, 12: 121 (Fig. 139a-e; authoritative redescrip-
tion; slides of nootype specimens are deposited in the OberOsterreichische Landesmuseum in Linz, Up-
per Austria).

Rem ark s: Taxonomy, morphology, and ecology, see T. humicola. Differs from the
second subspecies, T. humicola longisetum (see below), ahnost exclusively in that the
posterior-most dorsal cilia are not elongated (Fig. 13ge).

Tachysoma humicola longisetum FOISSNER, 1998 (Fig. 138b-e, Table 25)

1998 Tachysoma humicola /ongisetum FOISSNER, Europ. J. Protistol., 34: 226 (Fig. 138b--e; slides of type
specimens are deposited in the OberOsterreichische Landesmuseum in Linz, Upper Austria).

Rem ark s: Taxonomy, morphology, and ecology, see T. humicola. Differs from the
second subspecies, T. humicola humicola (see above), ahnost exclusively in that the
posterior-most dorsal cilia are distinctly longer (about 10 J-lm) than the other cilia
(2-3 J-lm; Fig. 138c, e). Detailed description of this subspecies, see ForssNER (1998).

Tachysoma balatonica GELLERT & TAMAs, 1958 (Fig. 139g)

1958 Tachysoma balatonica GEI.LERT & TAMAs, Annis Inst. bioI. Tihany, 25: 229, 240.

T a x 0 nom y: I have some doubt that this is a Tachysoma because the adoral zone is
about 50 % of body length and the posterior postoral ventral cirrus is displaced distinctly
posteriad, indicating that it belongs to the Stylonychinae (for example, Histriculus). The
posterior-most marginal cirri are enlarged and elongated similarly to in T. bicirratum;
however, this species is smaller, has only one micronucleus, and probably long dorsal
cilia (Fig. 139h). Detailed redescription necessary; possibly better classified as species
indeterminata.

M 0 r ph 0 log y: In life 130-140 J-lm long. Elliptical. Flexibility not mentioned (see
also taxonomy). Each macronuclear nodule with a single micronucleus. Adoral zone of
membrane lIes about 50 % of body length. About 41 adoral membranelies, 13 distal mem-
branelles high, proximal membranelles low. Frontal, ventral, and transverse cirri dis-
tinctlyenlarged. GELLERT & TAMAs (1958) drew only 2 frontoventral cirri, which would
be very unusual; presumable they omitted 2 cirri. Two pretransverse ventral and 5 fringed
transverse cirri. Posterior-most cirrus of each marginal row conspicuously enlarged and
 Taehysoma 457

elongated. Dorsal cilia very likely short and inconspicuous because not mentioned by
GELLERT & TAMAs (1958).

o c cur r e n c e and e colo g y: Locus classicus is the eastern shore of Lake Bala-
ton, Hungary, where GELLERT & TAMAs (1958) discovered it in detritus drifts. GELLERT &
TAMAs (1959b) found it in the same biotope on the southern shore of this lake. Feeds on
diatoms, according to GELLERT & TAMAS (1959b, 1961) exclusively on Navicula crypto-
eephala, and small globular green algae.

Tachysoma bicirratum (FOISSNER, BLATIERER, BERGER & KOHMANN,

1991) FOISSNER, BLATIERER, BERGER & KOHMANN, 1991 (Fig. 139h)

1932 Tachysomafurcata KAHL, Tierwelt Otl., 25: 605 (Fig. 139h).

1972 Tachysomafurcata KAHL, 1932- BORROR, J. Protozool., 19: 15.
1988 Tachysomafurcatum KAHL, 1932 - FOlSSNER, Hydrobiologia, 166: 30 (emendation).
1991 Oxytricha bicirrata FOISSNER, BLATfERER, BERGER & KOllMANN, lnfonnationsberichte Bayer. Landesam-
tes ftlr Wasserwirtschaft, 1191: 302 (new name for T. furcata KAHL; see taxonomy).
1991 Tachysoma bicirratum (FOIsSNER, BLATfERER, BERGER & KOllMANN, 1991) - FOlSSNER, BLATfERER,
BERGER & KOllMANN, Infonnationsberichte Bayer. Landesamtes ftlr Wasserwirtschaft, 1191: 302 (see no-
menclature).

Nom e n cia t u rea n d t a x 0 nom y: KAHL (1932) classified Taehysoma as sub-

genus of Oxytrieha, so that the correct name in the original description is Oxytrieha
(I'aehysoma) fureata KAHL, 1932. However, this is a junior primary homonym of O. fur-
eatus SMITH, 1897b, which was emended to O.fureata by KAHL 1932. KAHL's (1932) spe-
cies therefore required a new name (ForSSNER et al. 1991).
Taehysoma bicirratum differs from the somewhat doubtful T. baiatoniea, which also
has two elongated posterior-most marginal cirri, in body size, number of micronuclei, and
length of dorsal cilia.
Taehysomafureata sensu GELEI (1954) has short dorsal cilia; furthermore, I believe
that the elongated cirri at the posterior end of GELEI'S (1954) specimens are caudal cirri,
suggesting a classification in Oxytricha (see O. pseudofurcata, Fig. 53a, b). Tachysoma
bicirratum should not be confused with Oxytricha setigera or O. balladyna, which also
have a small size, long dorsal cilia, and a single micronucleus between the macronuclear
nodules. Detailed redescription necessary.

M 0 r p hoI 0 g y: In life 60-90 J.1m long. Slender ovoid, especially anteriorly distinctly
flattened. Adheres to substrate with elongated posterior-most marginal cirri during food
gathering. All adoral membranelles conspicuously strong. Dorsal cilia delicate and soft:.

o c cur r e n c e and e colo g y: Rare in the aufwuchs, detritus, and sapropel of

freshwater habitats. Europe. After KAHL (1932) common but always only scattered in the
458 SYSTEMATIC SECTION

sapropel, according to FOISSNER (1979b) characteristic for eutrophic, temporal small wa-
ter bodies.
Locus classicus not indicated (KAHL 1932). DINGFELDER (1962) found Tachysoma bi-
cirratum in puddles with decaying plant material mainly during summer and in flooded
meadows near Erlangen, Germany, sometimes up to 70 indo ml- I (PH 7.0-7.4); possibly
he confused it with a terrestrial form, for example, T. humicola. TAMAs & GELLERT (1958)
recorded T. bicirratum from the periphyton of the littoral stones of Lake Balaton, Hun-
gary. Further records: Maritza River, Bulgaria (DETCHEVA 1981); running waters in
Czechoslovakia (BtL y et al. 1952).
Feeds on bacteria (" Rhodobacteria") and small algae (KAHL 1932). Biomass of 106
individuals about 15 mg (FOISSNER et al. 1991). MAnoN! & GHETTI (1981a) observed T. bi-
cirratum scattered in a betamesosaprobic to alphamesosaprobic portion of an Italian
brook at 18-21 °C, pH 7.4---8.0, 440-470 IlS cm- I specific conductivity, 8.3-9.7 mg I-I
Oz, 11-13 mg I-I COD, 0.49-0.79 mg I-I NH/-N, 0.09-0.16 mg I-I NO£-N, 1.6-5 mg I-I
NO)--N, and 0.1-0.18 mg I-I POlo-po DETCHEVA (1983c) found it with low frequency
(0.6 %) in the Maritza River, Bulgaria, under following conditions: h-a, pH 6.8-7.4, 3.4
to 10.1 mg I-I Oz, (39-89 % Oz saturation), 12.5 mg I-I BODs, 50-69 mg I-I Ca2+, 0 to
19 mg I-I Mg2+, 91-293 mg I-I HCO)-, 14-107 mg I-I SOlo, 8.5-23 mg I-I CI-, 7-14 °dR,
0.03-0.05 mg I-I NH/, 1-1.5 mg I-I NO)-, 0-0.02 mg I-I NOz-, 0.03-0.05 mg I-I Fe z+.
Tachysoma bicirratum is classified as alphameso- to polysaprobic indicator of water
quality (a-p; b = 2, a = 4, P = 4, I = 2, SI = 3.2; FOISSNER 1988a, FOISSNER et al. 1991,
SLADECEK et al. 1981, WEGL 1983). This does not agree very well with the above men-
tioned habitats and autecological data, except with that ofKAHL (1932), who found it in
the sapropelic area. Since his record is the only one which is substantiated by an illustra-
tion, the classification is preliminarily accepted; however, it has to be verified on reliably
determined populations.

Tachysoma terricola HEMBERGER, 1985 (Fig. 139i-n, Tables 3, 25)

1982 Tachysoma terricola HEMBERGER, Dissertation, p 257 (Fig. 139i-n).

1985 Tachysoma terricola HEMBERGER, Arch. Protistenk., 130: 414 (Fig. 139i; 1 slide ofholotype-specimens
is deposited in the Institut filr landwirtschaftliche Zoologie of the University of Bonn, Germany).

T a x 0 nom y: The present species is reminiscent of T. pellionellum as concerns body

size and long dorsal cilia; however, they can be easily distinguished by the number of mi-
cronuclei and dorsal kineties, the beginning of morphogenesis, the habitat, and some mi-
nor differences in the infraciliature. Live observations necessary.

M 0 r p hoI 0 g y and b i 0 log y: In life (?) 70-100 x 28-40 /lID. Elliptical, slightly
sloped posteriorly. Invariably 2 micronuclei. Cytoplasm granulated. Adoral zone ofmem-
branelles about 25 % of body length. Frontal cirri not enlarged. Marginal cirri about
12 11m long. Invariably 2 pretransverse ventral cirri and 5 enlarged, about 20 11m long
 Tachysoma 459

OP

Fig. 139j-n Tachysoma terricola (from HEMBERGER 1982. j-n, morphogenetic stages after protargol impreg-
nation). j) Very early stage showing the 4 oral primordia k, I) Early stages. Note that all postoral ventral cirri
are involved in primordia formation. m, n) These middle stages differ significantly from that of T. pellionellum
in that no primary primordia are formed. OP = oral primordium; p 458.

transverse cirri, which are not as steeply arranged as in T. pellionellum. Three dorsal ki-
neties of body length and 1 shortened kinety. Dorsal cilia about 10 J,lm long.
Morphogenesis commences with prolifemtion of basal bodies at four sites (Fig.
139j). Some middle stages show that no so-called primary primordia are formed as in T.
pellionellum, indicating that Tachysoma is not monophyletic (Fig. 139k-n). This is sup-
ported by the fact that dorsal morphogenesis is in Urosomoida pattern (Fig. 24b) and not
in Tachysoma pattern.

o c cur r e n c e and e colo g y: Locus classicus is the soil of a woodland in Peru.

Not found since. Biomass of 106 individuals about 45 mg (FOISSNER 1987a).

Tachysoma granuliferum BERGER & FOISSNER, 1987 (Fig. 140a-e,

Table 25)

1987 Tachysoma granulifera BERGER & FOISSNER, Zoo!. lb. Syst., 114: 228 (Fig. l40a-e; I slide ofholotype
specimens and 1 slide of paratype specimens are deposited in the Oberosterreichische Landesmuseum
in Linz, Upper Austria).
1988 Tachysoma granuliferum BERGER & FOISSNER 1987 - FOISSNER & FOISSNER, Catalogus Faunae Austriae,
Ie: 92 (emendation).

T a x 0 nom y: Tachysoma granuliferum can be easily distinguished from the other

Tachysoma species by the conspicuous cortical granules. In life it may be confused with
some species of Cyrtohymena (for example, C. cifrina) because the colour and arrange-
 ~
0\
o
~,\y~
~,.~. ,
s'
\
~,~\ ··:f'~;:SI " ~"
'. ~ ~ §S ~I •
"'*\l
!J:,~,:',
~~'JI'!I,: 'ff'",,,'. ",....~
. ... .:..
/iti~~I~;, ~+~ "
.,, ., , 1') \
4,: r'.~: ~'~A/l
~~;~:~;~.... ~;,
"" , i \~
~~ , ~ 'kF' I :
"
0' ,.
:::::~?~.""'''<~~':':~V,~ ~: ~
r:)\' ,
~@r .:~j<4( ," t:~"~ "'"•
!:
:. . ','
: #
' '
C/:J
-<
~j~;~(/~~'f~ .-~J­ ::: :. # .'~';:.:/
\ .'1 ~;"~clT
,..., , .. \ C/:J
tJ
, ....,
-it'·'~·I~~',~ :- . .\: :: • • '"
"
.- •• !:-
,. :: ~ #
. .':
tJ # (1
i~~~f , ~
C/:J
~~.tij.:'~;'''t;,.~.·~.;,~ # tTl
(1
~t:,l·~~l\'l~~;;,~
, ....,
V' ~~ . ,f,f)'"
Xi,,'\'•. '.:~.'"i~.'I'~""~iJ ;·;r'-
..:~.';'-~
~ '''IIIir--,'/ "'#'~'~\ ~
':~i~ ~"'.'/'~' . •
•.:•
"r~: b c d
Fig. 140a-e Tachysoma granuliferum (from BERGER & FOISSNER 1987a. !H:, from life; d, e, protargol impregnation). a-c) Ventral, right lateral, and dorsal view, a = 77
11m, b, c = 80 J.lm. The yeUow to orange-yeUow, about 0.5 J.lm sized cortical granules are shown in (c) only. d, e) Ventral and dorsal infraciliature, 55 11m. Note the linear
arrangement offrontoventral cirri (arrow) which is reminiscent of Urosoma; p 459.
 Tachysoma 461

ment of the granules are rather similar; furthermore, Tachysoma granuliferum has a peri-
stomiallip simulating a Cyrtohymena-like oral apparatus.

M 0 r ph 0 log y and b i 0 log y: In life about 70-85 x 25-35 J.1m. Elliptical, both
ends rounded. Macronuclear nodules after protargol impregnation 13 x 6 J.1m on average,
with small nucleoli. Constantly (n = 11) two, in life about 3 x 2 J.1m sized micronuclei.
Contractile vacuole with an anterior collecting canal. Cortical granules 0.5 J.1ffi across,
yellow to orange-yellow, irregularly distributed in loosely arranged groups (Fig. 140c); in
small, perhaps precystic specimens they are distinctly orange. Cytoplasm with some small
crystals, many 2-5 J.1m sized homogenous globules, and food vacuoles. Rapid movement.
Adoral zone of membranelles about 35 % of body length, bases of largest mem-
branelles in life about 6 J.1m broad, cilia of distal membranelles in life about 15 J.1m long.
Buccal area small, deep, and rather distinctly curved anteriorly. Undulating membranes
bent, superimposed anteriorly. Bases of frontal cirri only slightly enlarged. Frontoventral
cirri arranged in a line. Invariably (n = 11) only 1 pretransverse ventral cirrus, arranged
very close to the slightly enlarged, in life about 22 J.1m long transverse cirri. Marginal
cirri in life about 15 J.1m long, going backward bases become smaller and distances be-
tween them become wider; marginal rows distinctly separated posteriorly. Dorsal cilia in
life about 4 J.1m long; dorsal kinety 1 slightly shortened anteriorly (Fig. 140d, e).

o c cur r e n c e and e colo g y: Very likely autochthonous soil species (FOISSNER

1987a). Locus classicus is an arable soil near Vienna, Austria. We found Tachysoma
granuliferum also in an arable soil in Styria, Austria (FOISSNER et al. 1990). Feeds on
amoebas and their cysts, fungal spores (7 x 2 J.1m), green algae, and heterotrophic flagel-
lates. Biomass of 106 individuals about 26 mg (FOISSNER 1987a).

Marine species

Tachysoma rigescens (KARL, 1932) BORROR, 1972 (Fig. 141a-c)

1932 Tachysoma? (ev. Oxytricha) rigescens KAlIL, 1932, Tierwelt Dti., 25: 605 (Fig. 141a).
1933 Tachysoma rigescens KAlIL 1932 - KAm., TierweltN.- u. Ostsee, 23: 113.
1967 Tachysoma rigescens KAlIL, 1932 - ALAoRO LUBEL & L6PEZ-OcHOTERENA, Revta Soc. mex. Rist. nat.,
28: 63 (Fig. 14Ib).
1972 Tachysoma rigescens KAlIL, 1932 - BORROR, J. Protozooi., 19: 15.
1990 Tachysoma rigescens KAlIL, 1932 - ALAoRO LUBEL, MARTiNEz MURILLO & MAvEN ESTRADA, Manual de
ciliados, p 139 (Fig. 141c; includes only data from ALAoRO LUBEL & L6PEZ-OCHOTERENA 1967).

Nom e n cIa t u rea n d t a x 0 nom y: KAHL (1932) classified Tachysoma as sub-

genus of Oxytricha; the correct name in the original description is thus Oxytricha (Tachy-
soma) rigescens KAHL, 1932. I fix BORROR (1972a) as the author ofthe combination with
462 SYSTEMATIC SECTION

Tachysoma. Detailed description necessary, especially

the lack of the caudal cirri has to be checked.

M 0 r p hoI 0 g y and b i 0 log y: In life 150 to

180 x 30-40 J.l.m (KARL 1932), according to ALAnRO Lu-
BEL & LOPEZ-OCHOTERENA (1967) 175-260 x 39-55 J.l.m.
Outline long elliptical, at level of buccal vertex slightly
broadened. Moderately flattened dorsoventrally. Macro-
nuclear nodules 17-23 x 9 J.l.m (ALAnRO LUBEL & LoPEZ-
OCHOTERENA 1967). Very likely two micronuclei (KAHL
1932). ALAnRO LUBEL & LOPEZ-OcHTOTERENA (1967) de-
scribed a 7 J.l.m sized contractile vacuole in the posterior
end of the cell, which is very likely a misobservation.
Ring-shaped cytoplasmic inclusions ("Reservek~rper").
According to KAHL (1932) Tachysoma rigescens is eas-
ily recognisable because it often lies immobile on the
detritus.
Adoral zone of membranelles about 1/3 of body
length, extends far along right body margin. Marginal
cirri short and strong, protrude beyond body margin
only posteriorly. Five distinctly enlarged transverse
cirri, protrude by 113 to 112 beyond posterior end of
cell.

o c cur r e nee and e colo g y: Locus classicus is

c Heligoland, Gennany, where KAHL (1932) discovered
Fig. 141a--c Tachysoma rigescens
Tacfzysoma rigescens in the betamesosaprobic detritus
(3, from KAHL 1932; b, from ALA. of a water tank. ALAnRO LUBEL & LOPEZ-OcHOTERENA
ORO LuBEL & L6PEZ-OcHOTERENA (1967) found it in a brackish lagoon, Mandinga, near the
1967; c, from ALAoRO LuBEL et aI. port of Veracruz, Mexico (see also LOPEZ-OcHOTERENA
1990. a-c, from life). a--c) Ventral & ROURE-CANE 1970).
views, a = 160 flm, b = 200 flDl, C
= 175 flm;p461.
Records not substantiated by illustrations: with a
frequency of 4.3 % in the mesopsammon of the Bulgar-
ian coast of the Black Sea at 23-27.5 °C and 0.5 to
1.78 % salinity (DETCHEVA 1977, 1980, 1982a, 1983d);
sediment of the Caspian Sea (AGAMALIEV 1971); brackish waters in Spain (MARGALEF
LOPEZ 1945).
The records from a Donor water reservoir (ALEKPEROV 1989) and from agricultural
soils in Czechoslovakia (TIRJAKovA 1988) are also not substantiated by illustrations and
are very likely misidentifications.
 Tachysoma 463

Fig. 14ld-i Tachysoma dragescoi (from SONG & WILBERT 1997. d, g-~ from life; e, f, protargol
impregnation). d) Ventral view, 47 ~m. e, 1) Ventral and dorsal infraciliature, 36 ~m. Arrow in (e) marks ante-
rior end of right marginal row, arrowhead denotes anterior-most postoral ventral cirrus (cirrus lVl2). Tiny ar-
rows in (f) denote very small micronuclei. g) Dorsal view showing long transverse cirri, which protrude dis-
tinctly beyond posterior body end, and short rows of cortical granules. h) Part of pellicle showing dorsal cilia
and cortical granules about 1 ~m across. i) Lateral view. BC = buccal cirrus, TC = transverse cirri.

Tachysoma dragescoi SONG & WILBERT, 1997 (Fig. 141d-i, Table 25)

1997 Tachysoma dragescoi SONG & WILBERT, Europ. J. Protistol., 33: 58 (I holotype and 2 paratype slides of
protargol-impregnated specimens are deposited in the Laboratory ofProtozoo\ogy, College of Fisheries,
Ocean University ofQingdao, China).

M 0 r ph 0 log y and b i 0 log y: In life about 40-50 x 20-25 /lIn. Outline elliptical
to oval, shape and size of life specimens quite constant. Body rigid (which is not typical
for a member of the Oxytrichinae and possibly due to the small size). Macronuclear nod-
ules oval with large nucleoli, widely separated from each other; each nodule with 1 very
small micronucleus (Fig. 141 f). Contractile vacuole not present. Cytoplasm colourless
and very transparent, with several to many large food vacuoles. Cortical granules in life
464 SYSTEMATIC SECTION

about 1 ~m across, usually in groups of2-4 and arranged in irregular lines (Fig. 141g, h);
colour unfortunately not mentioned, possibly more or less colourless. Movement typi-
cally slow to medium-fast and continuous on bottom of Petri dish.
Adoral zone of membranelles about 33-40 % of body length, basis of largest mem-
branelles about 5 ~m wide. Buccal area vel)' narrow. Endoral and paroral short and al-
most straight. Buccal cirri in front of undulating membranes. Frontoventral cirri in life
8-10 ~m long. Arrangement of frontal-ventral-transverse cirri, see Figure 14le. Invaria-
bly (n = 16) 8 frontal, frontoventral, and buccal cirri, and 5 postoral and pretransverse
ventral cirri; that is, in total 17 frontal-ventral-transverse cirri because invariably (n = 16)
4 transverse cirri present; transverse cirri distinctly enlarged, about 15-20 ~m long, pro-
trude distinctly beyond posterior end of cell, sometimes used as thigmotactic structures
with which ciliate anchors to substrate. Marginal cirri distinctly separated posteriorly,
right row begins slightly above mid-body (Fig. 141e, arrow). Dorsal kineties as long as
body, cilia in life about 3-4 ~m long.

o c cur r e n c e and e colo g y: Marine. Locus classicus is Taipingjiao (the Peace

Cape) of Qingdao, China, where SONG & WILBERT (1997) discovered T. dragescoi in a
eutrophic pond used for storing marine shellfish in mid-October, using a polyurethane
foam unit for collecting. Feeds on flagellates and bacteria

Tachysoma ovala SONG & WILBERT, 1997 (Fig. 141j-q, Table 25)

1997 Tachysoma ovala SONG & WILBERT, Europ. J. Protisto!., 33: 55 (l holotype and 2 paratype slides of
protargol-impregnated specimens are deposited in the Laboratory of Protozoology, College of Fisheries,
Ocean University of Qingdao, China).

M 0 r ph 0 log y and b i 0 log y: In life about 35-50 x 20-30 Jlm. Delicate and
slightly contractile. Outline elliptical, that is, both ends usually slightly narrowed (Fig.
141j, 0), although shapes as shown in Figures 1411, m not seldom observed. Slightly
twisted due to spiral arrangement of marginal cirral rows (Fig. 14Il, m). Ventral side
slightly concave (Fig. 141k). Macronuclear nodule closely spaced, with few large nucle-
oli, always lying left of median. One large micronucleus between macronuclear nodules
(Fig. 141p, q). No contractile vacuole to be recognised. Cytoplasm colourless, packed
with 2-3 Jlm sized, shining granules and several food vacuoles. Cortical granules ellip-
soid, about 1.2-1.5 ~m long, sparse and irregularly arranged either singly or in pairs and
not in distinct lines (Fig. 1411~). Movement typically slow and continuous on bottom of
Petri dish.
Adoral zone of membrane lies 40-50 % of body length (which is rather long for a
member of the Oxytrichinae), bases of largest membranelles (in life?) about 5 Jlm wide.
Buccal area vel)' narrow; paroral and endoral of usual length, slightly curved. Cirri usu-
ally associated with pronounced argyrophilic fibres. Frontoventral cirri in life about
8-10 ~m long. Arrangement of frontal-ventral-transverse cirri, see Figure 141 p. Invaria-
 Tachysoma 465

Fig. 141j-i) Tachysoma ovata (from SONG & WILBERT 1997. j-o, from life). j) Ventral view, 35 /lm. k) Left
lateral view showing concave ventral surface (arrow). I, m) Ventral views showing arrangement of cortical
granules and variability of body shape. Note furrows caused by spiral marginal cirra1 rows. n, 0) Part of pelli-
cle and dorsal view (35 /lm) showing ellipsoid, 1.2-1.5 /lm long, cortical granules and arrangement of dorsal
cilia; p 464.

bly (n = 16) 8 frontal, frontoventral, and buccal cirri and 5 postoral ventral and pretrans-
verse ventral cirri, that is, a typical 18-cirri oxytrichid, because invariably (n = 16) 5
transverse cirri present (Table 25). Postoral ventral and pretransverse ventral cirri not dis-
tinctly separated (Fig. 141p). Transverse cirri in life about 12 J.lm long, protrude only
slightly beyond posterior end of cell. Marginal rows distinctly spiral and widely separated
posteriorly. Right marginal cirral row beginning dorsally near anterior end of cell, termi-
nating close to rightmost transverse cirrus. Left marginal cirral row curved from ventral
to dorsal side, making nearly a half turn to posterior end of cell (Fig. 141 p, q). Three
complete dorsal kineties each with only 4-6 pairs of widely spaced basal bodies, some in-
466 SYSTEMATIC SECTION

\ ,
I '

!
/
•

/
,

p q

Fig. 141p, q Tachysoma ovala (from SONG & WILBERT 1997. p, q, protargol impregnation). p) Ventral infra-
ciliature and nuclear apparatus, 34 j.lm. Tiny arrows mark frontoventral cirri, which are not distinctly separated
from postoral ventral cirri. Arrowhead denotes right frontal cirrus. q) Dorsal infraciliature and nuclear appara-
tus, 34 j.lm. Arrow denotes micronucleus; p 464.

dividual basal body pairs in anterior half of cell; cilia in life about 4-5 11m long (Fig.
141q).

o c cur r e n c e and e colo g y: Marine. Locus classicus is Taipingjiao (the Peace

Cape) ofQingdao, China, where SONG & WILBERT (1997) discovered T. ovala in the open
water (salinity 32 %0, pH 8.1-8.3, 5-13°C) in early December, using a polyurethane
foam unit for collecting. Feeds on diatoms and flagellates.

Species indeterminata

Taclrysoma parvisty/um STOKES, 1887, Ann. Mag. nat. Hist., 20: 109 (Fig. 225a). Re-
marks: The body shape, the arrangement of the cirri, and especially the single macronu-
clear nodule indicate a misobservation; perhaps a postconjugant of another species. KAHL
(1932, p 605) considered this form as valid (T. parvistyla); the correct name in his revi-
sion is Oxytricha (rachysoma) parvistyla because he classified Tachysoma as subgenus
of Oxylricha. In life about 63 11m long; pyriform; colourless; 20 frontal, ventral, and
 Tachysoma 467

transverse cirri in a very unusual arrangement; contractile vacuole in about mid-body;

dorsal cilia short; rapid movement. Locus classicus is a shallow pool near Trenton in
New Jersey, USA. Further records from freshwater habitats: Greece (STEPHANlDES 1948);
Italy (MAooNI & GHETII 1977); Latvian rivers (LIEPA 1978); China (SHEN 1980); USA
(BOVEE 1960, CAIRNS & YONGUE 1973a, EDMONDSON 1920, MURRAY 1974, RUlHVEN 1972,
WANG 1928); Antarctica (HAWTHORN & ElLIs-EVANS 1984). DETCHEVA (1979c, 1981,
1983c) provided some autecological date of this unreliable species. According to CuRDs
(1975), Tachysoma parvistyium occurs also in activated sludge. AGAMALIEV (1975) re-
corded it from the west coast of the Central Caspian.

Insufficient redescriptions

Unless otherwise indicated, the following taxa are very poorly described and/or illus-
trated or the habitats do not match (especially the terrestrial records of T. pellionellum),
so that the identifications cannot be accepted.

Oxytricha agilis STOKES - CONN, 1905, Bull. Conn. St. geol. nat. Hist. Surv., 2: 60 (Fig.
225t). Remarks: Freshwater habitats in Connecticut, USA.

Oxytricha parvistyla STOKES - CONN, 1905, Bull. Conn. St. geol. nat. Hist. Surv., 2: 60
(Fig. 225b, c). Remarks: Freshwater habitats in Connecticut, USA.

Oxytricha pellionela (0. F. M) EHRBG. - BRODSKY & YANKOWSKAYA, 1929, Acta Univ.
Asiae mediae, 6: 33 (Fig. 226t). Remarks: Incorrect subsequent spelling. Two micronu-
clei. Soil from Central Asia.

Oxytrichapellionella, EHR. - BELTRAN, 1928, Mems Revta Soc. cient. 'Antonio Alzate',
49: 133 (Fig. 228c). Remarks: In life 70-110 x 3~6 J.1m. Two micronuclei. Lake Xo-
chimilco, Mexico.

Oxytricha pellionella - DUJARDIN, 1841, Zoophytes, p 417, Planche 11, fig. 10. Remarks:
STEIN (1859b) already mentioned the insufficiency of this drawing.

Oxytricha pellionella - QUENNERSTEDT, 1865, Acta Univ. lund., 2: 58 (Fig. 225n). Re-
marks: Freshwater habitats in Sweden (see also QUENNERSTEDT 1869).

Oxytricha pellionella - SMITH, 1978, Scient. Rep. Falkld lsI. Depend. Surv., No. 95: 75,
77 (Fig. 225m). Remarks: Edaphic habitats in Antarctica.

Oxytricha pellionella EHRENBERG - BUIsAN, 1944, Publnes Inst. BioI. apI., 1: 20 (Fig.
233d). Remarks: Among Cladophora and Spirogyra in freshwater habitats near Barce-
lona, Spain.
468 SYSTEMATIC SECTION

Oxytricha pellionella MOLL. - CONN, 1905, Bull. Conn. St. geol. nat. Hist. Surv., 2: 60
(Fig. 2251). Remarks: Freshwater habitats in Connecticut, USA.

Oxytricha pellione/la MOLL. - EDMONDSON, 1906, Proc. Davenport Acad. Sci., 11: 100
(Fig. 234i). Remarks: Freshwater habitat in Iowa, USA.

Oxytricha pellionella MOLL. - FELLERS & ALLISON, 1920, Soil Sci., 9: 9 (Fig. 225r). Re-
marks: Soils near New Brunswick, New Jersey, USA.

Oxytricha pellionella, MOLL. - KOFFMAN, 1926, Acta zooI., Stockh., 7: 310, Fig. 87. Re-
marks: Garden soil near Stockholm, Sweden.

Oxytricha pel/ionella MOLLER - SAMANO & SOKOLOFF, 1931, Monografias Inst. BioI.
Univ. naco Mex., 1: 34 (Fig. 225g). Remarks: Pond in Mexico.

Oxytricha pellionella MULL. - SMIlH, 1914, Kans. Univ. Sci. Bull., 9: 166 (Fig. 233g).
Remarks: In life 86-160 J.lm long. Two micronuclei. Possibly an Oxytricha. Infusion of
hay and leaves in Kansas, USA.

Oxytricha pellionella MULLER - WANG, 1925, Contr. bioI. Lab. Sci. Soc. China, 1: 50
(Fig. 242d). Remarks: Pond in Nanking, China.

Oxytricha pellionella (0. F. MOLL) EHR. - BIERNACKA, 1959, Polskie Archwm

HydrobioI., 5: 54, 57 (Fig. 225t). Remarks: Perhaps a Sterkiella histriomuscorum.
Sewage-treatment plant in Danzig, Poland.

Oxytricha pe/lionella STEIN, 1859 - LEPINIs, GELTZER, TSCHIBISOVA & GEPTNER, 1973, Key
to soil protozoa, p 155 (Fig. 225j). Remarks: Soil in the USSR

Tachysoma (Oxytricha) pellionella (MOLLER-STEIN) - SOKOLOFF & ANCONA, 1937, An.

Inst. BioI. Univ. Mex., 8: 175 (Fig. 234j). Remarks: Drinking water from towns in Mex-
ico.

Tachysoma parvistyla STOCK. - CHARDEZ, 1971, Revue verviet. Hist. nat., 28: 38 (Fig.
234n). Remarks: Freshwater habitat in Belgium.

Tachysoma parvistyia STOKES - LUNDIN & WEST, 1963, Free-living protozoa, p 67 and
WEST & LUNDIN, 1963, Pap. Mich. Acad. Sci., 48: 108 (Fig. 225d). Remarks: The outline
does, in fact, remind of the STOKES (1887) drawing (see Fig. 225a). The macronucleus of
the present form, however, obviously consists of several nodules. Freshwater habitats in
the Upper Peninsula of Michigan, USA.
 Tachysoma 469

Tachysoma pellionella (0. F. MOLLER, 1786) - ALADRO LUBEL, MARTiNEz MURILLO &
MAYEN ESTRADA, 1990, Manual de ciliados, p 138 (Fig. 241d). Remarks: Dorsal cilia
rather short. Marine habitats in Mexico.

Tachysoma pellionella - FERNANDEZ-LEBORANS & CASTRO DE ZALoUMBIDE, 1985, Micro-

bios, 42: 225, Fig. 1-11. Remarks: Two micronuclei. Soil near Madrid, Spain. Morpho-
genesis very insufficiently described.

Tachysoma pellionella (MOLLER-STEIN) - LOUSIER & BAMFORTH, 1990, Soil protozoa,

p 129, Fig. 6.29g. Remarks: Soil of North America.

Tachysoma pellionella MULLER-STEIN - RosA, 1957, Lesn. Cas., 3: 228, 235, 236 and
RosA, 1957, Pi'irodov. Sb. ostrav. Kraje, 18: 45 (Fig. 233a, c). Remarks: Soil of wood-
land in Czechoslovakia.

Tachysoma pellionella - TCHANG, PANG & Gu, 1984, Zoo!. Res., 5: 189 (Fig. 232j, k).
Remarks: The following characters indicate that this form is not identical with T. pe/-
lionel/um: (i) the bases of the transverse cirri are not distinctly enlarged; (ii) the number
of marginal cirri is distinctly higher; (iii) the frontoventral cirri and the dorsal kineties are
differently arranged; and (iv) the morphogenesis deviates from the description by HEM-
BERGER (1982), who determined the species correctly (Fig. 135h-o). Furthermore,
TCHANG et al. (1984) did not figure the nuclear apparatus and the length of the dorsal
cilia, the most important characters for a reliable identification of T. pellionellum (how-
ever, the Chinese text was not translated, so that I cannot exclude that these two charac-
ters are mentioned there!). China.

Tachysoma sp. - LEPSI, 1957, Trav. Mus. Hist nat. Gr. Antipa, 1: 90 (Fig. 225e). Re-
marks: LEPSI (1957) found this unreliable form in a mesosaprobic culture with roots from
a mountain lake in Romania. According to him, perhaps a "form" of T. pellionellum or an
unknown species which could not be studied sufficiently. In life 70 x 20 llm, both ends
rounded. Only one (!) ellipsoidal macro nuclear nodule in mid-body. The 3 left transverse
cirri are longer than the 2 right ones, which have a knob (?) at their posterior end.

Tachysoma sp. - LEPSI, 1965, Protozoologie, p 736 (Fig. 241c). Remarks: In life
100-160 llm long. No description. Romania?
470 SYSTEMATIC SECTION

Australocirrus BLAITERER & FOISSNER, 1988

1988 Australocirrus BLATIERER & FOISSNER, Stapfia, 17: 65 - Type (original designation): Australocirrus os-
citans BLATIERER & FOISSNER, 1988.
1997 Australocirrus BLATIERER & FOISSNER, 1988 - BERGER & FOISSNER, Arch. Protistenk., 148: 145.

C h a r act e r i sat ion: Adoral zone of membrane lIes formed like a question mark.
Undulating membranes curved, not distinctly intersecting. Frontoventral cirri form V-
shaped pattern. Postoral ventral cirri behind buccal vertex. Two pretransverse ventral and
5 transverse cirri. One right and 1 left row of marginal cirri. More than 6 dorsal kineties.
Caudal cirri present. Dorsal morphogenesis in Oxytricha pattern, however, more than 2
dorsomarginal kineties and kinety 3 not with simple, but with multiple fragmentation.

T a x 0 nom y: The characterisation is according to BERGER & FOISSNER (1997); for de-
tailed description of patterns mentioned, see chapter 2 in the general section. BIATIERER
& FOISSNER (1988) included two species, namely A. oscitans (type) and A. octonucleatus.
Both show multiple fragmentation of dorsal kinety 3, but differ significantly in body
flexibility. In the very flexible A. oscitans, cirrus V/3 is involved in primordia formation
(W. FOISSNER, unpubl. data; see also BERGER & FOISSNER 1997), supporting the classifica-
tion in the Oxytrichinae. Conversely, the body of A. octonucleatus is rigid, suggesting
that it is not the sister group of A. ascitans. I thus transfer A. octonucleatus to a new
taxon (Rigidocortex) belonging to the Stylonychinae. Multiple fragmentation of dorsal ki-
neties occurs also in the rigid taxa Pattersoniella, Laurentiella, and Onychodromus, and
in the flexible Territricha, indicating parallel evolution of this character. The large oral
apparatus of A. oscitans does not match the basic pattern of the flexible oxytrichids,
where the adoral zone is usually only 1/3 of body length; possibly, this is a further apo-
morphy (beside the multiple fragmentation of dorsal kinety 3) of Australocirrus.

Single species

Australocirrus oscitans BLAITERER & FOISSNER, 1988 (Fig. 142a-j,

Table 29)

1988 Australocirrus oscitans BLATIERER & FOISSNER, Stapfia, 17: 66 (the slide ofholotype specimens is de-
posited in the OberOsterreichische Landesmuseum in Linz, Upper Austria).

T a x 0 nom y: At superficial live observation A. oscitans can be confused with Holosti-

cha stueberi FOISSNER, 1987b (mid-ventral cirri present), which also has a very large buc-
cal area; furthermore, the resting cysts are very similar. Oxytricha iongissima has more
marginal cirri and only 5 dorsal kineties (no multiple fragmentation).
 )~ ,I J
') { ) (
) ) ( '\ I
') }) \
») I ) I \ I ( (
)}\ \ 1 \
') \ > 1
,') I ) J I
~' \
I~ 'I I I I 1
'~ I ) II II . I
') I ) 'I
~
~ ~ \l \ \ \ 1:;
::;-
\ ') ') I( \ ~
') \ \ \ I \ (';)
, =:;.
') \ \' I I )
~ ') • I \ I ~
)
') ') \.
,,
I
,, , .. ') I \ II I
') \ \
b c
, ~ I \ ) )j
~
,
~-. ..- ') \ I r
'"l r
~,

", d e
Fig. 142a-e Australocirrus oscitans (from BLATTERER & FOISSNER 1988. 11-<:, in life; d, e, protargol impregnation). a, c) Ventral views, a = 260 ~m. The large oral appa-
ratus suggested the species name oscitans (= yawning). b) Right lateral view. d, e) Ventral and dorsal infraciliature, d =205 ~m, e = 165 ~m. Arrow marks distal end of
adoral zone of membranelles. Arrowhead denotes posterior postoral ventral cirrus V/3 which is involved in primordia formation; the latter character and the flexible
body strongly indicate thatA. oscitans belongs to the Oxytrichinae although the adoral zone is exceptionally long, as in members of the Stylonychinae; p 470.
-...I
"'"
472 SYSTEMATIC SECTION

Fig. 142f-b Cyst of Australocirrus oscitam (original micrographs, kindly supplied by W. FOISSNER. f-h, Nor-
marski differential interference contrast). t) Total view, diameter about 70 11m. g) Conspicuous crystals in wall.
b) Cyst wall; p 470.

M 0 r ph 0 log y and b i 0 log y: In life 180-260 x 80-120 ~m. Broad elliptical,

that is, both ends broadly rounded, right and left body margin almost straight to slightly
convex, body usually slightly converging posteriorly. Flexible like, for example, Oxytri-
cha granuiijera, flattened about 2:1 , ventral side plain, dorsal convex. Invariably two (n
= 10) ellipsoidal macronuclear nodules, in life about 31-48 x 17-23 ~m, arranged slight-
ly left of median; numerous small spherical nucleoli. Usually 2, sometimes 3, in life about
7-8 x 6-7 ~m sized micronuclei adjacent to macronuclear nodules. Contractile vacuole
in about mid-body, during diastole often with conspicuous, bulbous collecting canals.
Pellicle colourless, distinct cortical granules lacking. Endoplasm densely granulated, with
moderately many, 1-5 ~m sized, greasily shining globules and about 8 ~m sized cyto-
plasmic crystals. Movement rapid.
Adoral zone of membranelles about 45 % or more of body length, makes a 3/4 turn
(Fig. 142d). Buccal area very broad. Bases of largest membranelles in life about 22 ~m
broad, distance between adoral membranelies conspicuously large. Undulating mem-
branes distinctly curved, arranged almost in parallel but widely separated, intersecting at
proximal end, both membranes probably consist of 2 rows of basal bodies each. Invaria-
bly (n = 10-11) 18 frontal-ventral-transverse cirri. Frontal cirri arranged almost longitu-
dinally, distinctly enlarged, in life about 30-40 ~m long. Buccal cirrus immediately left
of right frontal cirrus, about in middle of paroral (Fig. 142d). Postoral ventral cirrus V/3
 A ustralocirrus 473

, ..
.•.. •
•••
'

,
:
J
.
~.! i i :
: !~ ..
. '."
.• t# ~
'. ~
'.
~
~
j
•
Fig. 142i, j Australocirrus
asci/ans (original photomi-
crographs, kindly supplied
by W. FOISSNER. i, j, protar-
gol impregnation). i) Ventral
infraciliature of a late mor-
phogenetic stage. Large ar-
rows mark caudal cirri at
new dorsal kineties 1, small
arrow denotes new trarISverse cirrus VIII of opisthe, and arrowhead marks dorsomarginal kineties of opisthe.
j) Multiple fragmentation of dorsal kinety 3; p 470.

displaced distinctly posteriad (Fig. 142d) but, nevertheless, involved in primordia fonna-
tion (see taxonomy of genus section). Transverse cirri distinctly enlarged, arranged in
hook-shape, displaced distinctly anteriad and thus not protruding beyond posterior end of
cell (Fig. 142a). Ventral and marginal cirri in life about 25 l1m long. Marginal rows al-
most confluent posteriorly; right marginal row shortened anteriorly, that is, tenninates be-
side frontoventral cirrus VI/3. Dorsal kineties I and 2 of about body length; fragments of
474 SYSTEMATIC SECTION

Table 29 Morphometric data of Australocirrus oscitans (from BLAlTERER & FOISSNER (988). All data are
based on protargol impregnated specimens. All measurements in micrometres. CV = coefficient of variation
(in %), Max = maximum value, mean = arithmetic mean, Min = minimum value, n = sample size, SD = stan-
dard deviation

Character mean SD CV Min Max n

Body, length 208.7 16.8 8.1 190 250 10
Body, width 82.6 5.7 6.9 72 90 10
Macronuclear nodule, length 27.3 4.0 14.6 20 32 10
Macronuclear nodule, width 14.3 2.1 14.8 12 18 10
Micronuclei, number 2.1 0.3 14.4 2 3 11
Micronucleus, length 6.7 0.7 10.4 6 8 10
Micronucleus, width 4.8 0.3 6.3 4.5 5.3 10
Adoral zone of membranelles, length 89.4 2.6 2.9 85 94 10
Adoral membranelles, number 34.1 1.4 4.3 33 38 10
Right marginal row, number of cirri 21.2 0.9 4.3 20 23 10
Left marginal row, number of cirri 22.7 I.S 6.6 21 26 10

kinety 3 rather irregularly arranged in left half (in dorsal view) of cell (Fig. 142e). During
morphogenesis 3 dorsal primordia and more than 2 (3-5) dorsomarginal kineties occur in
both proter and opisthe (Fig. 142i). Dorsal cilia about 3 J.1m long. Three caudal cirri, one
each at posterior end of kineties 1 and 2 and at posterior-most fragment originating from
kinety 3 (Fig. 142j); sometimes (2 of 15 specimens) 4 caudal cirri.
Cyst spherical, about 70-80 J.1m across (Fig. 142f). Wall smooth, with a bluish shim-
mer, about 5 J.1m thick, consists of many thin layers. In the wall many conspicuous, about
2 J.1m sized crystals with 6 regularly arranged arms, each with an distinct knob at its distal
end (Fig. 142g, h). Mucous layer lacking.

o c cur r e n c e and e colo g y: Locus classicus is the Royal National Park south
of Sydney, Australia, where BLATTERER & FOISSNER (1988) discovered A. oscitans in the
upper soil layer (0-5 cm; pH 4.5) with much litter and brown sand of a woodland
("bush"). Further records: soil (0-8 cm, pH 5.1) from black-water inundation primary (?)
rain forest on Anavilhanas archipelagos in the Rio Negro, Brazil (FOIssNER 1997b); light
brown soil mixed with much leaf litter (PH 6.2) from Amazonian rain forest near Iquitos,
Peru (FOISSNER 1997b); soil from South Africa (W. FOISSNER, pers. comm.).

Feeds on fungal spores, testaceans (Euglypha sp., Trinema lineare), ciliates (Colpoda
sp., Blepharisma hyalinum), heterotrophic flagellates, and small green algae. Biomass of
106 individuals about 570 mg.
 Onychodromopsis 475

Onychodromopsis STOKES, 1887

1887 Onychodromopsis STOKES, Ann. Mag. nat. Hist., 20: 107 - Type (by monotypy): Onychodromopsis
jIexilis STOKES, 1887.
1888 Onychodromopsis, STOKES - STOKES, J. Trenton. nat. Hist. Soc., 1: 280.
1932 Onychodromopsis STOKES, 1887 - KAm., Tierwelt Dtl., 25: 620.
1974 Onychodromopsis STOKES - STILLER, Fauna Hung., 115: 107.
1985 Onychodromopsis - SMALL & LYNN, Ciliophora, p 458.

C h a r act e r i sat ion: Adoral zone of membranelles formed like a question mark.
Undulating membranes short, straight, and arranged almost side by side. Frontoventral
cirri in short transverse row or lacking. Postoral ventral cirri in longitudinal row right of
buccal vertex. Usually 2 pretransverse ventral and 5 transverse cirri. Two or more (mar-
ginal and frontoventral) cirral rows right of median and 2 left rows of marginal cirri. Cau-
dal cirri (very likely) present.

Nom e n cia t u rea n d t a x 0 nom y: "Onychodronum sp." in BUCHHOLZ &

BUCHHOLZ (1974, P 251) is likely an incorrect subsequent spelling of Onychodromopsis
or Onychodromus.
The taxonomy of Onychodromopsis is rather complicated. It was accepted by KAHL
(1932), STILLER (1974b), CORLISS (1979), and TUFFRAu (1987), but was differently classi-
fied. CORLISS (1979), for instance, mentioned it as incertae sedis in the Sporadotrichina,
whereas TUFFRAu (1987) classified Onychodromopsis in the Holostichidae. BOTsCHLI
(1889), BORROR (1972a, p 14), and CURDS et al. (1983) synonymised it with Pleurotricha,
which has, however, a rigid body and a different cirral pattern.
PETZ & FOISSNER (1996) redefmed Onychodromopsis, identifying their Sub-Antarctic
population, which has a typical Oxytricha cirral pattern (Fig. 92h), with the type, Ony-
chodromopsisflexilis (Fig. 143a). They assumed that the illustrations by STOKES (1887b,
1888; Fig. 143a, b), KAHL (1932; Fig. 143c), and SMALL & LYNN (1985; Fig. 143d) lack
some frontoventral cirri and are thus very likely incomplete, or, more likely, influenced
by STOKES' (1887b) figure. Thus, and in the absence of any type material, PETZ & FOISS-
NER (1996) fixed their population as neotype of Onychodromopsis.
By contrast, I consider the original description of 0. flexilis as more or less correct
because (i) STOKES usually made rather detailed and accurate descriptions, (ii) the protar-
gol stained Onychodromopsis sp. in SMALL & LYNN (1985; Fig. 143d) matches the figure
in STOKES (1888; Fig. 143a) in every detail (thus it is incomprehensible that they did not
identify their population as O. flexilis), (iii) the STOKES (1887b) and SMALL & LYNN
(1985) populations are very likely from the same geographic area, namely North
America, whereas the PETZ & FOISSNER (1996) populations are Sub-Antarctic, and (iv)
Paraurostyla viridis sensu PATSCH (1974) has almost the same infraciliature as O.flexilis
(see below), indicating that this cirral pattern is not a misobservation as supposed by PETZ
& FOISSNER (1996). The protargol-impregnated specimen has rather short and straight un-
dulating membranes (Fig. 143d); possibly the reason STOKES (l887b) did not illustrate
476 SYSTEMATIC SECTION

these structures, although he wrote ''peristome about one third as long as the body, the in-
ner or right-hand margin bearing a large and, in lateral view, conspicuous membrane".
Accordingly, o.jlexilis sensu PETZ & ForSSNER (1996) and some other taxa classified
with Onychodromopsis (for example, Pieurotricha variabilis REUTER) by these authors
have to be removed from Onychodromopsis. Since all have a typical Oxytricha cirral pat-
tern, I reactivate Allotricha STERKI (for details, see there). Parurosoma also has more than
2 marginal rows, but the frontoventral cirri are arranged as in Urosoma, that is, in a line.
Apoamphisiella has 2 frontoventral rows and lacks distinct pretransverse ventral cirri.
Two further Onychodromopsis species have been described, namely 0. kahli and O.
tihanyiensis. The latter species is described in detail by ForssNER (1 997b) and transferred
to Apoamphisiella (see p 782). Both species now included in Onychodromopsis have to
be redescribed in detail. I have no idea about the phylogenetic relationships, that is, the
sister-group of Onychodromopsis. It is included it in this book because it can be easily
confused with some taxa (for example, Allotricha, Parurosoma) which unequivocally be-
long to the flexible oxytrichids.

Key to species

Identifications of these little-known species should be substantiated by protargol impreg-

nation. For flexible oxytrichids with more than two cirral rows, see also Allotricha, Paru-
rosoma, Paraurostyia, Apoamphisiella or even Gastrostyla.

1 3-4 distinctly enlarged frontoventral cirri (including buccal cirrus) behind the 3
frontal cirri (Fig. 143a, d) ................... Onychodromopsisjlexilis (p 476)
Only I enlarged cirrus (= buccal cirrus?) behind the 3 frontal cirri (Fig. 143e) ..... .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. Onychodromopsis kahli (p 478)

OnychodromopsisjIexilis STOKES, 1887 (Fig. 143a--<i)

1887 Onychodromopsisjlexilis STOKES, Ann. Mag. nat Hist., 20: 107 (Fig. 143a).
1888 Onychodromopsisjlexilis, STOKES - STOKES, J. Trenton nat. Hist. Soc., I: 280 (Fig. 143b).
1932 Onychodromopsisjlexilis STOKES, 1887 - KAHL, Tierwelt Dtl., 25: 620; in part: Fig. 113 1 = redrawing
from STOKES 1888 (Fig. 143c); not Fig. 1I0s (see O. kahli).
1985 Onychodromopsis sp. - SMALL & LYNN, Ciliophora, P 458 (Fig. 143d).

T a x 0 n m y: See relevant chapter in genus section for complicated taxonomy. BORROR

(1972a) synonymised Onychodromopsis flexilis with Pleurotricha grandis, which has,
however, a larger, rigid body and a different cirral pattern. The KUSANO (1985) and
LUNDIN & WEST (1963; Fig. 223b) populations are insufficient redescribed. Detailed rede-
scription necessary. Onychodromopsis kahli (Fig. 143e, f; see below) is possibly a junior
synonym. Onychodromopsisflexilis sensu PETZ & FOISSNER (1996) is the authoritative de-
scription of Allotricha antarctica.
 Onychodromopsis 477

...
.. .,
·. •
.
·. ,.
,.

.. .. , .
e
a b c
. •••
.
•
. •••
.,

Fig. 143a-d Onychodromopsisflexilis (a, b, from STOKES 1887b, 1888; c, after STOKES 1888 from I<AHL 1932;
d, from SMALL & LYNN 1985. a-c, from life; d, protargol impregnation). a-c) Ventral view, 125 J,lm. The illus-
tration by STOKES (1888) differs from the original (a) in that the frontal-ventral-transverse cirri are enlarged and
in that the transverse cirri are distinctly separated from the inner left marginal row. It is unknown if these al-
terations are only due to redrawing, or if STOKES (1888) made use of new observations. d) Ventral infracili-
ature, 120 J,lm. Note that the cirral pattern matches the original illustration exactly; p 476.
Fig. 143e, f Onychodromopsis kahli (e, from I<AHL 1932; f, from SRAMEK-HUSEK 1957). Ventral views in life,
e = 120 J.llfl, f= 130 to 180 J.lm. Note single cirrus (= buccal cirrus?) at anterior end of undulating membranes;
p478.

M 0 r p hoi 0 g y: In life about 90-130 ~m long, about 113 as broad as long; after protar-
gol impregnation 120 ~m long. Soft and flexible. Elliptical, that is, both ends rounded.
Two ellipsoidal macronuclear nodules near left body margin, in either anterior or poste-
rior body-half. Contractile vacuole in about mid-body. No cortical granules mentioned.
Adoral zone of membranelles about 1/3 of body length. Undulating membranes not
illustrated (Fig. 143a, b) but mentioned in the text by STOKES (l887b, 1888; see taxonomy
chapter in the genus section); according to SMALL & LYNN (1985; Fig. 143d) inconspicu-
ous, that is, short and more or less straight. Buccal area narrow. Three distinctly enlarged
frontal cirri, somewhat behind (about at level of anterior end of undulating membranes) 3
transversely arranged cirri; the left one is very likely the buccal cirrus. Three or 2 longitu-
dinally arranged postoral ventral cirri, anterior one possibly always slightly in front of
level of buccal vertex (Fig. 143a, d). Two or 3 longitudinally arranged pretransverse ven-
478 SYSTEMATIC SECTION

tral cirri. Five transverse cirri, often furcate or fImbriate, protrude beyond posterior end
of cell. The almost "transverse" arrangement of the transverse cirri is rather unusual, but
of course not impossible! Cirri of outermost marginal rows longest and strongest. Left
outer cirral row almost overlapping on right margin, possibly posterior-most cirri of this
row are caudal cirri. Dorsal cilia "short" (that is, likely 2-4 11m).

o c cur r e n c e and e colo g y: Obviously very rare; possibly confIned to freshwa-

ter. Locus classicus is a Lemna pond in the USA, probably near Trenton, New Jersey,
where STOKES lived and worked. SMALL & LYNN (1985) mentioned no sample site, but very
likely they also found Onychodromopsis flexilis in North America. Further records from
the USA, which are, however, not substantiated by illustrations: Conestoga River Basin in
Lancaster County, Pennsylvania (CAIRNS 1965a); Douglas Lake Region, Michigan (CAIRNS
& YONGUE 1966); Copper Creek, Colorado (CAIRNS & YONGUE 1973b); predominantly in
cold sites (2-3 0c) oflimestone waters in Kentucky (GIITLESON & FERGUSON 1971).
PATRICK et al. (1967) found Onychodromopsisflexilis at two stations of the Savannah
River, USA, at following conditions: 23-28°C, 5-<10 mg 1-\ CI-, >3-7.5 mg 1-\ CO2,
5-7 mg 1-\ O2,0.1-0.4 mg 1-\ Fe2+, 0.02-0.04 mg 1-\ NH3-N, 0.001-<0.007 mg 1-\ N02--N,
0.07-0.2 mg 1-\ NOl--N, 0.03-0.08 mg 1-\ POlo, 6-12 mg 1-\ Si02, 10-<30 mg 1-\ SOl",
0.5-1 mg 1-\ BODs. MADaNI & GHEITI (1977) counted 16 indo cm·2 in an alphamesosapro-
bic to polysaprobic section of the Parma River, Italy. Two records from terrestrial habi-
tats, namely Abaco Island, Bahamas, and Italy are also not substantiated by illustrations
(CAIRNS & RUTIIVEN 1972, LUZZAITI 1938). Biomass of 106 individuals about 43 mg (NES-
TERENKO & KOVALCHUK 1991; incorrectly spelled as "Pleurotrychaflexilis").

Onychodromopsis kahli SRAMEK-HuSEK, 1957 (Fig. 143e, f)

1932 Onychodromopsisjlexi/is STOKES, 1887 - KARL, Tierwelt Dtl., 25: 620; in part: Fig. llOs (Fig. 143e);
not Fig. 113,.
1957 Onychodromopsis kahli SRAMEK-HuSEK, Vl!st. ~s!. zoo!. Spo!., 21: 15 (Fig. 143f).

Nom e n c I at u rea n d t a x 0 nom y: SRAMEK-HuSEK (1957) established this spe-

cies on his own and KAHL's observations. Thus the expression ''nom. nov." in the table
and in the headline in SRAMEK-HuSEK (1957, P 3, 15) is wrong. It differs from the type
species mainly in that it has only one enlarged frontoventral cirrus, which is very likely
the buccal cirrus. However, synonymy with Onychodromopsisflexilis or Allotricha (fron-
toventral cirri overlooked?) cannot be excluded. Detailed redescription necessary.

M 0 r p hoI 0 g y: After KAHL (1932) in life 120 11m, according to SRAMEK-HuSEK (1957)
130-180 ~m long. Elliptical, posterior end broadly to narrowly rounded, left margin
slightly indented at level of buccal vertex. Contractile vacuole slightly in front of mid-
body. No cortical granules mentioned. Behind the 3 frontal cirri only I enlarged cirrus,
 Rubrioxytricha 479

probably the buccal cirrus. After KAHL (1932) anterior cirrus of middle and inner right
marginal row possibly also enlarged. Three (KAHL 1932) or 2 (SRAMEK-HuSEK 1957) pos-
toral ventral cirri. Two pretransverse ventral cirri. Five transverse cirri arranged in hook-
shape project slightly beyond posterior end of cell. Very likely 3 slightly elongated cau-
dal cirri.

o c cur r e n c e and e colo g y: Locus classicus of this rare species is a beta- to al-
phamesosaprobic brook (Schwarzbach) in the Morava River system in Czechoslovakia
(SRAMEK-HuSEK 1957). KAHL (1932) isolated 0. kahli from "mesosapropel" water bodies,
probably near Hamburg, Germany, whereKAHL lived an worked. Feeds on rhodobacteria
and flagellates (KAlIL 1932) and diatoms (SRAMEK-HUSEK 1957).

Insufficient redescriptions

Onychodromopsis jlexilis STOKES - LUNDIN & WEST, 1963, Free living protozoa, p 68
(Fig. 223b). Remarks: Poorly figured so that identification is impossible. Natural waters
of the Upper Peninsula of Michigan, USA (see also WEST 1953 and WEST & LUNDIN
1963).

Onychodromopsis sp. (flexilis STOKES) - KUSANO, 1985, Rep. Inst. nat. Stu., 16: 107, 110,
Fig. 2b. Remarks: The scanning electron micrographs do not confirm the identification.
About 11 0-125 ~m long, obviously only 1 left cirral row. Feeds on bacteria, flagellates,
and algae. Litter layer of Mizutori-no-numa pond in Tokyo (?), Japan.

Rubrioxytricha gen. nov.

C h a r act e r i sat ion: Adoral zone of membranelies formed like a question mark.
Undulating membranes in Oxytricha pattern. Frontoventral cirri form V-shaped pattern.
Postoral ventral cirri in dense cluster behind buccal vertex. Two pretransverse ventral and
5 transverse cirri. One right and 1 left row of marginal cirri. Four or 5 dorsal kineties.
One or 2 caudal cirri. Cytoplasm usually distinctly, rarely only slightly homogeneously
orange to reddish or brownish.

T y pes p e c i e s: Oxytricha haematoplasma BLATTERER & FOISSNER, 1990.

Add i t ion a I c h a r act e r s: In life 150---260 ~m long, about 3.5-4.0 times as long
as broad. Body margins more or less parallel, both ends rounded. Flattened about 2: 1.
Very flexible, scarcely contractile. Two macronuclear nodules. Usually 2, R ferruginea
rarely with 3 micronuclei. Contractile vacuole distinctly in front of mid-body, during di-
astole with short collecting canals. Cortical granules present (see, however, taxonomy
chapter in Rferruginea). Adoral zone of membrane lIes about 1/3 of body length. Buccal
480 SYSTEMATIC SECTION

area rather small but deep. Peristomial lip anteriorly hooked to the left, similarly to in
Cyrtohymena and Notohymena; however, paroral only slightly curved. Undulating mem-
branes consist of2 rows of basal bodies each. Buccal cirrus near anterior end ofundulat-
ing membranes, about at level of right frontal cirrus, which is arranged very close to dis-
tal end of adoral zone of membranelles (Fig. 144b, arrow). Rather invariably full set (18)
of frontal-ventral-transverse cirri. Anterior frontoventral cirrus (VIJ4) slightly separate
from the other 3 cirri. Postoral ventral cirri arranged in a short longitudinal row. Trans-
verse cirri arranged in hook-shape and distinctly displaced anteriad, protrude only
slightly, if at all, beyond posterior end of cell. Dorsal cilia short, that is, in life 2-5 11m
long. Caudal cirrus (cirri) inconspicuous. Probably confmed to freshwater habitats.

T a x 0 nom y: For the description of the Oxytricha pattern mentioned in the characteri-
sation, see chapter 2 in the general section. Rubrioxytricha differs from Oxytricha mainly
in that it usually has only 1 or 2 caudal cirri and from Urosomoida in that it invariably
has 5 transverse cirri. Tachysoma has no caudal cirri. Furthermore, no species of these
taxa has a homogeneously coloured cytoplasm. In Oxytricha aeruginosa and Cyrtohy-
mena muscorum the reddish colour of the cytoplasm is induced by small granules. The
classification of Rubrioxytricha ferruginea in Steinia and Cyrtohymena was based on the
anteriorly distinctly curved peristomiallip; the undulating membranes, however, are only
slightly curved and arranged as in Oxytricha (Fig. 144b). The latter feature separates Ru-
brioxytricha from Cyrtohymena and Notohymena.
The cortical granules, the flexible body, and the loss of fragmentation of dorsal ki-
nety 3, that is, a simplification of dorsal morphogenesis (at least in R haematoplasma),
assigns Rubrioxytricha to the Oxytrichinae. This position can, however, only be eluci-
dated after the detailed description of divisional morphogenesis.

Key to species

The colours of the cytoplasm and the cortical granules are similar in both species (use
bright field for studying the colours!). Species distinction thus requires counting of dorsal
kineties (use Normarski interference contrast or protargol impregnation).

1 Cortical granules lemon-yellow, sometimes with a greenish shimmer, regularly

arranged in single rows; cytoplasm usually distinctly, rarely only slightly homoge-
neously orange to reddish coloured; 4 dorsal kineties (Fig. 144c, d) ............. .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . , R. haematoplasma (p 481)
Cortical granules brownish, irregularly arranged in broad longitudinal rows; cyto-
plasm homogeneously rusty brown; 5 dorsal kineties (Fig. 145c, d) ............ .
o •••••••••••••••••••••••••••••••••••••••••••••••• , R·ferruginea (p 484)
 Rubrioxytricha 481

Rubrioxytricha haematoplasma (BLATIERER & FOISSNER, 1990) comb. nov.

(Fig. 144a-k, Tables 3, 30)

1990 Oxytricha haematoplasma BLAITERER & FOISSNER, Arch. Protistenk., 138: 106 (Fig. 144a-h; 1 slide of
holotype specimens and 1 slide of paratype specimens are deposited in the OberOsterreichische Landes-
museum in Linz, Upper Austria).
1991 Oxytricha haematoplasma BLAITERER & FOISSNER, 1990 - FOISSNER, BLAITERER, BERGER & KOHMANN,
Informationsberichte des Bayer. Landesarntes filr Wasserwirtschatl, 1191: 287.
1993 Oxytricha haematoplasma BLAITERER & FOISSNER, 1990 - SHIN & KIM, Korean J. Zoo\., 36: 227 (Fig.
144i-k, see taxonomy).
1994 Oxytricha haematoplasma - BLAITERER, Katologe des OberOsterreichischen Landesmuseums Linz, 71:
155, Abb. 14 (colour microphotograph ofa living specimen).
1994 Oxytricha haematoplasma BLAITERE & FOISSNER, 1990 - SHIN, Dissertation, p 139 (Fig. 144i--k; incor-
rect spelling ofBLAITERER; see taxonomy; mixed up the figures of Oxytricha hymenostoma and R. hae-
matoplasma).

T a x 0 nom y: Unfortunately, SHIN & KIM (1993b) did not mention the cortical granules
and the colour of the cytoplasm. Their population has a distinctly lower number of adoral
membranelles (26 against 38; Table 30) and there are also some minor differences in the
dorsal ciliature (dorsal kinety I shortened anteriorly, kinety 4 interrupted) which are pos-
sibly a kind of geographic variation. For comparison with other coloured oxytrichids, see
the relevant chapter in Rubrioxytrichaferruginea.

M 0 r p hoi 0 g y and b i 0 log y (see also genus section): In life 120-180 x

40-55/-lm (BLATIERER & FOISSNER 1990), after SHIN & KiM (1993b) 100-180 x
50-90 /-lm. Macronuclear nodules in life up to 35 x 20 /-lm, with moderately sized nucle-
oli. Invariably 2 (n = 13, type material; n = 8, Korean population) micronuclei, in life up
to 8 x 5 /-lm. Cortical granules 0.5-1.4 /-lm across, absent around cirri (Fig. 144d). Cyto-
plasm often with many colourless, 1-5 11m sized, greasily shining globules, so that cells
appear darkish at low magnification. Moderately rapid movement, usually creeping on
detritus.
Bases of all cirri of about same size. Usually 4 (very rarely only 1; mean = 3.8; n =
18; BLATIERER & FOISSNER 1990) frontoventral cirri. One of 20 specimens with only 2
postoral ventral cirri. Usually 5, rarely 6 (mean = 5.1, n = 18, BLATIERER & FOISSNER
1990) transverse cirri, in life about 20-30 /-lm long, only posterior-most cirrus reaches
end of cell (Fig. 144a, b). Marginal cirri in life 15-20 /-lm long, marginal rows slightly
overlapping because left row terminates just on right margin. Dorsomarginal row (dorsal
kinety 4) only slightly shortened at both ends. Usually I caudal cirrus at end ofkinety 3,
somewhat elongated (Fig. 144c, h, arrow).
Morphogenesis is only incompletely known (Fig. 144f-h). However, a middle stage
shows a peculiarity in comparison to other taxa of the Oxytrichinae, namely cirrus V/3
(posterior postoral ventral cirrus) is not involved in primordia formation (Fig. 144f,
arrow). This would assign R haematoplasma to the Stylonychinae (Fig. 25a, Tables 4,
5). It cannot be excluded that this detail in Figure 144fis a misobservation because cirrus
 482 SYSTEMA TIC SECTION

(ill)
I ~\ ,/ 1)
. t:;)
\~ \' /
~ J}
\
~ ~ / J
~ ,
~/
,;
,; 1
t#
t# \ /
\
t#
t#
t#
/
~
t#
/
~
;
; \ \ /
~ ;
.,. \, \ /
)
\
~
~
.,.,; )
~
f
/
~
~
•• f \., ~

",
~

.
~ "-, /
!J
I
, "-, .iJ
# "-'(, ./

...---.
\,
# ---,
I I,," ~
-, ,/
b C

d
c e

Fig. 144a-e Rubrioxytricha haematoplasma (from BLAlTERER & FOISSNER 1990. a, d, e, from life; b, c, protar-
gol impregnation). a) Ventral view, 170 Jlm. b, c) Ventral and dorsal infraciliature, b, c = 100 Jlm. Arrows
mark right frontal cirrus and caudal cirrus. d) Ventral view. The lemon yellow, about 0.5-1.4 Jlm sized cortical
granules are arranged in longitudinal rows. e) Lateral view; p 481.

V/3 is displaced distinctly posteriad, whereas all postoral ventral cirri are arranged rather
close to the buccal vertex during interphase (Fig. 144b). Detailed morphogenetic studies
are needed to check the behaviour of this cirrus.

o c cur r e n c e and e colo g y: Common in the aufWuchs and detritus of freshwa-

ter habitats, especially in running waters. Locus classicus is the Amper River, Bavaria,
Germany, about 1 km below a lake (Ammersee). According to BLATIERER & FOISSNER
(1990) and FOISSNER et al. (1992b) this slow flowing river is strongly eutrophic and beta-
 Rubrioxytricha 483

.. ' - • • I

(j

""

..,,'
0
o

• •••
••....
i
~
;

,, . ':
,,
,

! ! .O~~'
,"
'.
.
.
".

o
0

0 0
q,0
0
• 0
h
0° ~

Fig. 144f-h Rubrioxytricha haematoplasma (from BLATfERER & FOISSNER 1990. f-h, protargo! impregnation).
f) Ventral infraciliature of a middle morphogenetic stage, 105 )lm. The arrow marks the (unchanged) parental
postoral ventral cirrus V/3 which is usually involved in primordia formation in the Oxytrichinae (see
taxonomy); possibly a misobservation. g, h) Ventral and dorsal infraciliature of a late morphogenetic stage,
130 )lm. Arrow marks the single caudal cirrus at posterior end of dorsal kinety 3 of proter; p 481.

to alphamesosaprobic. AOOLR (1996, 1996a, b, 1997, 1997a, b), BLATIERER (1994), FOISS-
NER (l997a), FOISSNER & MOOG (1992) and FOISSNER et al. (1991, 1992a) found Rubrioxy-
tricha haematoplasma in mesosaprobic rivers and brooks in Upper Austria and Bavaria.
BLATIERER (1989) observed it in the hypertrophic pond at Salzburg University, Austria. I
found it, for instance, in the Glasenbach, an oligo- to betamesosaprobic brook near the
city of Salzburg and in a slow flowing part of the alpha- to betamesosaprobic Salzach
River in Salzburg, Austria. SHIN & KIM (1993a, b) isolated R haematoplasma from the
Han River, Seoul, Korea, in summer.
Feeds on Oscil/aloria sp., diatoms (Synedra ulna), coccale green-algae, Euglena spi-
rogyra, E. viridis, testaceans (Trinema sp.), and ciliates (Chlamydonella sp.). Biomass of
106 individuals about 80 mg (FOISSNER et al. 1991).
 484 SYSTEMATIC SECTION

\.

\.
\. I
\.

0:
\. \. J
\. \. )
\.
\. ,I

\. )

\. ,)
•
.
fI
./
,J
,
•
, #
J
# , • ,)

," . I.
• k

Fig. 144i-k Rubrioxytricha haematoplasma (from SHIN & KIM I993b. i, from life; j, k, protargol impregna-
tion). i) Ventral view, 130 11m. j, k) Ventral and dorsal infraciliature, 135 11m; p 481.

Rubrioxytricha haematoplasma is classified as beta- to alphamesosaprobic indicator

of water quality: b-a; b = 6, a = 4, 1= 3, SI = 2.4 (ForSSNER et al. 1991). This has to be
proved by further autecological and saprobiological investigations on reliably determined
populations.

Rubrioxytrichaferruginea (STEIN, 1859) comb. nov. (Fig. 145a-o, Table 30)

1859 Oxytrichaferruginea STEIN, Organismus der lnfusionsthiere I, p 187 (Fig. I45t: g).
1862 Oxytrichaferruginea STEIN - ENGELMANN, Z. wiss. Zoo\., II: 365 (Fig. 145k).
1865 Oxytrichajerruginea - QUENNERSTEDT, Acta Univ. lund., 2: 59 (Fig. 145h).

Fig. 145a-e Rubrioxytrichaferruginea (from SONG & WILBERT 1989. a, d, from life; b, c, e, protargol impreg- -)
nation). a) Ventral view, 205 11m. b, c) Ventra! and dorsal infraciliature, b, c = 135 11m. The arrow marks the 3
posterior frontoventral cirri. d) Posterior portion of cell in dorsal view. The cortical granules are arranged ir-
regularly in longitudinal rows. e) Ventral infraciliature of anterior portion, bar = 50 11m. BC = buccal cirrus,
CC = caudal cirri, e = endoral, p = paroral, 1 = dorsal kinety 1.
 Rubrioxytricha 485

!
1 ' ,II i
- --I ' /
\ (I
'\ . \

~
,,-' '-
,,
( ,
, I
~ \

, , , , ,I
, - -,-"

,
,
,, r
, !
, I
,
, ,/
, .-
\'t .-
\.,
'-, '\ (
\.
j
/

I
, • '. ( /

145a

~ I \ ..,

:11 e
486 SYSTEMATIC SECTION

Table 30 Morphometric data of Rubrioxytrichajerruginea (fer, from SONG & WILBERT 1989), Rubrioxytricha
haematoplasma (hal, from BLATTERER & FOISSNER 1990; ha2, from SIllN & KIM 1993b), and Pseudostrombid-
ium planctonticum (PIa, from WILBERT 1986a). All data are based on protargol-impregnated material (PIa, after
WILBERT'S 1975 method). All measurements in micrometres. CV = coefficient of variation (in %), Max =
maximum value, mean = arithmetic mean, Min = minimum value, n = sample size, SD = standard deviation

Character Species mean SD CV Min Max n

Body, length fer 143.4 12.9 9.0 122 162 10
hal 113.3 16.4 14.5 91 143 19
ha2 133.2 18.1 13.6 112 160 9
pIa 223.8 18.1 8.1 194 267 20
Body, width fer 47.2 3.9 8.3 39 51 10
hal 41.4 6.8 16.6 29 51 19
ha2 62.8 13.6 21.8 42 80 9
pIa 86.0 11.2 13.0 67 108 20
Macronuclear nodule, length hall 15.9 3.2 20.4 11 21 19
ha2 19.6 1.9 9.6 17 22 9
pIa 44.9 7.7 17.1 32 52 20
Macronuclear nodule, width hall 8.6 1.3 15.2 7 12 19
ha2 10.7 l.l 10.5 9 12 9
pia 19.7 3.7 18.8 16 26 20
Macronuclear nodules, distance hal 9.8 4.2 43.0 0 21 19
between ha2 0
pIa 21.9 2.8 12.8 16 24 20
Micronucleus, length hall 4.2 0.7 17.2 3 5 13
ha2 3.3 0.5 15.0 3 4 9
pIa 9.6 1.9 19.8 7 12 10
Micronucleus, width hall 3.6 0.7 18.8 3 5 13
pIa 6.6 0.8 12.1 6 8 10
Adoral zone of membranelles, length fer 45.1 3.8 8.5 40 49 7
hal 35.8 4.0 11.0 30 46 19
ha2 44.0 6.1 13.9 34 51 9
pIa 111.4 8.9 8.0 98 125 17
Adoral membranelles, number fer 38.3 1.6 4.2 35 41 8
hal 38.4 4.3 11.2 30 48 19
ha2 25.8 3.5 13.6 21 30 10
pIa 51.2 2.4 4.5 48 55 17
Right marginal row, number of cirri fer 32.7 3.3 10.2 28 38 12
hal 33.4 3.6 10.9 25 39 18
ha2 31.0 3.0 9.6 27 35 10
pIa 26.7 2.2 8.2 20 30 15
Left marginal row, number of cirri fer 33.0 1.6 4.9 31 36 12
hal 35.8 3.9 11.0 28 45 18
ha2 32.3 2.4 7.4 27 35 9
pIa 20.8 1.0 4.8 20 22 15
Dorsal kineties, number fer 5.0 0 0 5 5 11
hal 4.0 0 0 4 4 15
ha2 4.1 0.4 8.6 4 5 8
pIa 6.0 6 6 20
 Rubrioxytricha 487

Table 30 Continued

Character Species mean SD CV Min Max n

Caudal cirri, number fer 1.7 0.6 92.1 1 2 15
hal 0.8 0.4 44.5 0 1 19
ha2 0.7 0.5 69.0 0 10
pla2 6.5 0.9 13.8 5 8 20
pla2 4.8 1.0 20.8 3 6 20
pla2 4.1 0.9 21.9 3 6 20

I Anterior one.
2 Caudal cirri on dorsal kineties 1 (fourth line), 2 (fifth line), and 4 (sixth line).

1866 Steiniajerruginea DIESING - DIESING, Sber. Akad. Wiss. Wien, 53: 114.
1901 Oxytrichajerruginea ST. - Roux, Mem. lust. nato. genev., 19: 102 (Fig. 1450).
1908 Oxytricha jerrujinea STEIN - BORGER, An. Univ. Chile, 122: 187 (Fig. 145m; incorrect subsequent
spelling).
1932 Steinia (Oxytricha) jerruginea STEIN, 1859 - KAlIL, Tierwelt Otl., 25: 612 (Fig. 145i).
1961 Steiniajerruginea STEIN, 1859- RElITER, Actazool. fenn., 99: 19 (Fig. 145j).
1968 Oxytricha (Steinia) jerruginea STEIN, 1859 - CHORIK, Free-living ciliates, p 137 (Fig. 1451).
1972 Oxytrichajerruginea STEIN, 1859 - BORROR, J. Protozool., 19: 13.
1974 Oxytricha jerruginea STEIN, 1859 - JONES, Univ. South Alabama Monogr., 1: 41 (Fig. 145n).
1989 Cyrtohymena jerruginea (STEIN, 1859) - FOISSNER, Sber. Ost. Akad. Wiss., 196: 239.
1989 Oxytrichajerruginea STEIN, 1859 - SONG & WILBERT, Lauterbomia, 3: 162 (Fig. 145a--e).
1991 Oxytrichajerruginea STEIN, 1859 - FOISSNER, BLATIERER, BERGER & KOHMANN, Informationsberichte
Bayer. Landesamtes filr Wasserwirtschaft, 1/91: 283.

Nom e n c I at u rea n d t a x 0 nom y: KAHL (1932) classified Steinia as subgenus

of Oxytricha; the correct name in his revision is thus Oxytricha (Steinia) ferruginea. In-
correct subsequent spellings: Oxytrichaferuginea (SONG & WILBERT 1989, P 219); Oxy-
tricha terruginea STEIN (ALlEv 1982a, P 807).
STEIN (1859b) stated that this species is easily recognisable by its body shape and the
more or less intense rusty colour of the cytoplasm. He assumed that the colour is due to
digestion products and did not mention a cortical granulation. It is hardly imaginable (but
not impossible!) that he overlooked the granules because he described them exactly in
Urostyla grandis and Paraurostyla weissei. By contrast, KAHL (1932) stated that the
brown colour is caused by granules lying close underneath the pellicle and in the cyto-
plasm. SONG & WILBERT (1989) described a (homogeneous?) brownish colour and a corti-
cal granulation. That means, if a population without cortical granulation is found a new
species has to be established for the KAHL (1932) and SONG & WILBERT (1989) popula-
tions. Each future record of this problematic species should contain a remark about the
colour of the cytoplasm and whether cortical granules are present or not! The marine
population described by WANG & NIE (1932, P 360; Fig. 145p) is probably not R ferrugi-
nea because the cytoplasm is yellowish and the cirri are differently arranged (see insuffi-
cient redescriptions). The redescription in PENARD (1922, Fig. 223e-g) is insufficient.
488 SYSTEMATIC SECTION

Fig. 145f-l Rubrioxytricha jerruginea «( g, after STEIN 1859b; h, after QUENNERSTEDT 1865; i, from KAHL
1932; j, after REUTER 1961; Ie, from ENGELMANN 1862; 1, from CHORIK 1968. f-l, from life). f-j, I) Ventral
views, f-h = size not indicated, i = 250 )lm, j = 160-180 )lm, 1 = 150 )lm. Note that in «( i, j) an increased
number of postoral and pretransverse ventral cirri is shown. k) Conjugation, 200 )lm; p 484.

The infraciliature of Rubrioxytricha jerruginea differs from that of the type species
mainly in the number of dorsal kineties (5 against 4); consequently, dorsal morphogenesis
must also be different. Rubrioxytricha haematoplasma shows the Urosomoida pattern,
that is, no fragmentation of kinety 3 and only 1 dorsomarginal kinety occur (Fig. 144g,
h). Conversely, in R jerruginea the 5 dorsal kineties vel)' likely originate according to
the Oxytricha pattern, that is, fragmentation ofkinety 3 and I dorsomarginal row occur.
Rubrioxytricha spp. remind on Oxytricha aeruginosa and Cyrtohymena muscorum;
however, in these species the reddish colour of the cytoplasm is not homogeneous, but
caused by granules. Furthennore, C. muscorum has an anteriorly distinctly curved paroral
and lives in terrestrial habitats only.
 Rubrioxytricha 489

Fig. 14501-0 Rubrioxytrichaferruginea (m, from BORGER 1908; n, from JONES 1974; 0, from Roux 1901).
Ventral views from life, m = size not indicated, n = 160 11m, 0 = 170 11m; p 484.
Fig. 145p Oxytrichaferruginea sensu WANG & NIE (1932). Ventral view from life, 150 11m; p 491.

M 0 r p hoi 0 g y: According to STEIN (1859) large specimens 170 Jim long; length after
SoNG & WILBERT (1989) about 150-200 Jim, after KAHL (1932) 180-260 Jim. Margins
sometimes slightly concave (Fig. 145t). Cortical granules (not described by STEIN 1859b;
see taxonomy) in life about 1 Jim sized, colour not mentioned by SONG & Wn..BERT
(1989), according to KAHL (1932) they cause a brownish colour. Frontal and transverse
cirri distinctly enlarged. Marginal rows not overlapping posteriorly. All 5 dorsal kineties
of about body length indicating that only 1 dorsomarginal row is formed.

o c cur r e n c e and e colo g y: According to the numerous records, which are,

however, not substantiated by morphological data, widely distributed but only moderately
frequent in the aufwuchs and detritus of freshwater habitats; also inhabits brackish and
marine waters. Obviously prefers stagnant and slow running oligosaprobic and be-
tamesosaprobic waters.
490 SYSTEMATIC SECTION

Locus classicus is a brook (puftbach) at Niemegk, Brandenburg, Germany, where

STEIN (1859b) discovered it in Oscillatoria tufts. Furthermore, he found R ferruginea
rather frequently in similar habitats near Prague, Czechoslovakia. Records documented
by illustrations: brackish water (salinity 0.5 %) of a shallow bay (Kalvassa) in the Baltic
Sea, Finland (REUTER 1961); near Leipzig, Germany (ENGELMANN 1862); on plants of
rather clean ditches and in non-putrefying detritus of large water bodies, probably in Ger-
many (KAlIL 1932); rarely in a eutrophic pond (poppelsdorfer Weiher) in Bonn,
Germany, in August (SONG & Wll.BERT 1989); Sweden (QUENNERSTEDT 1865); ponds and
Lake Geneva, Switzerland (Roux 1901; see also ANDRE 1912, 1916, BOURQUIN-LINDT
1919, FOREL 1904); small water bodies and cooling plant in Moldova (CHoRIK 1968, CHO-
RIK & VIKOL 1973); at a salinity of 0.2-3.0 % in Mobile Bay, Alabama, USA (JONES
1974); Chile (BORGER 1908). Records not documented by illustrations: mosses from
springs near Vienna, Austria (KOlIN 1940); pool with ground water near Eisenstadt, Aus-
tria (SPANDL 1926a); Belgium (BERVOETS 1940); freshwater mosses and rain puddle in
Czechoslovakia (TIRJAKovA & MATIS 1987a, SPANDL 1926a); near Perth, England
(CRAIGIE 1921); pond with mosses in Estonia (JACOBSON 1928); brackish and sea-water in
Finland (LEVANDER 1894a, b, 1901); aufwuchs of mesottophic and eu- to polytrophic
Eifelmaar lakes, Germany (PACKROFF 1992, PACKROFF & Wll.BERT 1991); Latvian rivers
(LIEPA 1978, 1990); freshwater and brackish water in the Netherlands (VERSCHAFFELT
1930); rain-wet mosses from Espegrend, Norway (REUTER 1963a); littoral of Lake Majeur
and up to 100 indo I-I in Swiss ponds in September (ANDRE 1915, RIGGENBACH 1922);
Ukrainian pond during winter (KovALCHUK & SVINTSOVA 1994); rivers in the USSR (NEIS-
WESTNOWA-SHADINA 1935, SASSUCHIN 1930, SASSUCHIN et al. 1927, SHADIN et al. 1931);
running waters in Azerbaijan (ALlEv 1982a); saline lake near the coast of the Black Sea
(DAGAJEVA 1930); River Plate, Argentina (CARBONELL 1935); Brazil (CUNHA 1913).
Terrestrial records unreliable (FANTHAM & PATERSON 1926, HEINIS 1959, TIRJAKovA &
MATIS 1987a); probably, confused with Notohymena rubeseens or Cyrtohymena museo-
rum, which are rather common in edaphic mosses and soil.
Feeds on bacteria, cyanobacteria, and diatoms. Biomass of 106 individuals about
125 mg (FOISSNER et al. 1991). Swims with about 400 ~m S-I (BULLINGTON 1925). MAnoNi
& GHEm (1981a) recorded Rubrioxytriehaferruginea in a betamesosaprobic brook in
northern Italy under the following conditions: SI = 2.1, 23-25 °C, pH 8.1-8.4, 700 to
800 ~S cm- I specific conductivity (25°C), 8.1-11.8 mg 1-1 0 2, 19-21 mg I-I COD, 0.2 to
0.38 mg I-I PO/-P, 0.16-133 mg I-I NHl, 0.63-1.6 mg I-I NOl'-N, 0.02-{).03 mg 1.1 N02--
N, 40-43 mg I-I CI-. MATIS (1975) found Rubrioxytriehaferruginea during the summer in
running waters in Czechoslovakia at low oxygen levels (1.2-2.6 mg I-I O2, pH 6.7-6.9,
12.5-17 0q. NOLAND (1925) reported it from various freshwater habitats in Wisconsin at
4.5-28.5 °C (mean = 15.6 °C; n = 5), pH 7.5-9.8 (PH 8.2), 4.1-9.9 mg}-I O2 (6.9 mg I-I),
and 53-174 % O2 saturation (99 %).
Rubrioxytrieha ferruginea is classified as oligosaprobic indicator (FOISSNER et al.
1991, SLADECEK et al. 1981, WEGL 1983): 0; 0 = 7, b = 3, 1=4, SI = 1.3. This has to be
verified on reliably determined populations.
 Parurosoma 491

Insufficient redescriptions

Oxytrichaferruginea STEIN 1859 - PENARD, 1922, Etudes Infusoires, p 237 (Fig. 223e-g).
Remarks: In life 125-150 ~m long. Possible a Holosticha or Pseudokeronopsis species.
Terrestrial(?) mosses in Switzerland.

Oxytrichaferruginea STEIN 1859 - WANG & NIE, 1932, Contr. bioI. Lab. Sci. Soc. China,
8: 360 (Fig. 145p). Remarks: In life 150 x 35 ~m. Body linearly oblong, about five times
as long as broad, widest at mid-body or slightly in front of posterior end, bluntly pointed
anteriorly, rounded posteriorly. More flexible than freshwater population. Two macronu-
clear nodules, one in front of contractile vacuole, the other centrally or sub-centrally.
Contractile vacuole without canals. Cytoplasm deeply, homogeneously yellowish. Buccal
area rather wide, about one third of body length; undulating membranes (obviously par-
oral) curved to the left anteriorly. 18 frontal-ventral-transverse cirri arranged as shown in
Figure 145p; frontal and transverse cirri distinctly enlarged, transverse cirri not or only
slightly projecting beyond posterior end of cell. Marginal cirri uninterrupted, projecting
posteriorly only. Feeds on diatoms and other marine algae. Rather abundant in the sea
nearby Amoy, China
According to WANG & NIE (1932), this population may be recognised as a marine va-
riety of Rubrioxytricha ferruginea. It differs from the type population in that it has a
deeply, homogeneously yellowish tint which definitely did not result of the temporary re-
tention of coloured food particles. Detailed redescription necessary.

Parurosoma GELEI, 1954

1954 Parurosoma GELEI, Acta bioI. hung., 5: 332 - Type (by monotypy): Holosticha (Parurosoma) dubium
GELEI, 1954.
1997 Parurosoma GELEI, 1954 - BERGER & FOISSNER, Arch. Protistenk., 148: 148.

C h a r act e r i sat ion: Adoral zone of membrane lIes, undulating membranes, and
frontoventral cirri similar to in Urosoma. Postoral ventral cirri in dense cluster behind
buccal vertex. Two pretransverse ventral and five transverse cirri. Two right and 2 left
rows of marginal cirri. Four, sometimes 5 dorsal kineties. Caudal cirri present. Dorsal
morphogenesis very likely in Urosomoida pattern.

T a x 0 nom y: The characterisation is according to BERGER & FOISSNER (1997). For a

detailed description of patterns, see chapter 2 in the general section. Parurosoma is
monotypic, that is, includes only the type species. GELEI (l954b) classified Parurosoma
as subgenus of Holosticha, that is, Holosticha (parurosoma); thus, CORLISS (1979) and
TUFFRAU (1987) included it in the Holostichidae. TUFFRAu & FLEURY (1994, P 141) classi-
fied Parurosoma without foundation in the Amphisiellidae. However, according to the
492 SYSTEMATIC SECTION

ventral infraciliature it is a typical member of the Oxytrichidae. This was already recog-
nised by BORROR (1972a), who synonymised Parurosoma with Pleurotricha, which has,
however, a rigid body and lacks caudal cirri.
Parurosoma tekirghiolica mentioned in TUCOLESCU (1965, P 178) is obviously a no-
men nudum, that is, a new species without description. Very likely, the same holds for
Leptocirra tekirghiolica (TUCOLESCU 1965, P 178), a new genus name not mentioned in
CORLISS (1979).

Single species

Parurosoma dubium (GELEI, 1954) comb. nov. (Fig. 146a-c)

1954 Parurosoma dubium GELEI, Acta bioI. hung., 5: 332 (Fig. 146a; for the correct name in the original de-
scription, see nomenclature).
1954 Holostichamononucleata GELEI, Acta bioI. hung., 5: 327 (Fig. 146b, c).
1972 Pleurotricha dubium (GELEI, 1954) - BORROR, J. Protozool., 19: IS.
1972 Pleurotricha mononucleata (GELEI, 1954) - BORROR, J. Protozool., 19: 15.
1974 Paruroleptus dubium GELEI - STILLER, Fauna Hung., liS: 86, 87.
1974 Pleurotricha mononucleata (GELEI, 1954) - STILLER, Annis hist.-nat. Mus. natn. hung., 66: 132.

Nom en c I at u rea n d t a x 0 nom y: Parurosoma was classified as subgenus of

Holosticha (GELEI 1954b); the correct name in the original description is thus Holosticha
(Parurosoma) dubium. It was never formally transferred to Parurosoma, which is here
undertaken. Incorrect spelling: Porurosoma dubio (GELEI 1954b, P 342).
The linear arrangement of the frontoventral cirri is reminiscent of Urosoma, indicat-
ing that they are sister-groups which differ in the number of marginal rows. Detailed re-
description necessary.
GELEI (1954b) found Holosticha mononucleata together with the rather fragile Paru-
rosoma dubium; I thus assume that H. mononucleata is simply the anterior portion of P.
dubium. This is indicated especially by the corresponding infraciliature, the position of
the contractile vacuole, and the single macronuclear nodule. Furthermore, many speci-
mens are pointed at the right posterior cell portion, which is typical of hypotrich frag-
ments. PETZ & ForssNER (1996) assumed that H. mononucleate is a reorganiser. BORROR
(l972a) transferred H. mononucleata to the Pleurotricha; obviously erroneously, STILLER
(l974a) repeated this nomenclatural act.

M 0 r p hoi 0 g y and b i 0 log y: In life 150-250 x 50-60 /lm. Anterior end

rounded, posterior portion with an "immovable" tail-like process. Moderately flexible.
Rather fragile, that is, bursts rather easily in thin water-layers. Two ellipsoidal macronu-
clear nodules, about 20 /lm long with large spherical nucleoli; each nodule with a single
micronucleus usually at distal, rarely at proximal end. Contractile vacuole with short col-
lecting canals, located at end of anterior third of cell. Cytoplasm near cytostome and at
 Parurosoma 493

Fig. i46a-c Parurosoma dubium (from GELEI

1954b; formaldehyde-sublimate fixation). a) Ven-
tral infraciliature, 215 Ilm. b, c) The synonym,
Holosticha mononuc/eata, in ventral and dorsal
view; very likely it is only the anterior portion of the
rather fragile Parursoma dubium; p 492.

base of tail granulated grey-brownish by

pigments (cortical granules?) and cyto-
plasmic crystals; after fIXation with
formaldehyde-sublimate entire cell shows
a "cortical granulation"(?). Usually cree-
ping but also swimming; often stands still
for long periods.
Adoral zone of membranelles ap-
proximately 25 % of body length, about
30 adoral membrane lIes. Buccal field nar-
row. Usually 9, rarely 8 frontoventral
cirri, that is, 3 frontal cirri, 1 buccal
cirrus, 2 cirri behind right frontal cirrus
(that is, one more than in Urosoma), and
3 cirri right behind them; however, protar-
gol impregnation is needed to ascertain
correct cirral pattern. Three postoral and
2 pretransverse ventral cirri. Invariably 5
fringed transverse cirri at base of tail-like
process; usually they protrude beyond left
body margin. Outer right marginal row
with 35-50, inner right marginal row with
20 (up to 31) cirri. 34-45 cirri in outer
left marginal row. Three dorsal kineties of
body length, 1 or 2 dorsomarginal kine-
ties with only 4-5 basal body pairs. Dor-
sal cilia "short", that is, about 2-4 f.1m.
One, rarely 2 caudal cirri.

o c cur r e n c e and e col 0 g y: Rare species. Locus classicus is an ephemeral, hy-

pertrophic pond ("Tiimpel I") on a meadow in the Borzsony Mountains, Hungary, where
GELEI (l954b) discovered Parurosoma dubium in the benthal and in dry mud from the lit-
toral. Furthermore, he found it in the pelagial of another pond ("Tiimpel V") in the same
area. For detailed description of these habitats and synecological data, see GELEI (1954a)
494 SYSTEMATIC SECTION

and GELEI et al. (1954). Feeds mainly on flagellates, rarely on diatoms and unicellular
green algae (GELEI 1954b, GELEI et aI. 1954). No further records published.

Pseudostromhidium HORVATH, 1933

1933 Pseudostrombidium HORVArn, Arch. Protistenk., 80: 291 - Type (by monotypy): Pseudostrombidium
planctonticum HORVAnJ, 1933.
1997 Pseudostrombidium HORVArn, 1933 - BERGER & FOISSNER, Arch. Protistenk., 148: 148.

C h a r act e r i sat ion: Adoral zone of membraneIIes formed like a question mark.
Undulating membranes almost straight and side by side. Frontoventral cirri in V -shaped
pattern. Postoral ventral cirri behind buccal vertex. Two pretransverse ventral and 5
transverse cirri. One right and I left row of marginal cirri. Six dorsal kineties. Number of
caudal cirri distinctly increased (>4). Dorsal morphogenesis in Oxytricha pattern.

T a x 0 nom y: The characterisation is according to BERGER & ForssNER (1997). For a

description of the Oxytricha pattern of dorsal morphogenesis, see chapter 2 in the general
section. Further details on taxonomy, see single species P. planctonticum.

Single species

Pseudostromhidiumplanctonticum HORVATH, 1933 (Fig. 147a-n, Table 30)

1933 Pseudostrombidium pianctonticum HORVAlH, Arch. Protistenk., 80: 291 (Fig. 147a-g).
1935 Urosoma (fiIr Parastrombidium) planctonicum J. HORVArn, 1933 - KAm., Tierwe1t Otl., 30: 841 (incor-
rect subsequent spelling, see also taxonomy).
1972 Parastrombidium pianctonicum HORVArn, 1933 - BORROR, J. Protozool., 19: 13 (incorrect subsequent
spelling).
1972 Urosoma (= Parastrombidium) pianctonicum (HORVAnJ 1933), KAm. 1935 - BlcK, Binnengewasser,
26: 76 (incorrect subsequent spelling).
1986 Oxytricha pianctontica (HORVAlH, 1933) - WILBERT, Arch. Protistenk., 131: 67: (Fig. 147h-j).
1994 Urosoma planktonicum - SKIBBE, Arch. Hydrobiol., 130: 346 (Fig. 147k, I; incorrect subsequent spelling).
1999 Pseudostrombidium planctonticum HORVArn, 1933 - FOISSNER, BERGER & SCHAUMBURG, Informations-
berichte des Bayer. Landesamtes fUr Wasserwirtschaft, in press (Fig. 147m, n).

Nom e n cIa t u rea n d t a x 0 nom y: KAHL (1935) obviously confused Pseu-

dostrombidium with Parastrombidium FAURE-FREMIET, 1924, an oligotrich. Many incor-
rect subsequent spellings: Oxytricha planctonica (FOISSNER & O'DONOGHUE 1990, P 692;
WILBERT 1986a, p 67); Urosoma pelagica (WILBERT 1969, P 459; probably he mixed up
the names Urotricha pelagica and Urosoma planctonicum); Urosoma planctonica HOR-
VATII (STILLER 1974b, P 115, 116); Urosoma planctonicum J. HORVATII (WILBERT 1969,
P 458); Urosoma planctonisum HORV. (LEPSI 1965, p 975); Viosoma planctonicum (HOR-
WATII, 1933) (DRAGESCO 1970, p 129).
 Pseudostrombidium 495

Monotypic, that is, includes only the euplanktonic type species. BORROR (l972a) syn~
onymised P. p/anctonticum with Urosoma acuminata, which is clearly different (see
there). It should not be confused with Oxytricha /ongicirrata, which has only three caudal
cirri and only one micronucleus between the two macronuclear nodules. According to the
original description and the redescription by WILBERT (1986a), the undulating membranes
of Pseudostrombidium are rather straight and do not intersect, similarly to in Stylonychia
(Fig. 147a, h). However, the micrographs of a protargol-impregnated specimen in SKIBBE
(1994) indicate an oral apparatus almost in the Oxytricha pattern. Pseudostrombidium
differs from Stylonychia spp. not only by the supple body but also by the significantly
higher number of caudal cirri.
The phylogenetic relationships of Pseudostrombidium are unknown. The flexible
body (HORVATH 1933) assigns it to the Oxytrichinae. By contrast, the long adoral zone of
membranelles, the straight undulating membranes (see, however, above), and the some-
what posteriorly displaced postoral ventral cirrus V/3 (Fig. 147h) support a classification
within the Stylonychinae. A further investigation (flexibility of body; pattern ofundulat-
ing membranes; exact arrangement of cirri; symbiotic algae and cortical granules absent
or not, see below) and a morphogenetic study are needed to clarify the systematic posi-
tion of this curious oxytrichid.

M 0 r ph 0 log y and b i 0 log y: After HORVATH (1933) in life about 120-160 11m
long, according to WILBERT (1986a) after protargol impregnation 194-267 x 67-108 11m;
broadest at adoral zone of membranelIes. Outline variable (Fig. 147c-g), usually anterior
end broad, posterior end very narrowly rounded, that is, margins taper towards posterior
end. Ventral surface only flattened in area of adoral zone of membranelIes, posterior por-
tion ± round in cross-section. Body flexible (HORvATH 1933). Invariably (n = 20) 2 mac-
ronuclear nodules with one micronucleus each. Contractile vacuole at about level ofbuc-
cal vertex; slightly behind a smaller vacuole which discharges somewhat earlier
(HORvATH 1933). Cytoplasm after HORVATH (1933) greyish, filled with numerous large
food vacuoles. He also mentioned uniformly distributed symbiotic algae ("Zoochlor-
ellen") between food vacuoles and in distal portion of cell; however, Pseudostrombidium
is pelagic, so that it cannot be excluded that HORVATH observed specimens with ingested
coccale planktonic algae; WILBERT (1986a) did not describe symbiotic algae. Slowly
swimming under rotation (from right to the left) about longer axis of cell, always ascend-
ing or descending. During swimming adoral zone of membranelIes turned up; only rarely
continuously creeping.
Adoral zone of membrane lies about 45-50 % of body length. Buccal area small and
flat. Undulating membranes almost straight and arranged in parallel, similarly to in Stylo-
nychia (Fig. 147a, h); however, in the micrograph by SKIBBE (1994; Fig. 147k) the undu-
lating membranes seem to be curved as in Oxytricha. Frontal, ventral, and transverse cirri
slightly to distinctly enlarged. Left and middle frontal cirrus at about same level, right
frontal cirrus just behind distal end of adoral zone of membranelles. Invariably (n = 20)
18 frontal-ventral-transverse cirri; arrangement of cirri (for example, frontoventral cirri
 .;:..
'-0
0'1

ml:;::!::::::;;::::"~::~::::::::::::

\ ...

\ "
I , '
,/ / \ ............
'" , ...
, .I ,
/
"" '""-
I "- ,
I "
/
I '> "- CIl
I I ' ....:::
" "- CIl
I I ' ...,
"- "
, ' , ' tTl
I \
'- "-
I '
I ...
\J, ~...,
/
(1
I I
I
-
CIl
tTl
(1
...,
- ,,~,- 0
-
z

h
rF.cc
Fig. 147a-j Pseudostrombidium planctonticum (a-g, from HORvArn 1933; b-j, from WILBERT 1986a. a, sublimate-toluidinblue stain; b, opalblue stain after BRESSLAU;
c-g, from life; h, i, composite from life and after protargol impregnation; j, protargol impregnation). a, b) Ventral and dorsal view, 190-200 11m. Very likely, HORvArn
did not recognise the dorsal pattern correctly. c-g) Shape variants in ventral, lateml, and dorsal view. h, i) Ventral and dorsal infraciliature, h, i = 200 11m. Arrangement
of posterior frontoventral cirri and posterior postoral ventral cirrus very likely not exactly shown (cp. Fig. 147k). j) A single adoral membranelle from middle portion of
adoral zone; CC = caudal cirri; p 494.
 Pseudostrombidium 497

Fig. 147k, 1 Pseudostrombidium planctonticum (from SKIBBE 1994 and original of same population, kindly
supplied by O. SKIBBE. k, I, protargol impregnation). Ventral and dorsal infraciliature and nuclear apparatus.
Arrows in (k) mark frontoventral cirri IIII2 and IV/3, arrowhead denotes posterior postoral ventral cirrus (V/3).
The location of these cirri matches the original description (Fig. 147a) but differs slightly from WILBERT'S illus-
tration (Fig. 147h). Arrowheads in (I) mark caudal cirri on posterior end of dorsal kinetics 1,2, and 4; p 494.

Hm and IV/3 and postoral ventral cirrus V/3) in Figure 147h possibly not exactly shown,
because slightly different from original description (Fig. 147a) and SKIBBE'S micrograph
(Fig. 147k). Posterior postoral ventral cirrus (cirrus V/3) displaced slightly (Fig. 147a, k)
to distinctly (Fig. 147h; likely a preparation artefact) posteriad. After HORVATH (1933)
posterior-most 2 transverse cirri protrude beyond posterior end of cell, after WILBERT
(1986a) obviously no transverse cirrus protrudes (Fig. 147a, h). Right marginal row be-
gins at level of right frontal cirrus. Bases of anterior-most 2 cirri of left marginal row
slightly enlarged (4-5 kineties, all other marginal cirri consist of2 kineties only; WILBERT
498 SYSTEMATIC SECTION

1986a). Dorsal cilia in life about 2 ~m long; after HORVATH (1933) 5 dorsal kineties of
body length, after WILBERT'S (1986a) table also only 5 rows (misleadingly designated as
"Zahl der Dorsalcirren"); however, the WILBERT illustration unequivocally shows the nor-
mal pattern with 6 dorsal kineties (Fig. 147i); very likely, HORVATH could not recognise
the exact pattern with the method used (147b). Caudal cirri form a rather dense corona
around posterior end of cell because 3-8 cirri on each dorsal kinety (Fig. 147a, h, I-n).
Cyst globular with smooth wall.
Morphogenesis commences near the left transverse cirrus. Later, 5 longitudinal anla-
gen (probably streak I not counted!) originate (WILBERT 1986a).

o c cur r e n c e and e colo g y: Rare species, pelagic, prefers probably eutrophic

water bodies. Abundant occurrence is possibly confmed to cold seasons, that is, winter
and spring (algal blooms?). Locus classicus is Szeged, Hungary, where HORVATH (1933)
discovered Pseudostrombidium planctonticum in the drain of the Cserepesisorer Pond at
6 °C in late March, together with Litonotus lamella, Chilodonella uncinata, and Stentor
roeselii. WILBERT (1969, 1972, 1986a) found Pseudostrombidium planctonticum (about
1000 ind.l· l ) in a eutrophic pond (Poppelsdorfer Weiher) in Bonn, Germany, only from
February to April. SKIBBE (1994) gave no site; probably he found it in north Germany. Dr.
HABERMEHL (Ansbacb, Germany) observed a mass occurrence under ice in a eutrophic
lake (AltmUhlsee) in Bavaria, Germany (autecological data, see FOISSNER et al. 1999).
Feeds on flagellates, planktonic diatoms and coccale algae (HORvATH 1933, WILBERT
1986a); possibly, HORVATH misinterpreted the latter as symbiotic green algae (see above).
Biomass of 106 individuals
110 mg. WILBERT (1986a)
found P. planctonticum at
0.4-11.2°C, pH 7.7-8.3,
4.9-12.5 mg I-I (h, 0 to
0.5 mg I-I NH4+-N, 0 mg II
N02--N, 3.5-14.0 mg II
N03--N. The AltmUhlsee
population was heavily in-
fected by a parasitic suctor
(FOISSNER et al. 1999).

Fig. 147m, D Pseudostrombid-

ium planclonlicum (from FOIss-
NER et al. 1999. m, n, protargol
impregnation). m, n) Ventral
and dorsal infraciliature. Arrows
in (n) mark caudal cirri rows.
I ! CC = caudal cirri, LMR = left
m n marginal cirri row; p.494.
 Stylonychinea 499

Stylonychinae BERGER & FOISSNER, 1997

1997 Stylonychinae BERGER & FOISSNER, Arch. Protistenk., 148: 149. - Type (original designation): Sty[ony-
chia EHRENBERG, 1830.

C h a r act e r i sat ion: Oxytrichidae with rigid body. Cortical granules lacking. Ado-
ral zone of membranelles usually >40 % of body length. Primarily 18 frontal-ventral-
transverse cirri clustered to six distinct groups, secondarily number of these cirri slightly
to distinctly increased or slightly decreased. Cirrus V/3 not involved in primordia fonna-
tion. Dorsal morphogenesis primarily in Oxytricha pattern, secondarily with multiple
fragmentation and/or retention of parental kineties.

Rem ark s: BERGER & ForssNER (1997) established this taxon as subfamily, as also indi-
cated by the defmed ending -inae. Here I ignore the category (for details, see Ax 1995,
P 18 and WESTIlEIDE & RiEGER 1996, P VII). However, to avoid inflation of names I use
the name without any change.
Cladistic analysis using mainly morphological and morphogenetic data divided the
18-cirri oxytrichids in two groups, namely the Stylonychinae and its sister group, Oxytri-
chinae (Fig. 25a). Autapomorphies of the Stylonychinae are the rigid body, the lack of
cortical granules, and an adoral zone of membranelles which is usually more than 40 %
of body length. The autapomorphy "origin of proter's primordia V and VI from cirrus
V/4" was proposed by the PAUP-program (Fig. 25b), but not supported by manual analy-
sis because of different defmition of the plesiomorphic state.
The consistency of the body was already used by STEIN (1859b) and later by KENr
(1882), KAHL (1932), BORROR (1972a), and SMALL & LYNN (1985) to distinguish several
oxytrichids. Suppleness is typical for the sister group of the Stylonychinae, the Oxytrichi-
nae, and for very many (all? except for euplotids and the curious Psilotricha) other hypo-
trichs. This strongly indicates that a flexible body is the plesiomorphic, and a stiff one the
apomorphic state. I am rather sure that this character evolved only once, inasmuch as it is
supported by two other autapomorphies. The difference in the body consistency is due to
ultrastructural specialisation. Oxytrichids with a rigid body have several, in Stylonychia
arranged crosswise, layers of subpellicular microtubules (CALVO et al. 1986a, PuYTORAC
et al. 1976), while hypotrichs with a supple body have only a single sheet of parallel, po-
lar microtubules (GRIMES 1972, WIRNSBERGER-AEsCHT & ForSSNER 1989). The consistency
of the body is not only of high cladistic value, but also important in identification
practice. Flexible hypotrichs bend laterally while moving unhampered on the substratum,
when swimming they are often curved and when touching an obstacle their anterior half
contracts slightly. Conversely, rigid hypotrichs never bend in any direction while creep-
ing and look like a rotating board when swimming.
Cortical granules (previously often called subpellicular granules) are widespread in
flexible oxytrichids and non-oxytrichid hypotrichs, except euplotids (see ForSSNER et al.
1991 and Table 2), strongly suggesting, that the possession of such structures is a plesio-
500 SYSTEMATIC SECTION

morphy. Conversely, cortical granules are entirely lacking in rigid oxytrichids. Obvious-
ly, the granules were lost in the last common ancestor of the rigid oxytrichids, possibly in
consequence of the altered ultrastructural architecture.
The third autapomorphy of the Stylonychinae is the relative elongation of the adoral
zone of membranelles. It is less than 40 % (usually 25-35 %) in oxytrichids having a
flexible body and in most other hypotrichs (except euplotids), suggesting that such values
are plesiomorphic. Exceptions are rare, for example, Cyrtohymena quadrinucleata and
Gonostomum spp., whose adoral zone is 40-50 % of body length, as in all oxytrichids
with a rigid body. A correlation between the relative length of the adoral zone and the
mode of nutrition is neither known nor apparent.
All other features mentioned in the characterisation of the Stylonychinae are either
plesiomorphies (for example, cirrus V/3 not involved in primordia formation) or con-
spicuous autapomorphies of "atypical" members of this group (for example, number of
frontal-ventral-transverse cirri secondarily increased in Laurentiella strenua, Onychodro-
mus spp., Pattersoniella vitiphila). It was mainly the paper by SCHLEGEL et al. (1991) that
encouraged me to include "atypical" taxa in the Stylonychinae, inasmuch as there is no
conflict with the three autapomorphies discussed above. However, it is very likely rather
difficult to find their exact placement in the cladograrn and I thus arrange them at the end
of the Stylonychinae section.
The IS-cirri oxytrichids Pleurotricha, Rigidocortex, and the little-known Parastylo-
nychia have a rigid body, no cortical granules, and a long oral apparatus. These features
assign them unequivocally to the Stylonychinae. However, they cannot be considered in
the cladistic analysis because detailed morphogenetic data are lacking; thus, the clado-
grams presented in Figure 25 are rather incomplete and they will change more or less sig-
nificantly as further taxa are analysed.
Still, some genera are not defmed by a unique character (autapomorphy), but by a
specific combination of plesiomorphies, that is, their monophyly is questionable. Thus,
and because I did not name and characterise the sister groups within the Stylonychinae,
the characterisation of a genus includes all features necessary to separate it from the other
genera of this group.
Recently, EIGNER (1997) transferred most taxa of the Stylonychinae to the Para-
kahliellidae EIGNER.

Key to genera
For identification of genera, see key at the beginning of the systematic section
(p 104-113). Most (all?) rigid, non-euplotid hypotrichs belong to the Stylonychinae.
 Stylonychia 501

Stylonychia EHRENBERG, 1830

1829 Stylonychia cimex EHRENBERG, Abh. preuss. Akad. Wiss., year 1829: 12, 17, 19 (see nomenclature and
taxonomy).
1830 Stylonychia mytilus - EHRENBERG, Abh. preuss. Akad. Wiss., year 1830: 43 - Type (subsequent desig-
nation by FROMENTEL 1875, P 163): Trichoda mytilus MOLLER, 1773.
1830 Stylonychia histrio - EHRENBERG, Abh. preuss. Akad. Wiss., year 1830: 43.
1838 Stylonychia - EHRENBERG, lnfusionsthierchen, p 370.
1859 Stylonychia. EHRBG. - STEIN, Organismus der lnfusionsthiere I, p 146.
1875 Stylonychia - FROMENTEL, Etudes microzoaires, p 162.
1882 Stylonychia, EHRENBERG - KENT, Manual Infusoria, p 790.
1932 Stylonychia EHRENBERG, 1838 - KAHr., Tierwelt Dtl., 25: 617 (classified as subgenus ofOxytricha).
1972 Stylonychia EHRENBERG, 1830 - BORROR, 1. Protozoal., 19: IS.
1974 Stylonychia EHRENBERG- STILLER, Fauna Hung., lIS: 153.
1997 Stylonychia EHRENBERG, 1830 - BERGER & FOISSNER, Arch. Protistenk., 148: 151.

C h a r act e r i sat ion: Undulating membranes in Stylonychia pattern. One right and 1
left row of marginal cirri, distinctly separate posteriorly. Six dorsal kineties. Caudal cirri
present and often distinctly elongated. Primordium II of proter originates from oral pri-
mordium. Primordia V and VI of proter originate from frontoventral cirrus IV/3 or from
postoral ventral cirrus V14. Primordium IV of opisthe originates from postoral ventral cir-
rus V/4. Dorsal morphogenesis in Oxytricha pattern.

Add i t ion a I c h a r act e r s: Body rigid. Two ellipsoidal macronuclear nodules,

except S. nodulinucleata which has a moniliform macronucleus. Contractile vacuole in
about mid-body. Cortical granules lacking. Rapid movement. Adoral zone of mem-
braneHes about 40-50 % of body length. Dorsal cilia short, that is, 3-5 !lm.

Nom e n c I at u rea n d t a x 0 nom y: Stylonychia was mentioned for the first time
in EHRENBERG (1829) in combination with the epithet cimex as follows: "Stylonychia? ci-
mex. n. sp." (p 17) and "Stylonychia cimex. n. G." (p 19). Since this is the only originally
included species in the nominal genus-group Stylonychia it would be its type species by
indication (type by monotypy; Article 68 [d] of the IcZN 1985). However, somewhat later
this species (which is obviously identical with Trichoda cimex MOLLER, 1773) was classi-
fied as Euplotes (EHRENBERG 1838, P 380); FROMENTEL (1876) mentioned this species for
the last time. In 1830, EHRENBERG transferred Trichoda mytilus MOLLER, 1773 and Para-
maecium histrio MOLLER, 1773, to Stylonychia. This paper (EHRENBERG 1830) is the gen-
erally accepted original description of Sty/onychia (for example, BORROR 1972a, CORLISS
1979). Trichoda mytilus was subsequently designated as type by FROMENTEL (1875,
P 163) and any change in this situation would disturb stability and cause great confusion.
KAHL (1932) classified Stylonychia as subgenus of Oxytricha; the correct name in his re-
vision is thus Oxytricha (Stylonychia).
Incorrect subsequent spellings: Stilonichia (EDDY 1925, P 85); Stilonychia (HENTSCHEL
1916, P 25); Stilonychia mytilus (ASPER & HEUSCHER 1886, p 170); Stylochia (ScHLICHTING
502 SYSTEMATIC SECTION

1961, P 87); Stylometra pustulata (JENNINGS & MOORE 1902a, p 191); Stylomychia (CAIRNS
et al. 1976, p 265); Stylonchia (GIESE & ALDEN 1938, P 125; LEE & KUGRENS 1992, P 538;
ZAK1AN 1989, p 584); Stylonchia curvata (SEN et al. 1967); Stylonchia /emnae (WEN et al.
1995, p 1704); Sty/onchia mytilus E. (CIENKOWSKY 1855, P 302; TORRES et al. 1979a); Sty-
/onichia EHRENBERG (this is the most frequent incorrect subsequent spelling: for example
ANON 1965; AWERINZEW 1900, P 324; BAILY 1845, P 34; BRUGEROUE & AooUTIE 1988,
P 257; BUOINGTON & HARVEY 1915, P 305; CHENEVEAU & BOHN 1903, P 800; FABRE-
DoMERGUE 1887, P 798; FAoo-FREMIET 1910, P 46; LQpEZ-OcHOTERENA 1967, P 165; MAu-
PAS 1883, P 619; PERKINS 1956, P 59; SCHULZ 1961, P 51; SCOURFIELD 1897, P 790; SERNOV
1929, P 557; SIMON 1988, P 16; SLAoECEK et al. 1958, p 74; SUOZUKI 1978, P 90; TURNER
1954, P 19; VAN HOlDE et al. 1979, p391; WANG 1925, p51; WATSON 1946a, b, p46,
127); Stylonichnia mytilus (BAillENSPERGER 1927, p 295); Stylonichus mytilus (SHARAVINA
1978, p 1573); Stylonycha (KURASAwA 1958, p 119); Stylonychi /emnae (WEFES & LIPPS
1990, P 25); Stylonychna mytilus (BAillENSPERGER 1927, P 211); Stylonychnia mytilus
(BAillENSPERGER 1927, P 197, 229, 272, 283); Sty/onychnia pustulata (BAillENSPERGER
1927, P 283); Stylonyehia sp. (DUTTA et al. 1990, p 22); Styloychia mytilus EHR. (GUHL
1987, p 455); Stynolychia monostylus (FROMENTEL 1876, P 359).
The characterisation of the genus is according to BERGER & FOISSNER (1997). For de-
scription of ''patterns'', see chapter 2 in the general section. Morphogenetic data suggest
that Sty/onychia is not monophyletic (Table 4; Fig. 25), that is, species of the S. mytilus
complex are very likely more closely related to Coniculostomum and Steinia than to other
well-known stylonychias, like S. pustulata. However, molecular biological data on
Steinia and Conicu/ostomum and morphogenetic data on further stylonychias should be
awaited before changing names. Recently, EIGNER (1997) transferred Sty/onychia pustu-
lata and S. vorax to the Parakahliellidae. Consequently he had to establish a new genus,
Clara (type species: Kerona pustu/ata MOLLER, 1786), which is, however, a junior homo-
nym that has to be replaced.
In the present book, Stylonychia includes 11 species, largely confined to freshwater.
Only four of them are described in detail, namely S. mytilus, S. lemnae, S. pustu/ata, and
S. hi/aria. The remaining species need detailed redescription, that is, live observations,
protargol impregnation, morphometric characterisation. Furthermore, study of morpho-
genesis is recommended to show whether they belong to the S. myti/us or to the S. pustu-
lata clade. Species of the S. mytilus complex and S. pustu/ata are one of the most com-
mon oxytrichids in lentic and lotic waters (FOISSNER et al. 1991).
When I wrote the main part of the Stylonychia section I did not have all papers now
included in the lists of synonyms. Thus, not all illustrations concerning a species are ar-
ranged in succession.

Key to species

Rem ark s: Some species, especially Sty/onychia putrina, S. vorax, S. curvata, and the
very common and well described S. pustulata are rather difficult to distinguish. Because
 Stylonychia 503

conspicuous and unambiguous differences are lacking, I had to use mainly the body
shape, which is of course unsatisfactory. Thus, use only healthy specimens, look at the
body shape without squeezing, and read the descriptions carefully. If identification is un-
certain, for example in water pollution assessment when only one or few specimens have
been observed, I recommend writing Stylonychia sp. or S. pustulata group. If caudal cirri
inconspicuous, see also Sterkiella and Rigidocortex, which have, however, undulating
membranes in Oxytricha pattern.

1 Macronucleus moniliform ......................... S. nodulinucleata (p 552)

2 macronuclear nodules ............................................... 2
2 Invariably 1 micronucleus between macronuclear nodules (Fig. 167b-d) ......... .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. S. stylomuscorum (p. 595)
2 or more micronuclei ................................................ 3
3 Caudal cirri inconspicuous, that is, not distinctly longer than marginal cirri (Fig.
157a, i-k) ................................................. S. hi/aria (p 557)
Caudal cirri distinctly longer than marginal cirri (for example, Fig. 148c, 15ge) .. 4
4 Transverse cirri distinctly separated into 2 groups (for example, Fig. 148d, 149a) . 5
Transverse cirri arranged like a hook, that is, do not form 2 distinct groups (for ex-
ample, Fig. 160c, 161a, c) ............................................. 7
5 Body distinctly broader anteriorly than posteriorly, tapering backwards from centre,
posterior end usually truncated, left margin usually concave, right convex; postoral
ventral cirrus V/4 in front of level of postoral ventral cirrus IVI2; caudal
cirri widely separated (for example, Fig. 148a-c, 0, 150k, 151k) ................ .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. 6 (S. mytilus complex, p 504)
Body roughly parallel-sided, posterior end rounded; postoral ventral cirrus V/4 ar-
ranged behind level of postoral ventral cirrus IV/2; caudal cirri closely spaced (Fig.
15ge, 166c) ............................................. S. notophora (p 553)
6 About 44--66 and 40-56 basal body pairs in dorsal kinety 3 and 4 respectively (Table
31) .................................................. S. mytilus (p 549)
About 33-40 and 30-39 basal body pairs in dorsal kinety 3 and 4 respectively (Table
31) .................................................. S. lemnae (p 550)
7 (4) Usually 10-12 frontal, frontoventral, and buccal cirri left of and beside adoral
zone of membranelies; caudal cirri displaced slightly forward on right margin (Fig.
167a; little-known, uncertain species) .................. S. pseudograndis (p 564)
Rather invariably 8 frontal, frontoventral, and buccal cirri; caudal cirri almost ex-
actly terminal (for example, Fig. 161a) ................................... 8
8 Outline ovoid, that is, posterior end broadly rounded (Fig. 167t) . S. curvata (p 593)
Outline elliptical, parallel-sided or slightly converging posteriorly but never ovoid,
posterior end rounded or somewhat tapered .............................. 9
9 Posterior half of cell tapering and obtusely pointed (Fig. 166a, b) . S. vorax (p 591)
Outline parallel-sided or elliptical, posterior end of cell usually rounded (for exam-
ple, Fig. 162a, 166a) ................................................ 10
504 SYSTEMATIC SECTION

10 Outline usually elliptical, posterior end often narrowly rounded; caudal cirri indis-
tinctly separate; left frontoventral cirrus III/2 immediately beside other frontoventral
cirri (Fig. 162a, c, x) ...................................... S. pustu/ata (p 565)
Outline usually pronouncedly parallel-sided, both ends broadly rounded; caudal cirri
distinctly separate; left frontoventral cirrus III12 arranged immediately beside undu-
lating membranes and distinctly separate from the 3 other frontoventral cirri, which
are arranged in a short oblique row (Fig. 165a, c, d) ........... S. putrina (p 586)

Stylonychia mytilus complex (Fig. 7a, b, 13a-p, 17a-k, 18b, 27a-h,

148a-p, 149a-d, 150a-z, 151a-z, 152a-o, q-t, 153a-l, 154f-p, 155,
159g, 163e, Tables 3, 4, 18,28,31-35)

1773 Trichoda mytilus MOLLER, Vermium Terrestrium et Fluviatilium, p 87.

1773 Trichoda silurus MOLLER, Vermium Terrestrium et Fluviatilium, p 88.
1786 Kerona myti/us - MOLLER, Animalcula Infusoria, p 242 (Fig. 150d-f).
1786 Kerona silurus - MoLLER, Animalcula Infusoria, p 244, Tab. XXXIV, Fig. 9, 10.
1830 Sty/onychia myti/us - EHRENBERG, Abh. preuss. Akad. Wiss., year 1830: 43.
1833 Sty/onychia mytilus - EHRENBERG, Abh. preuss. Akad. Wiss., year 1833: 174,330 (Fig. 150g; "Waf-
fenthierchen").
1838 Sty/onychia mytilus - EHRENBERG, Infusionsthierchen, p 370 (Fig. 150h, i).
1838 Sty/onychia silurus - EHRENBERG, Infusionsthierchen, p 372, Tafel XLII, Fig. II.
1841 Kerona myti/us - DuJARDIN, Zoophytes, p 425, Planche XII, fig. 2, 3.
1852 Cerona myti/us M. - PER1Y, Zur Kenntoiss kleinster Lebensformen, piSS.
1858 Sty/onychia mytilus EHR. - CLAPAREDE & LACHMANN, Mem. Inst. nato. genev., 5: 158 (Fig. 150j).
1858 Sty/onychia mytilus - EBERHARD, Programm der Realschule zu Koburg, Jahr 1858: 50 (Fig. 152t; very
low quality).
1859 Sty/onychia mytilus. EHRaG. - STEIN, Organismus der Infusionsthiere I, p 147 (Fig. 150k, I).
1860 Sty/onychia myti/us - BALBIANI, Journal de la physiologie de I'homme et des animaux, 3: 86, Planche
IV, Fig. 1-8.
1862 Sty/onychia mytilus- ENGELMANN, Z. wiss. Zoo!., 11: 359 (Fig. ISla).
1865 Sty/onychia mytilus - QUENNERSTEDT, Acta Univ. lund., 2: 56 (Fig. 151x-z).
1867 Sty/onychia mytilus. EHR. - WRZESNloWSKIEGO, Rozprawa, Krak6w, p 113 (Fig. 151u).
1867 Sty/onychia myti/us. EHR. - WRZESNIOWSKIEGO, Rocznik ces. kr61. Towarzystwa Naukowego Kra-
kowskiego. (Krakow). Poczet trzeci, 12: 341 (Fig. 151u).
1876 Kerona dubius FROMETEL, Etudes microzoaires, p 270, Planche XIII, fig. 3.
1876 Sty/onychia mytilus- FROMENTEL, Etudes microzoaires, p 275, Planche XIV, fig. 10, lOa
1877 Stylonychia myti/us O. F. MOLLER. EHR. - MERESCHKOWSKY, Trudy imp. S-peterb. Obshch. Estest., 8:
234 (Fig. 152c).
1882 Sty/onychia myti/us, EHR. - KENT, Manual infusoria II, p 790.
1899 Sty/onichia mytilus EHRENBERG - ScHUSTER, Sci. Gossip,S: 202 (Fig. 154n; incorrect subsequent spell-
ing).
1900 Sty/onychia mythilus - WALLENGREN, Bih. K. svenska VetenskAkad. Hand!., 26: 14 (Fig. 152f; incor-
rect subsequent spelling).
1900 Sty/onychia myti/us MOLL. - PoTTER, Arch. Anat. Physiol., 1900: 267 (Fig. 152j).
1901 Sty/onychia myti/us (O.-F. MOLL.) - Roux, Mem. Inst. nato. genev., 19: 103 (Fig. 151v).
1902 Sty/onychia mytilus- WALLENGREN, Zool. lb., 15: 20 (Fig. 163f).
1905 Sty/onychia sp. (?) - CONN, Bull. Conn. St. geo!. nat. Hist. Surv., 2: 61 (Fig. 152n).
 Stylonychia 505

1905 Stylonychia mytilus MOLL. - CONN, Bull. Conn. St. geol. nat. Hist. Surv., 2: 61 (Fig. 1540).
1905 Stylonychiafissieta C. & L. - CONN, Bull. Conn. St. geol. nat. Hist. Surv., 2: 61 (Fig. 1520; incorrect
subsequent spelling).
1906 Stylonychia mytilus EHR. - EDMONDSON, Proc. Davenport Acad. Sci., 11: 101 (Fig. 152q).
1906 Stylonychia sp. - EDMONDSON, Proc. Davenport Acad. Sci., II: 102 (Fig. 152r).
1908 Stylonychia mytilus O. F. MOLL. - BORGER, An. Univ. Chile, 122: 188 (Fig. 154m).
1909 Stylonychia mytilus- FAURE-FREMIET, Arch. Protistenk., 13: 127 (Fig. 151w).
1912 Stylonychia mytilus- LEWIN, Proc. R. Soc., 84: 336 (Fig. 152a).
1914 Stylonychia mytilus EHR. - SMmI, Kans. Univ. Sci. Bull., 9: 166 (Fig. I 54f).
1920 Stylonichia mytilis EHRBG. - FELLERS & ALUSON, Soil Sci., 9: 9, 24 (Fig. 152g; incorrect subsequent
spelling; identification uncertain).
1922 Stylonychia mytilus - PFEIFFER, Mitteilungen der Markischen mikrobiologischen Vereinigung, II: 131,
Abb. A, B.
1925 Stylonichia mytilus EHRENBERG - WANG, Contr. bioI. Lab. Sci. Soc. China, I: 51 (Fig. 152d; incorrect
subsequent spelling).
1925 Stylonychia mytilus - DEMBOWSKA, Arch. mikrosk. Anat. EntwMech., 104: 185 (Fig. 163e).
1928 Pleurotricha lanceolata - RAo, 1. Indian Inst. Sci., I1A: Legend of plate II (Fig. 1510; misidentifica-
tion).
1931 Stylonychia mytilus EHRBG. - TAl, Sci. Rep. natn. Tsing Hua Univ., 1: 52 (Fig. 154i).
1932 Stylonychia mytilus EHRENBERG, 1838 - KAIu., Tierwelt Dtl., 25: 618 (Fig. 151k, I).
1932 Stylonychia spec. - KAIu., Tierwelt Dtl., 25: 619 (Fig. 151m).
1938 Stylonychia mytilus- DEMBOWSKA, Arch. Protistenk., 91: 89 (Fig. 15Ip).
1944 Sty/onychia mytilus- CHEN, J. Morph., 75: 338 (Fig. I SOn).
1944 Stylonychia mytilus MULLER - BUIsAN, Publnes Inst. BioI. apI., I: 20 (Fig. 1541).
1954 Stylonychia clavi/ormis GELEI, Acta bioI. hung., 5: 336 (Fig. 151 e, f, t; was establish in Oxytricha, thus
the correct name is O. (Stylonychia) clavi/ormis).
1955 Stylonychia mytilus EHRENBERG - BARWICK, BEVERIDGE, BRAZIER, CLOSE, HIRSCHFELD, PILLA!, RAMSAY,
ROBINSON, STEVENS & TODD, Tuatara, 5: 96 (Fig. 154p).
1960 Stylonychia mytilus STEIN - STILLER, Fauna Hung., 57: II, 13. libra, B (redrawing after BOTSCHu).
1961 Stylonychia mytilus EHRENBERG, 1838 - REUTER, Acta zool. fenn., 99: 21 (Fig. ISis).
1961 Stylonychia mytilus EHR. - BucK, Jh. Ver. vaterl. Naturk. WUrtt., 116: 212 (Fig. 15In).
1963 Stylonychia mytilus EHRB. - BIERNACKA, Polskie Archwm Hydrobiol., 11: 50 (Fig. 154k).
1965 Sty/onychia myti/us (MOLLER) - TCHANG & PANG, Kexue Tongbao, 4: 358 (Fig. 150m).
1965 Sty/onychia mytilus Varietllt I - AMMERMANN, Arch. Protistenk., 108: III (Fig. 149d).
1965 Stylonychia mytilus Varietat 2 - AMMERMANN, Arch. Protistenk., 108: III (Fig. 150c).
1965 Stylonychia mytilus EHRENBERG - MACIlEMER, Arch. Protistenk., 108: 153 (Fig. 152h).
1965 Stylonychia mytilus- TVFFRAU, Archs. Zool. expo gen., 105: 84 (Fig. 152i).
1965 Stylonichia mytilus EHRENBERG, 1838 - L6PEZ-OcHOTERENA, Revta Soc. mex. Hist. nat., 24: 160 (Fig.
152k, I; incorrect subsequent spelling).
1966 Stylonychia mytilus EHR. - DRAGESCO, Protistologica, 2: 90 (Fig. 150s-z).
1967 Stylonichia mytillus MULLER - CHARDEZ, Revue Ecol. BioI. Sol, 4: 294 (Fig. 154g; incorrect subsequent
spelling).
1968 Oxytricha (Stylonychia) mytilus EHRENBERG, 1838 - CHORIK, Free-living ciliates, p 139 (Fig. 15Ig).
1969 Stylonychia mytilus - TUFFRAU, Protistologica, 5: 230,235 (Fig. 15li).
1969 Stylonychia mytilus EHRENBERG, 1838 - CURDS, Wat. Pollut. Res., No. 12: 70 (Fig. 151q).
1971 Stylonychia putrina STOKES (7) - DRAGESCO & NJINE, Annis Fac. Sci. Univ. fed. Cameroun, No. 7-8:
130; in part: "grande variete", Fig. 27; not "petit variete", Fig. 28 (Fig. 159g; misidentification).
1971 Stylonychia mythilus EHRENBERG - CHARDEZ, Revue verviet. Hist. nat., 28: 38 (Fig. 154j; incorrect sub-
sequent spelling).
1972 Stylonychia mytilus (MOLLER, 1773) EHRENBERG, 1830 - BORROR, J. Protozool., 19: 15.
1972 Sty/onychia mytilus EHRENBERG - BICK, Ciliated protozoa, p 174 (Fig. 151 b).
506 SYSTEMATIC SECTION

1973 Sty/onychia mytilus- AMMERMANN, Cell development, p 51 (Fig. 148q, r).

1974 Sty/onychia mytilus (MOLLER, 1773) EHRENBERG, 1830 - JONES, Univ. South Alabama Monogr., I: 41
(Fig. IS2e).
1974 Sty/onychia steinii STILLER, Annis hist.-nat. Mus. natn. hung., 66: 133 (see nomenclature and
taxonomy).
1975 Sty/onychia mytilus - WILBERT, Mikrokosmos, 64: 176, Bild 9.
1977 Stylonychia mytilus EHRB. 1838 - BERECZKY, AnnIs Univ. Scient. bpest. Rolando EOtvOs, 18-19: 170
(Fig. IS2m).
1979 Stylonichia mytilus EHRB., 1838 - MAMAEvA, Infusoria of the Volga Basin, p 72 (Fig. IS4h; incorrect
subsequent spelling).
1981 Stylonychia mytilus- TVFFRAU, FRYD-VERSAVEL & TUFFRAU, Protistologica, 17: 387 (Fig. 13a-p, ISlr).
1982 Stylonychia mytilus EHRENBERG, 1838 - FOISSNER, Arch. Protistenk., 126: 107 (Fig. IS0o-r).
1982 Spirotrich-JEFFRIES, J. bioI. education, 16: 37, Fig. 6 (microphotograph).
1983 Stylonychia mytilus, EHRENBERG 1838 - AMMERMANN & SCHLEGEL, J. Protozool., 30: 292 (Fig. 148n, 0,
IS1j).
1983 Stylonychia lemnae AMMERMANN & SCHLEGEL, J. Protozool., 30: 291 (Fig. 148m, p, 149c).
1983 Stylonychia mytilus EHRENBERG, 1838 - STEINBROCK & SCHLEGEL, J. Protozool., 30: 294.
1983 Stylonychia lemnae STEINBROCK & SCHLEGEL, J. Protozool., 30: 296 (see nomenclature and taxonomy).
1986 Sty/onychia mytilus - WIRNSBERGER, FOISSNER & ADAM, Arch. Protistenk., 132: 169 (Fig. 149a, b,
IS3d).
1986 Sty/onychia lemnae AMMERMANN & SCHLEGEL, 1983 - WIRNSBERGER, FOISSNER & ADAM, Arch. Protis-
tenk., 132: 167 (Fig. IS0a, b, I 53a-c, e-j).
1986 Sty/onychia myti/us (0. F. MOLLER, 1773) EHRENBERG, 1830 - DRAGESCO & DRAGESCO-KERNEIS, Faune
tropicale, 26: 486.
1986 Sty/onychia putrina STOKES, 188S - DRAGESCO & DRAGESCO-KERNEIS, Faune tropicale, 26: 487, in part:
Planche 14S A; not Planche 14S B-F (misidentification).
1987 Sty/onychia mytilus EHRENBERG, 1838 - TVFFRAU, Annis Sci. nat., Zoologie, 8: 113 (Fig. ISlh).
1987 Sty/onychia myti/us - MACHEMER & DEITMER, Progr. Protistol., 2: 217 (Fig. ISle, d; also figured in
SLEIGH 1989).
1987 Sty/onychia mytilus EHRENBERG, 1838 - LoKOT, Okologie der Wimpertiere, p 74 (Fig. IS2s).
1988 Sty/onychia mytilus (0. F. MOLLER, 1773) - SHIN & KIM, Korean J. syst. Zool., Special Issue No.2: 71,
3A-F.
1991 Sty/onychia mytilus-Komplex - FOISSNER, BLATTERER, BERGER & KOHMANN, Infonnationsberichte Bayer.
Landesamtes filr Wasserwirtschaft, 1/91: 31S (Fig. 148a-1).
1993 Sty/onychia mytilus EHRENBERG - YOSHINO, Island Studies in Okinanwa, 11:63, Plate VIII, Fig. 7 (mi-
crophotograph).
1997 Stylonychia mytilus-Komplex - BERGER, FOISSNER & KOHMANN, Bestimmung und Okologie der Mik-
rosaprobien, p 163.

Nom e n c I at u rea n d t a x 0 nom y: AMMERMANN (1965a) reported that the wide-

spread and very well-known species Stylo11)'chia mytilus consists of 2 morphologically
inseparable but distinguishable genetic species. He followed SONNEBORN (1957) and used
the term "syngen" and "variety", respectively, to designate these sibling species. About
20 years later, a fully distinctive taxonomic name for the S. mytilus syngen 1, namely Sty-
/o11)'chia lemnae was proposed. Unfortunately, this name was published several times by
various authors with the appendix "n. sp." CAMMERMANN' & SCHLEGEL 1983; STEINBROCK &

Fig. 148a-g Sty/onychia myti/us complex (from FOISSNER et al. 1991. a-c, f, g, from life, bright field; d, e, ~
protargol impregnation). a, f) Ventral views. b, c) Dorsal views. d) Ventral infraciliature of a specimen with
 Stylonychia 507

AZM
FC
RMR-
,
# Ma
.,
,
,
I'

,
VC
, f

-cc
d e 9

only 2 frontal cirri. The transverse cirri are separated into two distinct groups (arrows). e) Dorsal view showing
adoral zone of membranelles, caudal cirri, and nuclear apparatus; arrows mark micronuclei. g) Lateral view of
specimen creeping on detritus. AZM = adoral zone of membranelles, CC = posteriorly fringed caudal cirri, FC
= frontal cirrus, LMR = left marginal cirral row, Ma = macronuclear nodule, RMR = right marginal cirral row,
TC = transverse cirri, VC = right pretransverse ventral cirrus (cirrus VII2); p 504.
 508 SYSTEMATIC SECTION

Fig. 148b-1 Stylonychia myilus complex (from FOlssNER et aI. 1991. h, i, k, I, scanning electron micrographs;
j, from life, bright field). b, I) Ventral views. Note that transverse cirri are distinctly separated into two groups.
i) Anterior half in ventral view. Note large, bowl-shaped buccal area. Large arrow marks buccal cirrus. This
specimen has a supernumerary frontoventral cirrus (small arrow). j) Resting cyst. k) Right lateral view. Large
arrow marks "collar", small arrow denotes dorsal kinety 5. AZM = adoral zone of membrane lies, CC = caudal
cirri, FC = right frontal cirrus, p = paroml, pVC = posterior postoml ventral cirrus (Vl3), TC = right transverse
cirrus; p 504.
 Stylonychia 509

Fig. 148m, p Sty/onychia /emnae (from AMMERMANN & ScHLEGEL 1983. m, scanning electron micrograph; p,
phase contrast micrograph). m} Ventral view, 240 11m. p} Dorsal view of a cell starved for one day, 240 11m;
p504.

Fig. 148n, 0 Stylonychia mytilus (from AMMERMANN & SCHLEGEL 1983. n, scanning electron micrograph; 0,
phase contrast micrograph). n} Ventral view, 290 11m. o} Dorsal view of a cell starved for one day, 310 11m;
p 504.

Fig. 148q, r Stylonychia mytilus complex (from AMMERMANN 1973). q) Macronucleus anlage after conjuga-
tion, about 20 h old, with short spiralled and long de-spiralled chromosomes. r) Portion of a giant chromosome
of a macronucleus anlage, about 40 h old, with a large heterochromatin block; p 504.
 510 SYSTEMATIC SECTION

.,
, ,
,,
.
•
,

•
• §§
~

""......
I
~

~~~~ :-.. :
V/4 'IV/2 ..
•
..•.-. , .

~
--
....-a • • b
Fig. 149a-d Sty/onychia lemnae (a, b, from WIRNSBERGER et aI. 1986; c, from AMMERMANN & SCHLEGEL 1983;
d, from AMMERMANN 1965a. a, b, protargol impregnation; c, redrawn from a scanning electron micrograph; d,
from life). a-c) Ventral and dorsal infraciliature, a, b = 180 !lID, c = 215 Jlffi. Arrowhead in (a) marks fronto-
ventral cirrus lI112, which is arranged slightly behind level of cirrus IV/3 (arrow); possibly an important differ-
ence to S. mytilus (cp. Fig. 150a and see text). Arrangement of cirri in (c) obviously incorrect (cp. Fig. 149a).
The anterior end of dorsal kineties 1-4 is obviously less curved than in S. mytilus (cp Fig. 150b). d) Ventral
view, size not indicated; p 504.

SCHLEGEL 1983; FEYER & MACHEMER 1983, P 224; DEITMER et aI. 1983, P 286). Since the
description of the morphology, the reproductive behaviour, and the type location are pro-
vided by AMMERMANN & SCHLEGEL (1983) they have to be considered as the describers.
Of course it is impossible to determine which species of the S. mytilus complex MOI.-
LER saw in 1773. But, as in the case of the Tetrahymena pyriformis complex (CORLISS &
DAGGETI 1983, NANNEY & McCoy 1976), one variety, namely S. mytilus syngen2 of AM-
MERMANN (l965a) has to retain the original taxonomic name. This is in agreement with
Artkle 17 (I) of the IcZN (1985), which says, that the availability of a name is not af-
fected even if it is found that the original description relates to more than one taxon. The
authorship of S. mytilus is often erroneously credited to EHRENBERG (see list of syno-
 Stylonychia 511

I
,, .
~'
: '

•• \.\
, "'.

•,, ....
,
~.:~:.
. "
.
•• • .
~.

····1. .-: ~ ",

••,
""
I

"
I
••• .,
't •
I
I,
", b

Fig. 150a--c Sty/onychia mytilus (a, b, from WIRNSBERGER et al. 1986; c, from AMMERMANN 1965a a, b, protar-
gol impregnation; c, from life). a, b) Ventral and dorsal infraciliature, a, b = 215 11m. Arrowhead marks fron-
toventral cirrus 11II2, which is situated slightly in front of level of cirrus IV13 (arrow); possibly an important
difference to S. /emnae (cp. Fig. 149a and see text). Furthennore, the dorsal kineties 1-4 are obviously some-
what more curved anteriorly than in the sister group, S. /emnae (cp. 149b). c) Ventral view, size not indicated;
p504.

nyms). However, there is no doubt that it has to be attributed to MOLLER (1773), as cor-
rectly stated by BORROR (l972a). The classification in Stylonychia was already done by
EHRENBERG (1830) and not by EHRENBERG (1838), as often assumed.
KAlIL (1932) classified Stylonychia as subgenus of Oxytricha; the correct name in his
revision is thus Oxytricha (Stylonychia) mytilus. Incorrect subsequent spellings: miti/us
(ENRIQUES 1912, P 425); St. sy/urus (WEISSE 1848c, p 362); Styloichia miti/us (MAMAEVA
1979b, P 118); Stylonichia miti/us (MAoRAZO-GARIBAY & L6PEZ-OCHOTERENA 1973, P 67;
SKORIKOW 1902, P 563; WETZEL 1925, P 243); Sty/onichia muti/us (Km..EBOVICH 1987,
P 219); Stylonichia mytillus EHRENBERG (ClIARDEZ 1984, P 21; TUCULESCU 1965, P 190);
Sty/onichia mytylus EHR. (SACCHI 1888, p 51); Sty/onychia clavata GELEI (STILLER 1974b,
P 156 instead of S. c1aviformis); Sty/onychia mutilis (MULLER.) (CAIRNS 1965b, P 48); Sty-
lonychia mutilus (VEYLANDE & LIYEPA 1985, p 82); Stylonychia mysilus (DETCHEVA
1979a, p 72); Stylonychia mystilus (SCHARF 1961, p 389); Sty/onychia mytilis (BAMFORTH
512 SYSTEMATIC SECTION
 Stylonychia 513

,
,
,
,
,
..
,
,,
I

,
,
,
,
,
, •..••
.
,
#
,,
..

Fig. 150~r Sty/onychia myti/us complex (from FOlssNER 1982. 0, p, from life; q, r, protargol impregnation).
0, p) Specimens creeping on a soil particle, 0 = 85 /lIn, P = 100/lm. q, r) Ventral and dorsal infi-aciliature, q =
90 /lm, r = 95 /lm. The arrangement of the two posterior-most frontoventral cirri and the curving of dorsal ki-
neties is as in S. lemnae (cp. Fig. 149a, b); P 504.

1980b, P 148; BURKOVSKY 1984, P 10; FANG & CECH 1991, P 5515; KUTIKOWA 1984,
P 177; THOMPSON 1972, P 287); Stylonychia mytillus (DETCHEVA 1981, P 23; GRACIA et al.
1989, p 25); Stylonychia my/Huns (NOMOTO et al. 1988, p 255); Stylonychia mytylus (MA-
DONI 1988a, P 62).
The list of synonyms of S. mytilus (below) and S. lemnae (below) comprise - besides
the works of MOLLER (1773, 1786) and EHRENBERG (1830) - only those synonyms where

~ Fig. 150ll-f Sty/onychia mytilus (after MOLLER 1786). Various views from life, size not indicated; p 504, 549.
g-o Stylonychia mytilus complex (g, after EHRENBERG 1833; h, i, after EHRENBERG 1838; j, after CLAPAREDE &
LACHMANN 1958; k, I, after STEIN 1859b; m, from TCHANG & PANG 1965a; n, from CHEN 1944. g-I, from life; m,
method not indicated; n, haematoxylin stain). g-k, m, n) Ventral views, g-k, m = size not indicated, n =
905 /lm! (obviously a wrong value). Figure g shows a specimen with an ingested filiform cyanobacterium. Fig-
ure h, i seen from dorsal. I) Resting cyst, 55-70 /lm; p 504.
 514 SYSTEMATIC SECTION

Fig. 15Os-u Stylonychia mylilus com-

plex (from DRAGESCO 1966b. s-u, protar-
gol impregnation). s) Ventral inftacilia-
ture of a "nonnal specimen", size not in-
dicated and caudal cirri not shown in full
length. t) Dorsal inftaciliature schematic.
Dorsomarginal kineties obviously drawn
too long. u) Ventral inftaciliature of a
"small variety" from 111onon with an ad-
ditional frontoventral cirrus (arrow),
about 180 llm; p 504.

the identification is reliable. The other synonyms are shown in the comprehensive list of
the S. mytilus complex (see above) because an unequivocally relation to one of these two
species is impossible.
According to EHRENBERG (1838) and DIESING (1850, p 152) the following old taxa are
also synonyms of S. mytilus: Himantopus corona MUllER, 1786 (= Kerona corona (MOL-
LER, 1786) BoRY, 18242); Kerona haustellum MUllER, 1786; Kerona haustrum MUllER,
1786; Plagiotricha diana BORY, 1824; Trichoda acarus MUllER, 1773 (= Himantopus
acarus (MOILER, 1773) MOLLER, 1786 = Kerona acarus (MOILER, 1773) BORY, 1824);
Trichoda cypris MOILER, 1773 (= Kerona cypris (MUllER, 1773) MOILER, 1786); Tri-
choda erosa MUllER, 1786 (= Kerona erosa (MOLLER, 1786) BORY, 1824); Trichoda
fimbriata MUllER, 1786 (= Keronafimbriata (MOILER, 1786) BORY, 1824; = Oxytricha
fimbriata (MOILER, 1786) FROMENTEL, 1876); Trichoda ludio MOLLER, 1773 (= Himanto-
pus ludio (MOLLER, 1773) MOILER, 1786 = Kerona ludio (MOLLER, 1773) BORY, 1824);

2 BORY, 1824 = BORY DE SAINT-VINCENT in LAMOUROUX et a1., 1924.

 Stylonychia 515

Fig. lS0v-z Sty/onychia mytilus complex (from DRAGESCO 1966b. v-x, Z, protargol impregnation; y, Feu/gen
stain). v, x) Ventral and dorsal infraciliature, size not indicated. w) Nuclear apparatus. y) Resting cyst. z) Dor-
sal cilium including fibrillar system; p 504.

Trichoda rostrata MOLLER, 1786 (= Kerona rostrata (MOLLER, 1786) BORY, 1824);
Trichoda sannio MOLLER, 1773 (= Himantopus sannio (MOLLER, 1773) MOLLER, 1786 =
Kerona sannio (MOLLER, 1773) BoRY, 1824).
516 SYSTEMATIC SECTION

ell
.....
lO
.....
 Sty/onychia 517

Discussion of some synonyms of the S. mytilus complex (for a more comprehensive

discussion of pre-1838 synonyms, see EHRENBERG 1838): Sty/onychia silurus sensu EH-
RENBERG (1838) is rather small (87-117 11m), but the morphology, especially the con-
spicuous, widely spaced caudal cirri indicate that it is a member of the S. mytilus
complex. The fIrst detailed and comprehensive redescriptions of S. mytilus were provided
by CLAPAREDE & LACHMANN (1858, Fig. 150j) and especially by STEIN (1 859b, Fig. 150k,
1). STEIN studied the morphology, the morphogenesis, the conjugation, the cyst, and a suc-
torian infection. Kerona dubius FROMENTEL, 1876, is very likely a slightly deformed
specimen of a species of the S. mytilus complex. This was already supposed by FROMEN-
TEL (1876) himself; furthermore, in the legend to the fIgure he wrote "Kerona mytilus".
Stylonychia mytilus sensu MERESCHKOWSKY (1877, Fig. 152c) from a marine habitat has
only 2 caudal cirri, an anomaly also occurring in freshwater populations. Pleurotricha
lanceo/ata sensu RAo (1928, Fig. 1510) is rather surely a misidentifIcation. Obviously, he
misinterpreted a dorsal kinety as outer right marginal row; furthermore, Pleurotricha has
no caudal cirri. Stylonychia spec. sensu KAlIL (1932) is a redrawing of Tafel VIII, Fig. 3
of STEIN (l859b). The original fIgure by STEIN (l859b) shows a heavy suctorian infection
causing a conspicuous indentation of the left body margin. Furthermore, three deformed
macronuclear nodules are present. KAlIL (1932) obviously neglected these details, sepa-
rated it from S. mytilus, and stated that there is no closer relationship between S. mytilus
and Sty/onychia spec. STILLER (1974a) proposed the name S. steinii for this - in my opin-
ion - invalid taxon.
Sty/onychia clavi/ormis GELEI, 1954 (Fig. ISle, f, t) has already been synonymised
with S. mytilus by BORROR (1972a). The body shape and the arrangement of the trans-
verse and caudal cirri of the "grande variete" of S. putrina sensu DRAGESCO & NJINE
(1971, Fig. 159g; more or less identical with Planche 144 A in DRAGESCO & DRAGESCo-
.KERNElS 1986) are more reminiscent of a species of the S. mytilus complex than of S. pu-
trina.
Stylonychia mytilus sensu MAUPAS (1888, Fig. l63a) is classifIed as S. pustulata be-
cause the overall morphology does not agree in the least with that of species of the S.
mytilus complex. Obviously erroneously, MACHEMER (1966b, P 642) stated that AMMER-
MANN (1965a) recognised 3 syngens ("Varietliten") of S. mytilus.
The following redescriptions are insufficient: AGAMALIEV (1978; Fig. 230c, d), BAL-
DENSPERGER (1927, Planche VI, 204), BATIISH (1992; his Fig. 55), CIENKOWSKY (1855;
Fig. 222j), DUMAS (1929; Fig. 237f, i), L6PEZ-OCHOTERENA (1964), LUNDIN & WEST
(1963; identical with WEST & LUNDIN 1963; Fig. 234a), RAMIREz DE GUERRERO (1970,

+- Fig. 151a-4 Sty/onychia mytilus complex (a, from ENGELMANN 1862; b, from BICK 1972a; C, d, from MA-
CHEMER & DEITMER 1987. a, from life; b-d, from life?). a, b) Ventral views, a = 265 11m, b = 265 11m. c) Lateral
view (305 11m) showing adoral zone of membranelles (large arrow), that is, the most conspicuous part of the
prominent oral apparatus, the cirri (arrowheads) on the ± flat ventral side, and the ± distinctly arched dorsal
side with the dorsal kineties (small arrows). d) Ventral view mainly showing arrangement of cirri and oral ap-
paratus, 305 11m. CC = caudal cirri at posterior end of dorsal kineties I, 2, and 4, CV = contractile vacuole, p =
paroral; p 504.
518 SYSTEMATIC SECTION

..
~ ~
"
~.

'f~
? .;,.
t> Ei!
 Stylonychia 519

Fig. 151k-t Stylonychia mytilus complex (k, I, from KAliL 1932; m, after STEIN 1859b from KAliL 1932; n, af-
ter BUCK 1961; 0, afterRAo 1928; p, from DEMBOWSKA 1938; q, from CURDS 1969; r, from TUFFRAU et al. 1981;
s, after RElTfER 1961; t, from GELEI 1954b. k-q, s, from life; r, protargol impregnation; t, sublimate fixation).
k-o, q, s) Ventral views, Ie, n =250 JUII, I, s = 100 11m, m, 0 =size not indicated, q = 155 J1ffi. p) Ventral infra-
ciliature in dorsal view, 375 !lm. r) Ventral infraciliature of a specimen after conjugation, 95 !lm. Preceding
stages, see Figures 13a-g. t) Dorsal view of a very small (75 !lm) specimen; p 504.

E- Fig. 151e-i Stylonychia mytilus complex (e, f, from GELEI 1954b; g, from CHORIK 1968; h, after TUFFRAU
1965a from TUFFRAU 1987; i, from TUFFRAU 1969. e, sublimate fixation; f, g, from life; h, i, protargol impreg-
nation). e, g) Ventral views, e = about llO !lm, g = 150 !lm. 1) Arrangement of transverse cirri in ventral view.
b) Ventral infraciliature, 235 !lm. i) Early morphogenetic stage in ventral view showing oral primordium (ar-
row); p 504. j Stylonychia mytilus (from AMMERMANN & SCHLEGEL 1983). Ventral infraciliature redrawn from
a scanning electron micrograph, 300 11m; p 504, 549.
 520 SYSTEMATIC SECTION

z a
Fig. 151u-z, 152a Stylonychia mytilus complex (u, after WRZESNIOWSKlEGO 1867a, b; v, after Roux 1901; W,
from FAVRE-FREMIET 1909; x-z, after QUENNERSTEDT 1865; a, after LEWIN 1912. u-z, a, from life). u-w, z, a)
Ventral views, u = 330 f.lm, v = 340 f.lm, w, Z =? f.lm, a = 205 f.lm. x, y) Infection with suctors; p 504.
 Sty/onychia 521

Fig. 152b--e Stylonychia mytilus complex (b, from DRAGESCO 1966b; c, after MERESCHKOWSKY 1877; d, after
WANG 1925; e, from JONES 1974. b, protargol impregnation; c--e, from life). b) Ventral infraciliature of the sup-
posed synonym Stylonychia kohli, 200 11m. c-e) Ventral views, c = size not indicated, d = 165 11m, e =
133 11m. Note that the specimen in (c) has only 2 caudal cirri, an anomaly which can be observed relatively of-
ten in S. mytilus; p 504.

Fig. 2261). The illustration in REDEKE (1948) is a redrawing from STEIN (1859b). Stylony-
chia mytilus in ALEKl'EROV (1992, P 132) is a misidentification (possibly S. pustulata).

M 0r p hoI 0 g y and b i 0 log y: See also characterisation of Stylonychia, addi-

tional characters, and key to species for some characters. Size rather variable, for exam-
ple, 87-347 ~m (STEIN 1959b), 100-300 ~m (KAHL 1932), 280-375 ~m (BUISAN 1944),
200-400 ~m (EDMONDSON 1906), 80-160 ~m (JONES 1974), lOO-l30 x 40 ~m (GELEI
1954). According to AMMERMANN & SCHLEGEL (1983), S. lemnae - the smaller of the two
species of the complex - is about 230 ~m long on average, whereas S. mytilus measures
about 300 ~m. However, WIRNSBERGER et al. (1986) demonstrated that body size and
shape (see below) show transitions, indicating that they cannot be used for separation as
suggested by AMMERMANN & SCHLEGEL (1983; Table 31).
 522 SYSTEMATIC SECTION

. .
\

.::~
.~ \
~

~
f

'.
~.. ~ "
·· "'" \,,
~ ~ ~
t

· ., . •• 1

\ . \.
I • 1 •

·· .
-"\

.
"

''.
;·,, '" '\ ,
:
i .

h
9
Fig. lS2f-b Sty/onychia myli/us complex (f, from WALLENGREN 1900a; g, after FELLERS & ALUSON 1920; h,
slightly modified from MACHEMER 1965b. f-h, from life). 1) Arrangement of frontal, ventral, and transverse
cirri. Cirri originating from the same streak are connected by a dotted line. g, b) Ventral views, g, h = size not
indicated. Identification of (g) uncertain; p 504.

Outline usually very typical, namely with usually striking bulge in region of peri-
stome (for example, Fig. 148a, 0, 151k, n). Body margins slightly to distinctly converging
posteriorly, left margin usually concave, right margin slightly to distinctly convex. Poste-
rior end usually ± truncated, sometimes rather broadly rounded. Flattened about 2: 1 dor-
soventrally, first and last quarter of cell very thin (Fig. 151 c); especially posterior portion
thus usually rather translucent.

Fig. lS2i Sty/onychia myli/us complex (from TUFFRAU 1965a). Ventral infraciliature and associated fibrilles ~
after protargol impregnation. Explanation of original labels: A = "eventail" retrociliaire perpendiculaire au plan
des membraneles adorales, BI-B3 = lames fibreuse sous-ectoplasmiques issues des bases des cirres mar-
ginaux et ventmux, CI, C2 = fibres radiculaires des cirres de la face ventrale, D = fibres cytopbaryngiennes,
E I, E2 = grandes racines ciliaires des bases marginales, F = faisceau des fibres longitudinales issues des cirres
transversaux, G = fibres de connexion adorales, HI, H2 = fibres tangentielles des plages anterieure et poste-
rieure, I = "termina filament" de LUND, J = fibres ramifiees sous-adorales, K = petites racines ciliaires des cirres
marginaux; p 504.
 Stylonychia 523

81_-,.-,

;:;;;;;:--_ _ _ J

~~~~~~--- __----E2

~M,'~~~~_-----K
 524 SYSTEMATIC SECTION

I /'.
I , ,,
/ ~

Fig.152j-m Sty/onychia mytilus complex G, from POTIER 1900; k, I, from LQpEZ-OcHOTERENA 1965; In, from
BERECZKY 1977a.j-1, from life; m, method not indicated). j-m) Ventral views,j, I, m = size not indicated, k =
115 ~m; p 504.

Macronuclear nodules often connected by fme strand, usually arranged about in me-
dian of cell. Anterior nodule sometimes displaced more to right than posterior nodule.
Nodules in life about 40-45 x 15-25 11m (in large to middle-sized populations), after
protargol impregnation 28 x 11 11m (23-32 x 9-13 11m; n = 15 11m) to 39 x 14 11m
(31-45 x 11-17 11m; n = 25) in S. mytilus and 26 x 10 11m (19-38 x 9-11 11m; n = 15) to
32 x 13 11m (25-39 x 10-15 11m; n = 25) in S. lemnae. Usually 3-4, according to JICKELI
(1884) up to 6 micronuclei; in life 6-7 11m across. Cultivation at supra-optimum tempera-
ture (30°C) yields a significantly higher number of micronuclei (namely 6) than in con-
trol cells (3 on average; KAUL & SAPRA 1986). Many small crystals homogeneously dis-
tributed in cytoplasm (HAUSMANN 1982). Contractile vacuole pulsates about every 8-13 s
(GELEI 1935); takes 45 min at 18°C to excrete the amount of water corresponding to cell
volume (BRAND 1935, MAUPAS 1883). Rapid movement, comprising 4 fundamental forms,
namely swimming, running, resting, and reversing movements (see MACHEMER 1965b for
details).
Adoral zone of membranelles 50-55 % of body length. Frontal and transverse cirri
and buccal cirrus distinctly enlarged. Frontal and transverse cirri in life about 40-50 11m,
 Sty/onychia 525

Fig. 152n, 0 Sty/onychia myfilus complex (from CONN 1905). Ventral views from life, n = 170 11m, 0 =
155 11m; p 504. P Sty/onychia nOfophora (from LOPEZ-OcHOTERENA 1965). Ventral view from life (1), 120 11m;
p 553.

marginal cirri in life. 20-30 J.1m long. Frontoventral cirrus 11112 slightly behind level of
frontoventral cirrus IV/3 (Fig. 149a) or somewhat in front of it (Fig. 150a); possible this
is a small, but significant difference between S. mytilus and S. lemnae which can be seen
even in live specimens. Only the 2 right transverse cirri protrude distinctly beyond poste-
rior end of cell. Usually 6, rarely 7 dorsal kineties. Caudal cirri widely separated, in me-
dium- and large-sized specimens in life about 50-70 J.1m long, posteriorly usually fringed.
Resting cyst with a solid, wavy ectocyst (Fig. 148j, 1501; see also ILOWAISKY 1915,
1926b). After STEIN (l859b) in life 55-70 J.1m, rarely up to 105 J.1m, according to BRAND
(1923) 108-128 J.1m across, belongs to kinetosome-resorbing type (WALKER & MAUGEL
1976). All mineral concretions are eliminated from cell during encystment (RELlA et al.
1988). The observations ofCIENKOWSKY (1855) on cyst formation are rather superficial.
 526 SYSTEMATIC SECTION

Fig. 152q-t Stylonychia mytilus complex (q, r, from EDMONDSON 1906; s, from LoKOT 1987; 1, from EBERHARD
1858. q-t, from life). Ventral views, q = 250 Jlm, r = 245 Jlm, s = 150 Jlm, t = size not indicated. Figure q pos-
sibly shows a different species as correctly stated by EDMONDSON (1906) in that he identified it as "Stylonychia
sp."; p 504.

M 0 r p hog e n e sis. More or less comprehensive contributions to divisional morpho-

genesis of species of the S. mytilus complex are provided by BALBIANI (1860a), DEM-
BOWSKA (1925), FRICK (1968), GELEI (1927), HEMBERGER (1982), I<AMRA & SAPRA
(1993a), STEIN (1859b), TCHANG et al. (1965), WALLENGREN (1902), WATANABE & HORA
(1983), Qru et al. (1996), and in most detail by WIRNSBERGER et al. (1986). The latter
authors did not find any differences in morphogenesis of S. mytilus and S. lemnae. Thus,
the following statements and drawings (Table 4, Fig. 23f, 153a-j) are valid for both spe-
cies. Stomatogenesis commences with formation of a small group of basal bodies very
close to the left transverse cirrus (cirrus III I ; Fig. 151 i, 153a; TUFFRAU 1970). Sometimes,
this group originates also very close to transverse cirrus IIIII; however, transverse cirri

Fig. 153a-f Stylonychia mytilus complex (from WIRNSBERGER et aI. 1986. a--c, e, f, S. lemnae; d, S. mylilus. ~
a-f, ventral infraciliature of morphogenetic stages after protargol impregnation). a) Very early stage with ori-
gin of oral primordium (arrowhead), 175 Jlm. b, c) Early stages, b = 190 Jlm, c = 195 Jlm. Arrowhead marks
postoral ventral cirrus V/4, which is modified to a primordium. Arrow marks oral primordium. d) Early stage.
Only anterior portion of cell is shown. e, f) Middle stages with a supernumerary ventral cirrus (f, arrowhead), e
= 215 Jlffi, f= 190 Jlm; further explanation see text; p 504.
 Stylonychia 527

.'.:

:'...

'j
~

<'

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ow. "
" ~
l!
" ~
". "
~
" o t·
\~ ~~"i
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Cl 0 0
o ., :'/ Q
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153 3

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I) ()
q
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q
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528 SYSTEMATIC SECTION

"
"
"
o"
"
""
"
to
",':. ()
\\.0
\~~
0
()t::::J " "
() 00
o
'"'"
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<>() o 0
0 t:;
'"
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153g .a
a h
 Stylonychia 529

. .
. :
.. .,f
• ..,,:

(J
0
C)
0 (]

~
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I)

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¢
~
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~
k

Fig.lS3k, 1 Sty/onychia mytilus complex (from FLEURY & FRyo-VERSAVEL 1981. k, I. protargol impregnation).
Ventral and dorsal infraciliature of a reorganiser, 200 Jim. Arrowhead in (I) marks fragmentation of new dorsal
kinety 3. Details on reorganisation, see chapter 1·10·4 in the general section; p 504.

~ Fig. IS3g-j Sty/onychia /emnae (from WIRNSBERGER et aI. 1986. g-j, ventral infraciliature of morphogenetic
stages after protargol impregnation). g, b) Late stages, g = 160 Jim, h = 200 Jim. i, j) Very late stage and opis-
the immediately after cell division, i = 190 Jim, j = ISO Jim. Cirri originating from individual primordia are
connected by a broken line; new structures black, parental white; further explanation, see text. DP = primordia
of dorsomarginal kineties, MP = primordium of right marginal row of proter; p 504.
 530 SYSTEMATIC SECTION

..--
.
•• ,"'t ;;.

/.: .( ~
( )J
..
.

9
Fig. 154a-il Sty/onychia pustulata (a, from TAl 1931; b, from CELA 1972; c, from YANO & SUHAMA 1991; d,
after SMITH 1914. a, d, from life; b, from life and after silver impregnation (?); c, protargol impregnation). a, b,
iii) Ventral views, a = 130 j.lm, b = 60 j.lm, d = size not indicated. c) Ventral infraciliature of very early morpho-
genetic stage. OP = oral primordiwn; p 565.
Fig.l54e Sty/onychia notophora (after SMITH 1914). Ventral view from life, size not indicated; p 553.
Fig. 154f-i Sty/onychia myti/us complex «(after SMIDI 1914; g, from CHARDEZ 1967; h, from MAMAEvA
1979b; i, from TAl 1931. f-i, from life). f-i) Ventral views, f= size not indicated, g = 155 j.lm, h = 200 j.lm, i =
165 j.lm; p 504.
 Sty/onychia 531

I
I

I
I n

Fig. 154j-p Sty/onychia mytilus complex (j, from CHARDEZ 1971; k, from BIERNACKA 1963; I, from BUlsAN
1944; In, from BORGER 1908; n, from SCHUSTER 1899; 0, from CONN 1905; p, from BARWICK et al. 1955. j-p,
from life). Ventral views, j = 190 /lm, k = size not indicated, I = 315 /lIn, m = 200 /lIn, n = 240 /lm, 0 = size
not indicated, p = 225 /lm; p 504.
532 SYSTEMATIC SECTION

remain unchanged. The number of basal bodies increases and a longish oral primordium
is formed (Fig. 153b). Only the anterior portion of the parental undulating membranes are
reorganized (Fig. 153h, i). The primordium for the undulating membranes of the opisthe
originates from the oral primordium (Fig. 153e-g). At this time, the first new adoral
membrane lIes occur. In the opisthe the frontal-ventral-transverse cirri primordia I-III
originate from the oral primordium (Fig. 153f, g). The primordia IV-VI are produced by
the right postoral ventral cirrus (cirrus IVl4; Fig. 153d-g). In the proter, primordium I
originates from parental undulating membranes (Fig. 153g-i); primordium II from oral
primordium of opisthe (Fig. 153d-f); primordium III from frontoventral cirrus 11112 (Fig.
153d, f, g); primordium IV from frontoventral cirrus IV/4 (Fig. 153d, f); primordia V and
VI from right postoral ventral cirrus (Fig. 153d-f). The following cirri do not participate
in the formation of primordia: frontal cirri Ill, 1113, II1I3, the buccal cirrus 1112, the fron-
toventral cirri VII3 and VII4 (= migratory cirri), the postoral ventral cirri IV/2 and V/3,
and the transverse cirri. The left frontoventral cirrus IIII2 and the postoral ventral cirrus
V14 proliferate basal bodies and are partly dissolved (Fig. 153f-h). The rest is resorbed in
later stages of division. The development of the dorsal infraciliature proceeds in Oxytri-
cha pattern (Fig. 24a).
FRICK (1968) found that during regeneration of S. mytilus fragments different varia-
tions in the formation of cirri are possible. Especially primordium IV tends to vary.
Sometimes all cirri of this row are lacking, sometimes only cirrus IV12 or IV13 is absent.
Occasionally, 1-3 additional primordia occur, usually resulting in an increased number of
postoral ventral, pretransverse ventral, and transverse cirri. FRICK (1967) made experi-
ments on the regeneration of amicronuc1eate specimens obtained by B-irradiation and by
microsurgery. He found that regeneration is strongly disturbed and that all specimens
without micronucleus die after 3-4 divisions. Cells ofa species of the S. mytilus complex
cut in mid-body are completely regenerated after 5-7 h (DEMBOWSKA 1925, see also
DOROSZEWSKI & RAABE 1966 and FAuRE-FREMIET 1967); according to JANISCH (1959), this
kind of regeneration takes 6-24 h. Detailed investigations on regeneration were also per-
formed by ISHIKAWA (1912). TOTWEN-NOWAKOWSKA (1965) applied thermal shock consist-
ing in the rise of temperature from 20°C to 33-34 °C to S. mytilus cultures. As result,
doublets of two different types appeared: (i) regular spherical doublets consisting of2 in-
dividuals of a similar structure, joined together by their dorsal side, and (ii) irregular flat
doublets composed of individuals of uneven dimensions and shape, connected by their
sides. They reproduced by transversal cell division and could be maintained in culture for
many generations, that is, for several months. In tap water without food, S. mytilus lived
14-19 d reorganising the whole body several times (DEMBOWSKA 1938). There are sev-
eral papers concerning the morphogenetic processes during regeneration, reorganisation,
and conjugation, including that of doublets (BALBIANI 1858, FENG et al. 1990; FLEURY &
FRYD-VERSAVEL 1981, see Fig. 153k, 1; FRANKEL 1989, 1991, GRIMES & ADLER 1978,
GRIMES & L'HERNAULT 1979, Gv et al. 1990, JERKA-DzIADOSZ & FRANKEL 1990, Lv et al.
1991, NG 1990, PANG et al. 1984, PROWAZEK 1903, SCHILD 1921, SHI 1975, 1976, 1987,
SHI & FRANKEL 1990, SHI et al. 1987, 1990, 1991a, b, TCHANG & PANG 1965a, b, TCHANG
 Stylonychia 533

et al. 1965, 1981, TOTWEN-NoWAKOWSKA 1969, 1973, TUFFRAU & TOTWEN-NoWAKOWSKA

1988, TUFFRAU et al. 1978a, b, ZHANG & PANG 1979, 1981, Zou & NG 1991a-c, Zou et al.
1989).
Species of the Stylonychia mytilus complex are sometimes attacked by parasitic suc-
torians (see chapter 4 in the general section).
Further literature. Species of the Stylonychia mytilus complex are one of the most
popular hypotrichous ciliates. Most of their popularity stems from their intensive use in
experimental studies, especially in areas of genetics, physiology, biochemistry, and ultra-
structure. Since no comprehensive compilation of the literature is available I will list ad-
ditional papers, including some old ones; some further papers are mentioned at S.
lemnae. Biochemistry, genetics: ALLEN & OUNS (1984), ALONSO (1975,1978), ALONSO &
JARENO (1974), ALONSO & PEREz-SILVA (1966a, b, 1969), AMMERMANN (1965b, 1967,
1969, 1970, 1971a, 1973 [see Fig. 148q, r], 1979), AMMERMANN et al. (1978, 1981), BAL-
BIANI (1861, 1862), BOTsCHLI (1876, p 324), CHADHA et al. (1978, 1979), CORLISS (1990),
DAVIES & WALKER (1979), DUTTA et al. (1982), FAlIN et al. (1998), FANG & CECH (1991),
FEDRIANI et al. (1976a), GRELL (1951, 1965), ILOWAISKY (1926a), JARENO (1972, 1974,
1976, 1990, 1993), JARENO & ALONSO (1973), JARENO et al. (1969a, b, 1970, 1972), KAUL
(1989, 1991a, b), KAUL & SAPRA (1981), KAUL et al. (1990), KLOETZEL (1969), KNOWLES
et al. (1978), KRAMER (1988), KRAUT & LIPPS (1984), KRAUT et al. (1986), LINGNER et al.
(1994), LIPPS et al. (1974), MEYER & LIPPS (1980, 1981, 1984), MOLDENHAUER (1965),
MURTI et al. (1972), OLINS et al. (1993), PEREZ-SILVA et aI. (1969), POPOFF (1907), PREs-
corr (1989), PREscorr et al. (1971), RAIKov (1969), RAIKov & AMMERMANN (1974),
RESNITSCHEK (1972), RmWE & LIPPS (1977), SALVINI et al. (1993), SAPRA & AMMERMANN
(1973, 1974), SAPRA & KLOETZEL (1974, 1975), SAPRA et al. (1976), SCHLEGEL (1985),
SCHLEGEL & STEINBROCK (1986), Sm et al. (1987), STEINBROCK (1980, 1981, 1990, 1993),
STEINBROCK & JACOB (1997), TCHANG & PANG (1974, 1979a), TCHANG et aI. (1981), TOR-
RES et al. (1979a), TOTWEN-NoWAKOWSKA (1964), WATANABE et al. (1986), WILSON &

SOMMERVILLE (1987), WONNING & LIPPS (1983), Xu & Sm (1987), YOUNG (1926). Physi-
ology: BALBIANI (1882), BUJWm-CWIK & DRYL (1971), DEITMER (1981a, 1983a, 1987,
1989), DEITMER et al. (1986), DEMBOWSKA (1939), DRYL (1963, 1965), DRYL &
NOWAKOWSKA (1966), DRYL & TOTWEN-NoWAKOWSKA (1972, 1975, 1977, 1985), DRYL et
al. (1981), EFIMOFF (1924), ENGELMANN (1880 ), GELTZER (1967), IVENS (1986), lz.ruMov
(1948), JiROVEC & SOSNA (1954), KUSALSKI et al. (1978), LICHTENBERG (1955b), MA-
CHEMER (1965c, 1966c, 1969a, b, 1970, 1980), MAcHEMER & PEYER (1977), MEIER &
KAUNAT (1954), MOGAMI & MAcHEMER (1991a, b), MOGAMI et al. (1992), PEYER (1976),
PEYER & HUGGEL (1973), PEYER & MACHEMER (1978a, 1979), POPOFF (1908,1909), SAPRA
& DASS (1978), SCHWALBE (1866), SLEIGH (1968), STATKEWITSCH (1904, 1905), TCHANG et
al. (1964), TOTWEN-NOWAKOWSKA & DRYL (1976), YANK & MAcHEMER (1996, 1996a),
WESKAMP (1993), WESKAMP & MACHEMER (1993), ZAIKA (1970). Ultrastructure: CHADHA
(1980), CHADHA et al. (1983), DASS et al. (1976, 1982), ENGELMANN (1875), ESTEVE
(1972), FERNANDEz-LEBORANS & MARTIN-GONZALEZ (1982), GIL (1976, 1979), GIL et al.
(1972), GORTZ (1982), GRIMES (1982a, b), KAUL et al. (1982), KLOETZEL (1970), Kow-
534 SYSTEMATIC SECTION

Table 31 Morphometric data of Stylorrychia hi/aria (bil, bi2, two populations from the Schlossalm area in
Salzburg, Austria, from WIRNSBERGER et al. 1985a), S. lemnae (leI, population from gardening of Schilde in
Wendelsheim, Germany; le2, population from Spitzberg near Tuebingen, Germany; both from WIRNSBERGER et
aI. 1986), S. mytilus (myl, population from a lake near Reutlingen-Gonningen, Germany; my2, population
from a valley of the Kimbach, Olgahein, Germany; both from WIRNSBERGER et aI. 1986), S. pustulata (pul,
population from a lake [Mondsee] in Upper Austria; pu2, pu3, clones of a large and a small specimen from
population pul; from WIRNSBERGER et aI. 1985a pu4, population from a lake [Lunzer See] in Lower Austria;
from GSCHWIND 1991). All data are based on protargol-impregnated specimens. All data, except pul, are from
clones. All measurements in micrometres. CV = coefficient of variation (in %), Max = maximum value, mean
= arithmetic mean, Min = minimum value, n = sample size, SD = standard deviation
Character Species mean SD CV Min Max n
Body, length bil 80.0 6.0 8.0 70 91 28
bi2 91.0 8.0 8.0 75 110 25
leI 186.3 1l.5 6.2 168 211 25
le2 154.9 11.0 7.1 139 180 15
my 1 222.4 14.0 6.3 195 256 25
my2 166.1 14.4 8.8 147 203 15
pul 65.0 13.0 21.0 43 90 25
pu2 94.0 9.0 10.0 79 108 25
pu3 79.0 8.0 10.0 64 93 25
pu4 53.0 6.2 1l.8 44 66 21
Body, width bill 42.0 4.0 11.0 36 52 28
bi2 1 51.0 6.0 11.0 42 62 25
lel 2 78.7 5.4 6.8 70 95 25
le22 60.1 4.2 7.0 52 68 15
myF 1l0.9 8.9 8.1 96 127 25
my2 2 69.5 9.2 13.1 57 87 15
pull 34.0 6.0 18.0 22 50 25
pu2 1 47.0 6.0 13.0 35 62 25
pu3 1 36.0 5.0 15.0 27 48 25
pu4 1 26.4 3.0 1l.5 21 31 21
Body, width at level of middle lei 60.2 7.7 12.9 40 78 25
transverse cirrus le2 46.6 6.7 14.5 36 61 15
my I 66.6 9.3 13.9 45 86 25
my2 42.1 7.7 18.2 25 58 15
Macronuclear nodule, length bil 14.0 1.0 8.0 12 17 28
bi2 14.0 2.0 14.0 12 17 25
leI 31.6 3.6 11.4 25 39 25
le2 26.4 4.6 17.3 19 38 15
my I 38.5 4.9 12.7 31 45 25
my2 27.7 2.5 9.1 23 32 15
pul 14.0 2.0 16.0 9 17 25
pu2 18.0 5.0 28.0 10 28 25
pu3 15.0 2.0 13.0 10 20 25
pu4 11.5 1.5 13.3 9 14 21
Macronuclear nodule, width bil 7.0 1.0 15.0 4 9 28
bi2 7.0 1.0 13.0 5 9 25
leI 13.2 I.3 10.2 10 15 25
le2 9.6 0.8 8.6 9 II 15
my I 13.7 1.5 11.0 II 17 25
 Sty/onychia 535

Table 31 Continued

Character Species mean SD CV Min Max n

Macronuclear nodule, width my2 10.9 1.4 12.5 9 13 15
pul 7.0 1.0 14.0 5 8 25
pu2 9.0 2.0 17.0 7 13 25
pu3 7.0 1.0 14.0 5 8 25
pu4 6.1 0.5 7.3 6 8 21
Micronuclei, number leI 3.4 0.6 18.9 2 5 25
le2 3.5 0.7 21.4 2 5 15
my 1 3.7 0.5 14.2 3 5 25
my2 3.9 1.0 24.4 3 6 15
pu4 2.0 0.0 0.0 2 2 16
Adoral zone of membranelles, bil 40.0 2.0 5.0 36 44 28
length bi2 44.0 4.0 10.0 35 53 25
leI 98.4 3.9 4.0 88 105 25
le2 78.7 5.5 7.0 69 88 15
my 1 118.6 7.7 6.5 102 131 25
my2 92.3 7.6 8.2 82 112 15
pul 35.0 6.0 18.0 25 44 25
pu2 49.0 4.0 9.0 40 56 25
puJ 42.0 4.0 10.0 31 48 25
pu4 29.0 3.5 12.2 24 37 21
Adoral membranelles, number bil 37.0 2.0 5.0 32 40 28
bi2 34.0 2.0 7.0 30 39 25
leI 61.4 2.3 3.7 57 65 25
le2 58.5 2.5 4.4 52 61 15
my 1 76.3 2.5 3.3 71 81 25
my2 68.1 6.1 9.0 52 76 15
pul 35.0 5.0 15.0 24 42 25
pu2 38.0 2.0 6.0 34 42 25
puJ 34.0 2.0 6.0 31 37 25
pu4 32.1 2.8 8.6 27 39 21
Paroral, length leI 61.2 4.9 8.0 55 71 15
le2 65.4 9.8 15.0 47 84 15
my 1 72.4 6.4 8.9 59 81 15
my2 72.8 6.6 9.1 65 85 15
Endoral, length leI 51.4 4.0 7.8 46 57 15
le2 57.5 8.9 15.5 42 73 15
my 1 62.4 4.3 6.9 56 70 15
my2 62.4 7.1 11.3 46 74 15
Right marginal row, number of cirri bil 19.0 2.0 10.0 14 22 28
bi2 19.0 2.0 8.0 16 21 25
leI 34.3 1.6 4.6 31 37 25
le2 35.5 3.1 8.7 31 41 15
my 1 37.4 1.9 5.2 33 42 25
my2 39.3 2.9 7.5 33 44 15
pul 26.0 3.0 13.0 18 32 25
pu2 31.0 2.0 5.0 28 34 25
pu3 27.0 2.0 6.0 25 30 25
pu4 24.2 2.7 11.0 20 29 21
 536 SYSTEMATIC SECTION

Table 31 Continued

Character Species mean SD CV Min Max n

Left marginal row, number of cirri bil 17.0 1.0 5.0 IS 18 28
bi2 17.0 1.0 6.0 IS 19 25
lei 25.0 l.l 4.5 23 27 25
le2 26.5 1.6 6.0 24 29 15
my I 26.7 1.3 5.0 24 29 25
my2 28.0 1.6 5.9 25 30 15
pul 19.0 2.0 13.0 12 22 25
pu2 21.0 2.0 8.0 18 24 25
pu3 19.0 2.0 9.0 16 22 25
pu4 14.9 1.4 9.5 12 18 21
Dorsal kinety 1, number of basal leI 76.8 3.0 3.9 74 81 5
body pairs le2 72.0 5.1 7.2 65 78 5
my 1 97.2 7.1 7.3 88 106 5
my2 100.4 10.5 10.5 85 III 5
Dorsal kinety 3, number of basal leI 36.8 2.8 7.5 33 40 5
body pairs le2 37.6 0.9 2.4 37 39 5
my I 64.0 13.8 21.6 44 66 5
my2 60.6 5.8 9.5 53 66 5
Dorsal kinety 4, number of basal lei 33.6 3.5 10.4 30 39 5
body pairs le2 33.0 2.0 6.1 31 36 5
my I 51.8 3.3 1.9 47 56 5
my2 47.8 6.8 14.3 40 55 5
All dorsal kineties, total number leI 284.6 13.7 4.8 271 304 5
of basal body pairs le2 271.8 10.9 4.0 258 286 5
my I 369.9 11.8 3.2 358 389 5
my2 363.0 35.4 9.8 318 398 5
Distance between first leI 27.2 4.0 14.8 19 34 25
and third caudal cirrus le2 24.7 2.1 8.5 21 27 15
my I 35.3 3.8 10.7 26 42 2
my2 26.9 4.3 15.9 17 35 15

I Maximum width.
2 Width at level of buccal cirrus.

ALEWSKIEGO (1882), MAlER (1903), MAUPAS (1883), MURTI (1972, 1973, 1976), NOBill &
ROSATI RAFFAELU (1971), OLINS et al. (1981), PESCHKOWSKY (1927), PRowAZEK (l899a),
PuYTORAC & RODRIGUES DE SANTA ROSA (1976), PuYTORAC et al. (1976, 1994), STERKI
(1878, see also LUND 1941), TAYLOR (1964), TUFFRAU (1964), TUFFRAU et al. (1968),
WALKER (1976a)," WALKER & GOODE (1975,1976), WALKER et al. (1975).

Occurrence and e colo g y: Widely distributed in freshwater habitats through-

out the year with peaks in spring and autumn (for example, BICK & KUNZE 1971, HECK-
MAN 1982, own observations). Common in the benthal of stagnant (lakes, reservoirs,
ponds, temporal small water bodies) and running waters (for example, BICK & SCHMEREN-
BECK 1971, CZAPIK 1982, I<AHL 1932, RIGGENBACH 1922, SCHMERENBECK 1975, SPANDL
 Stylonychia 537

1926a). Sometimes planktonic in ponds and lakes (for example, FOISSNER et al. 1991) and
large rivers (for example, PoJIN et al. 1984); rarely in brackish and sea water; onlyocca-
sionally in terrestrial habitats.
Locus classicus is Denmark, where MOLLER (1773, 1786) discovered Sty/onychia
mYlilus in lentic water. EHRENBERG (1830, 1833, 1838) recorded it abundantly among Os-
cil/aloria in all infusions and in waters in Berlin, Germany; he found it also in Copenha-
gen in sea water. STEIN (1859b) observed S. mYli/us among Lemna, callitrichs, filamen-
tous algae ("Conferven"), and decaying macrophytes, where largest specimens occurred.
Locus classicus of the synonym, S. clavi/ormis, is a eutrophic pond in the Borzsony
Mountain, Hungary (GELEI 1954, GELEI et al. 1954).
Further records substantiated by illustrations: alpine grassland (2310 m above sea
level, alpine pseudogleyic on mica-slate) in the Glockner Area, Austria (FOISSNER 1981,
1982, 1987a); plankton of hypertrophic pond at Salzburg University, Austria (BLATIERER
1988, FOISSNER et al. 1991); Belgium (CHARDEZ 1971); very common in freshwater and in
flood overflows in England (SCHUSTER 1899); rock pools with 0.1 to 3.6 %0 NaCI in Fin-
land (REUTER 1961); France (DRAGESCO 1966a, DUJARDIN 1841, FROMENrEL 1876, TUFFRAu
et al. 1981); common, but usually only moderately abundant in mesosaprobic running wa-
ters in Germany (BUCK 1959, 1961); water bodies near Leipzig, Germany (ENGEIMANN
1862); mesosaprobic Danube River in Hungary (BERECZKY 1977a); estuaries in the Dan-
zig Bay with ~ %0 salt (BIERNACKA 1963) and other sites in Poland (WRZESNIOWSKIEGO
1867a, b); among algae in stagnant water in Spain (BUIsAN 1944); among Lemna in Swe-
den (QUENNERSTEDT 1965); clear and polluted ponds near Geneva, Switzerland (Roux
1901); plankton and benthos of large rivers (MAMAEVA 1979a, b) and other sites in the
USSR (CHoRIK 1968, LOKOT 1987); small stream on the campus of National South West
Associated University, China (CHEN 1944); common in pond water from the campus of
Tsing Hua University, China (TAl 1931); Lake Ho Hu and Lake Mo Chu Hu, Nanking,
China (WANG 1925) and other sites in China (TCHANG & PANG 1965a); Mysore soils, In-
dia (RAo 1928); Han River and Ullung Island, Korea (SHIN & KIM 1988, 1993a, 1995);
pond in Japan (yOSHINO 1993); freshwaters in Connecticut, USA (CONN 1905); infusions
of pond water in Iowa, USA (EDMONDSON 1906); northern area of Mobile Bay, USA, with
a salinity of2 %0 (JONES 1974); soils near New Brunswick, New Jersey, USA (FELLERS &
AlliSON 1920); pond water in Kansas, USA (SMITH 1914); Mexico (L6PEZ-OCHOTERENA
1965); Chile (BORGER 1908); Benin (DRAGESCO & DRAGESCo-KERNEIS 1986); abundant
among decaying vegetation in Waiwhetu stream, New Zealand (BARWICK et al. 1955).
Records from freshwater habitats (pools, ponds, lakes, brooks, rivers etc.) not sub-
stantiated by illustrations - Europe and Asia: Armenia (ZHARIKOV 1982, see also USSR);
rarely in beta- and beta- to alphamesosaprobic brooks and rivers in Austria (AOOLR
1994b, FOISSNER & MOOG 1992, HAsLAUER & HAIDER 1976, HAsLAUER & PICHLER 1979);
littoral of Danube River in Vienna, Austria (HUMPESCH & MOOG 1994, KALTENBACH
1960); alpine lakes (2000 m above sea level) in Tyrol and clear lakes in Styria, Austria
(DALLA TORRE 1891, KIwNER 1988, ZACHARIAS 1903b, ZSCHOKKE 1900); profundal (64 m)
ofCarinthian and Tyrolian lakes, Austria (FINDENEGG 1953, IMHOFF 1895); smalliotic and
538 SYSTEMATIC SECTION

lentic waters in the Austrian Alps (FOISSNER 1980c, FOISSNER & ADAM 1979); in a several
weeks old water with mouldy odour, Austria (SCHMARDA 1846); infusion of submerged
mosses from Upper Austria (FOISSNER 1974); abundant in long standing samples with
mud, algae, and decaying macrophytes, Austria (VORNATSCHER 1938; further records from
Austria: BRUNNTHALER et al. 1901, CZERNIN-CHUDENITZ 1985, FOISSNER & FOISSNER 1988,
HAsLAUER et al. 1979); rivers in Azerbaijan (ALlEv 1982a, 1984; see also USSR); Bel-
gium (BERVOETS 1940, CHARDEZ 1987, SORGELOOS 1970); brooks, lakes, ponds, peat-bogs;
marshland, pools, wet meadow, fishponds, road ditches, and rivers in Bulgaria (DETCHEVA
1979b, 1981, 1986, 1991, 1993, DETSCHEWA 1972, RUSSEV et al. 1984); saprobic brooks
and polluted river in Czechoslovakia (BuCHAR 1957, MATIS 1967, NOWAK 1940); moder-
ately frequent in betamesosaprobic water bodies in Czechoslovakia (SRAMEK-Hu~EK
1956b, 1957, V ~ICEK. 1964); common in the littoral of a pond in Czechoslovakia, large
specimens especially in the foam on the water (SVEC 1897); submerged and wet mosses in
Czechoslovakia (TIRJAKovA & MATIS 1987a; further records from Czechoslovakia: BtLy
et al. 1952, HAEMPEL 1908, HANu~KA 1962, KALMUS 1928, MATIS 1961, 1977, MATIS &
TIRJAKovA 1994, MORAVCOVA 1962, SRAMEK-HU~EK 1953, TIRJAKovA 1992b); pond in
Denmark (FENCHEL 1980a, c); chalk streams in England (BALDOCK et al. 1983, GRAY
1952); in 8 of 178 sites of sewage fungus outbreaks in England (CURTIS & CURDS 1971);
eutrophic Esthwaite water, England (WEBB 1961; further records from England: CRAIGIE
1921, DIXON 1920); Estonia (JACOBSON 1928); polluted rivers in France (GROLrERE et al.
1990, ORMANCEY 1852, SPARAGANO & GROLrERE 1991); pond and peat-bog moss habitats
in France (GROLrERE 1977, GROLrERE & NJINE 1973); in the clean Illach River in Germany
(ForSSNER 1997a); common in Rhine River plankton, Germany (CZERNIN-CHUDENITZ
1958); rarely in the beta- to alphamesosaprobic Rhine River, Germany (MAUCH 1990);
cooling system of a power station at the beta- to alphamesosaprobic Main River, Ger-
many (BERNERTH 1982); aufwuchs and sandy sediment of betamesosaprobic Elbe River
upstream of a dam, Germany (GRIMM 1968); among detritus and in surface film of ditches
regularly contaminated with pesticide sprays and in unsprayed ditches in the Elbe River
floodplain, Germany (CASPERS & HECKMAN 1982); beta- to alphamesosaprobic ditch in
Germany (HEuss 1975); rarely to abundantly in beta- to alphamesosaprobic brooks and
rivers in Germany (FOISSNER et al. 1992a, b, HEuss et al. 1972, JUTRCZENKI 1982, MAUCH
& WITILING 1991, SCHEUBEL 1986, SCHONBORN 1996); abundant at 10 cm and sparse at
25 cm depth of Fulda River Headwater sediment, Germany (LOPKES 1976); ponds near
Bonn and 66 indo mtl in small basins under laboratory conditions, Germany (BICK 1958,
1964a, 1967b, KRAMER 1964); road pools, flooded meadows, and infusions of agricul-
tural soils in Bavaria, Germany (DINGFELDER 1962); effluent ditch (3 mg 1-1 PtotaI,
13.4 mg 1-1 COD) of the Schillinger Lake, Germany (GUHL 1987); littoral of Lake Con-
stance~ Germany (KUTTNER 1923); mesotrophic lakes in Germany (MOCKE 1979, PACK-
ROFF 1992, PACKROFF & WILBERT 1991, ZACHARIAS 1902); rarely in an oligosaprobic to be-
tamesosaprobic reservoir and about 40 indo ml- l in an alphamesosaprobic reservoir in
Germany (NusCH 1970); pseudoperiphyton of a eutrophic pond (Poppelsdorfer Weiher)
in Bonn, Germany (MICHIELS & WILBERT 1973, SONG & WILBERT 1989, WILBERT 1969);
 Stylonychia 539

very abundantly in a marshy ditch (mud contained H2S) near the Schussen River, Ger-
many (WETZEL 1928a); aufwuchs community and detritus in Hamburg Harbour, a highly
stressed habitat in the freshwater section of the Elbe Estuary, Germany (BARTSCH &
HARTWIG 1984, HECKMAN et aI. 1990, RIEDEL-10m 1981, Roy 1938, TENT 1981); marshy
ditch and lakes in Germany (ZACHARIAS 1893, 1894, 1903a; further records from Ger-
many: BAUER 1987, DORR 1935, HENDERSON 1905, PACKROFF & ZWICK 1996, PATSCH
1974, QUIRMBACH 1912, SCHMIDT 1913, 1916, SCHNEIDER 1909, ScHONBORN 1984,
ZACHARIAS 1897); Greece (STEPHANIDES 1948); up to 16 indo 1'1 in plankton of mesosapro-
bic Danube River and in a oxbow in Hungary (BERECZKY 1971, 1972, 1977b, c, 1979,
1991, NOSEK & BERECZKY 1981); aufwuchs and pelagial of Tisza River, Hungary (J6SA
1974, STILLER 1942); at pH 8.1 in a fishpond in Hungary (JAcz6 1939); common and to-
gether with S, pustulata in marshy areas, in the aufwuchs community, and detritus drifts
of the littoral of Lake Balaton, Hungary (DADAY 1885, ENTZ 1897, FRANCE 1897, GELLERT
& TAMAs 1958, 1959b, 1960, TAMAs & GELLERT 1959; further records from Hungary:
BERCZIK 1966, DADAY 1892, DUDICH 1947, HAMAR 1971, KREPuSKA 1917, VOROSVARY
1950); river and lough in Clare Island, Ireland (DUNKERLY 1913); various waters in
Northern Italy (CANELLA 1954, MADaNI 1980, MADaNI & ROSSI 1977); characteristic for
medium hilly zone in Italy with narrow water course, pebbled bed with sand and with
Frontonia leucas as typical species (MAnoNI 1984); 2 indo cm,2 in an alphamesosaprobic
area of the Po River, Italy (MADONI 1979); rarely in beta- to alphamesosaprobic running
waters in Italy (MADaNI & GHETTI 1980, 1981a); Lake Como, Italy (CATTANEO 1882); rice
paddies in Italy (COPPA 1921, MADaNI 1988b; further records from Italy: CuNEo 1891,
ENRIQUES 1913, GARGIULO 1907, GRANDORI 1934, LONGHI 1894, 1895, MADaNI 1983, MA-
DONI & VIAROLI 1985, SACCHI 1888, STELLA & CAMPEA 1948); lakes and polluted rivers in
Latvia (LIEPA 1981, 1983, 1990); the Netherlands (DoRSMAN 1918, GRMF 1956, 1957,
GRMF & MEDER 1956); betamesosaprobic and polysaprobic areas in a river in Poland
(CZAPIK 1982); with 35 % frequency in the Sphaerotilus community of a Polish river
(HUL 1986); beta- to alphamesosaprobic and alphamesosaprobic areas of rivers in Poland
(HUL 1987a, b, WIACKOWSKI 1981); reservoirs in Poland (CZAPIK 1962, KRzYZANEK &
KRzyfANEK 1986); at a saprobic index of 1.9-2.6 in a Polish river (GRABACKA 1985); mud
of polluted fish ponds and fish ponds with mowed plants in Poland (CZAPIK 1959, GRA-
BACKA 1971, 1977, KWIATKOWSKA-GRABACKA 1964, 1965, SJEMINSKA & SJEMINSKA 1967);
aquarium with Lemna and decaying algae in Poland? (WRZESNIWOSKI 1861; further re-
cords from Poland: CZAPIK & FYDA 1995, JAWOROWSKI 1893, KOMALA & PRzYBOS 1980,
1988, PAWLOWSKI 1980, WRZESNIOWSKlEGO 1866); Azores, Portugal (BARROIS 1896, IMHOF
1896); abundantly in mud from Danube River in Romania (SPANDL 1926a); Turiec River
in Slovakia (TIRJAKovA 1993); marshy ditch in Slovenia? (SCHMARDA 1847); clean section
of Henares River, Spain (SOLA et al. 1996); up to 6.6 indo mI'l in upper sediment and up
to 2 indo ml'1 in neuston of a eutrophic reservoir near Barcelona, Spain (SALvAD6 CABRE
1993, SALVADO & GRACIA 1991; further records from Spain: FERNANDEZ-LEBORANS et al.
1990, GRACIA & IGUAL 1987a, b, GRACIA et aI. 1987a, 1989, IGUAL 1990, MARGALEF L6PEZ
1945); Danube River in Slovakia (SZENTIvANY & TIRJAKovA 1994); Sweden (QUEN-
540 SYSTEMATIC SECTION

NERSTEDT 1869); with low abundance in clean and polluted running waters and in lake
plankton in Switzerland (STEINMANN & SURBECK 1918, 1934); shallow lake with Phrag-
mites and leaflitter in Switzerland (HEusCHER 1890); fresh and old samples from marshy
waters in Switzerland (PERTY 1852b); littoral and profundal of Lake Geneva (ANDRE
1916, FOREL 1904, Roux 1900); alpine lakes in Switzerland (ASPER & HEUSCHER 1886);
peat-bogs in Switzerland (MERMOD 1914); ponds and ditches near Bern and near Neuen-
burg and Lake Neuenburg, Switzerland (CDANN 1907, MONARD 1920, SAKOWSKY-
CAMPION! 1906, THIEBAUD & FAVRE 1906; further records from Switzerland: ANDRE 1912,
1915, BOURQUIN-LINDT 1919, MESSIKOMMER 1954, RIGGENBACH 1922, ZSCHOKKE 1894);
alphamesosaprobic aufwuchs in the Save River, Yugoslavia (PRIMc 1981; further records
from Yugoslavia: DJUKIC et al. 1994, PRonc 1933, PuJIN 1994); in the epiphyton of
50-75 % of collected samples from karstic running waters in Yugoslavia (PRIMC-HABDUA
& HABDUA 1991); well in Yugoslavia (KARAMAN 1935); lake in China (SHEN 1980); to-
gether with Paramecium aurelia in long standing samples, USSR (EICHWALD 1844, 1847,
1849); meso- and eutrophic lakes in USSR (KoRNIYENKO 1972); common in Lake Baikal
(GAJEVSKAJA 1927, GAJEWSKAJA 1933); benthic and planktonic in slightly to distinctly pol-
luted reservoirs and rivers of the USSR (ALEKPEROV 1980, 1981, 1982b, c, 1983, 1984b,
c, BELOVA 1988, KRAVCHENKO 1969, MYLNIKOVA 1981, NEBRAT 1980, SHADIN et aI. 1931;
further records from the USSR: AoIL 1934, AGAMALIEV & ALlEv 1978, ALEKPEROV 1986,
ALlEv 1988, BELIKHOFF 1936, CHoRIK & VIKOL 1973, GORIDCHENKO 1984, KOSOVA 1965,
MERESCHKOVSKY 1880, MINKEWITSCH 1898, MOVCHAN & PROTASOV 1986, OLEKSIV 1985,
PASHITNOWA 1935, SKORIKOW 1902, WEISSE 1845, ZHARIKOV 1992, ZoLOTAREV 1987, ZYK-
OFF 1903); alphamesosaprobic and polysaprobic puddles with high amounts of rotting
leaves and flowers in Nagpur, India (KruSHNAMOORTHI 1969); Palestine (BODENHEIMER
1937, LEPSI 1929a); leaf litter of a small pond in Tokyo, Japan (HATANO & WATANABE
1981, KUSANO 1985); Sapporo, Japan (HAYASHI 1959); foul water in City ofHakodate, Ja-
pan (MURAMATsu 1957; further record from Japan: MATSUOKA et aI. 1983).
America: ponds, pools, still areas along stream margins, polluted streams, tree or
stump holes, marsh pools in USA (HAUSMAN 1917, LACKEY 1938a, SMITH 1878); soft to
medium hard water rivers in the eastern USA (PATRICK 1961); abundant in tundra pools
(Alaska) at 3-6°C and pH 6-6.4 (FENCHEL 1975, SULUVAN 1957); Brainard Lake
(3140 m above sea level) and other sites in Colorado (BEARDSLY 1902, CAIRNS & YONGUE
1973b, EDMONDSON 1912, HAMILTON 1943); sphagnous swamp near the village ofPilatka
and other sites in Florida (BAILEY 1851, EHRENBERG 1853); Iowa (SHAWHAN et aI. 1947);
south eastern Louisiana (BAMFORTH 1963); Potomac River, Maryland (CAIRNS 1966b); Sa-
lem, Massachusetts (COLE 1853); at 19-24 °C and pH 5.6-7.0 in Douglas Lake region,
Michigan (CAIRNS & YONGUE 1966, PRATT et al. 1985); common in shore collections from
East Twin Lake, Michigan (KOFOID 1896); wetland sites in Michigan (HENEBRY & CAIRNS
1984, PRATT et al. 1989); mesotrophic lakes and marsh with Typha spp., Michigan
(STEWART et al. 1986); natural waters of the Upper Peninsula of Michigan (WEST 1953);
mud in a lake in New York? (MURRAY 1974); Scioto River, Ohio (LACKEY 1938b); Brush
Lake, Ohio (RIDDLE 1905); common on floating algae in water well exposed to light, in
 Stylonychia 541

algae covered roots in clear, well lighted water, and on sand and algae in clear, fresh,
slowly running water at Mirror Lake, Ohio (STEHLE 1920); rarely in November in a shal-
low area of Crystal Lake, Oklahoma (BRAGG 1960); common in the Conestoga drainage
basin, Pennsylvania (CAIRNS 1965a); Reelfoot Lake, Tennessee (BEVEL 1938); Mountain
Lake region, Virginia (BOVEE 1960); pond in Blacksburg, Virginia (McCORMICK &
CAIRNS 1991, 1992); rarely in Lake Cromwell, Canada (PuYTORAC et al. 1972); Costa
Rica (RUIz 1961); Mexico (L6PEZ-OCHOTERENA & ROURE-CANE 1970, MAoRAZo-GARIBAY
& L6pEZ-OCHOTERENA 1973); Argentina (CARBONELL 1935, SECKT 1924); Brazil (CUNHA
1913, 1918, PROWAZEK 1910); Pantanal floodplains, Brazil (HARDOIM & HECKMAN 1996);
running water near Amazon River, Colombia (BOVEE 1957); Paraguay (DADAY E. 1905;
DADAY J. 1905); moss infusions, Venezuela? (SCORZA & NuNEZ MONTIEL 1954).
Africa: East Africa and Madagascar (SONDHEIM 1929); Cameroon (NJINE 1977).
Australia, New Zealand: Waterloo Swamps, Port Jackson, Australia (WIDTELEGGE
1889); Wellington (MASKELL 1886).
Antarctica: Coast Lake, Victoria Land (THOMPSON 1972).
Records from brackish water and marine habitats largely not substantiated by illustra-
tions: saline lakes in Germany (ZACHARIAS 1888); Baltic Sea (ANDRUSHAITIS 1990, BIER-
NACKA 1962, BOIKovA 1984a, b, 1986, JACOBSON 1928, KUMSARE et al. 1974, LEVANDER
1894a, b, 1901); Gulf of Kola (GASSOVSKY 1916); salt-pit in Cagliari, Italy (PARONA
1883); saline lake in Romania (TUCOLESCO 1962b); Divichinsky Estuary with 3.5 %0 salin-
ity and pH 7.0-8.5, Caspian Sea (AGAMALIEV 1986); saline lakes and brackish water on
Apsheron Peninsula, Azerbaijan (AGAMALIEV 1980); low salinity habitats (4 %0,27 0c) at
Black Sea in Bulgaria (DETCHEVA 1981, 1983d, VALKANOV 1936); Bay of Kertsch, Black
Sea (ANDRUSSOWA 1886); saline pools in central Asia (GURWITSCH 1934); polluted har-
bour on east coast of India (KALAVATI et al. 1995, RAMAN et al. 1993); Barents Sea
(AzOVSKY 1996); attached, or in debris and in bottom samples from the wharf, Atlantic
ocean at Woods Hole, USA (LACKEY 1936, 1938a); saline lakes in Canada (WILBERT
1995); Gulf of Mexico (ALADRo-LUBEL et al. 1988, 1990, BORROR 1962, MAoRAZo-
GARIBAY & L6PEZ-OcHOTERENA 1985, SMITH 1904). The experimental data on the tolera-
tion of increased salinity are rather varying; according to FINLEY (1930) and HAMMER
(1986), species of the S. mytilus complex tolerate a salinity of up to 1.6 %, after Ax & Ax
(1961) 3 %0, and after BICK (1967a, 1968b) only 1.8 %0. On the other hand, records from
marine habitats are available (see above). KuPPER (1963) found S. mytilus in a brook
with 6.6 g 1-1 ct, and ROSTKOWSKA (1961) stated that sodium chloride concentrations of
more than 5 %0 are noxious (Table 33). ALBRECHT (1983, 1984, 1986) classified S. myti-
Ius as oligo- to euryhaline (up to 5 g 1-1 Cn with a freshwater preference. However, sum-
marising the data listed above, species of the S. mytilus complex must be classified as
holo-euryhaline (>3 % salinity) with a freshwater optimum.
Records from terrestrial habitats: grey alluvial soil of a bottom-land in Lower Austria
(FOISSNER 1982, FOISSNER et al. 1985); Belgium (CHARDEZ 1967; Fig. 154g); Bulgaria
(DECHEVA 1973, DETCHEVA 1972b); spruce-forest in Czechoslovakia (ROSA 1957a); Ger-
many (GREEFF 1889, HARTMANN 1956); irrigated (?) agricultural soil in Romania (LEPSI
542 SYSTEMATIC SECTION

1951); garden soil from Stockholm, Sweden (KoFFMAN 1926, 1934); Salix litter and moss
from Scoresby Land, East Greenland (STOUT 1970); USSR (NIKOLJUK & GELTZER 1972);
Delta hardwood bottom-land forest soils in south-eastern Louisiana (BAMFORTH 1967);
soil from shrub in a cedar glade, Tennessee (MARTIN & SHARP 1983); mosses near Mont-
real, Canada (FANTHAM & PORTER 1946); loamy soil with pH 6.4 in South Africa (FAN-
THAM 1929, 1931, FANTHAM & PATERSON 1926, FANTHAM & ROBERTSON 1928, FANTHAM &
TAYLOR 1922, FANTHAM et al. 1927); pasture and seasonally flooded grassland in New
Zealand (STOUT 1981, 1984).
Stylonychia mytilus is not very common in sewage treatment plants such as activated
sludge (ALONSO et al. 1981, BANINA 1990, HARTMANN 1960; KUTIKOWA 1984, the figure is
a redrawing from CURDS 1969, Fig. 151q; LACKEY 1938a, MAnoNI 1988a) and trickling
filters (BANINA 1983, BARKER 1949, CURDS 1969, 1975, LACKEY 1938a). Not recorded
from Imhoff tanks. SCHERB (1968a) counted up to 900 indo mI- 1 in activated sludge with
351 mg I-I KMn04-consumption at influent and 91 % digestion; accordingly, S. mytilus
was more abundant in plants treating dairy wastes than in plants treating municipal sew-
age. LIEBMANN (1936) counted 40 indo ml-I in the Kahmhaut of35 d and 60 d old, stand-
ing municipal sewage. KAUFMANN (1958) found it in the mud of an oxidation pond.
Other habitats: drinking water in Austria (LIEB et al. 1956); wells (SPANDL 1926c,
VEJDOVSKY 1882); filter of drinking water plants in the Netherlands (VERSCHAFFELT 1930);
caves in Germany, Scandinavia, France, and Italy (BOSCOLO 1968, GRIEPENBURG 1934,
1935, LOFFLER 1951, MONIEZ 1889, Tosco 1956; see also review by GITILESON & HOOVER
1969).
Species of the S. my/ilus complex are omnivorous, that is, they feed on bacteria (for
example, Bacillus subtilis, rhodobacteria), cyanobacteria, algae (Chlamydomonas rein-
hardtii, Chlorogonium elongatum, Euglena, Phacus caudata, Scenedesmus acutus,
spores of Spirogyra sp., diatoms), yeast (Saccharomyces cerevisiae), fungi, heterotrophic
flagellates (Chilomonas paramaecium), ciliates (Colpidium sp., Cyclidium sp., Euplotes
octocarinatus, Tetrahymena pyriformis), starch, dried chicken egg yolk, and organic de-
tritus (for example, BENllTS-CLAUS 1974, see Table 32; BERECZKY 1969, BICK 1972a,
BRAND 1923, CHADHA et al. 1978, CURTIS & CURDS 1971, CZAPIK 1975, DEMBOWSKA 1925,
FENCHEL 1975, FRICK 1968, GAUSE 1934a, b, GELLERT & TAMAS 1959a, GRACIA & IGUAL
1987a, GROZDOV et al. 1982, HECKMANN 1995, JONES 1974, KAHL 1932, KAUL & SAPRA
1983, KOLKWITZ 1950, KUHLMANN & HECKMANN 1994, LICHTENBERG 1955a, LIYEPA 1984b,
PFISTER 1995, PFISTER & ARNoT 1995, POSCH & ARNoT 1995, 1996, PRArr & CAIRNS 1985,
PUYTORAC et al. 1976, RIGGENBACH 1922, SHIGENAKA et al. 1973, TUFFRAU et al. 1981,
ZINGHER 1934). Also feeds on eggs of the maw-worm, Ascaris suum (SHARAVINA 1978).
One specimen ingests 2400 diatoms and other algae per day with a total volume of
960000 J.lm3 and 361 000 J.lm3 respectively (ScHONBORN 1981, 1982). Maximum inges-
tion rate per hour about 20 % of cell volume (FENCHEL 1980b).
KAUL & SAPRA (1983) found that S. mytilus does not ingest non-nutrient particles,
such as charcoal, India ink, or carmine particles. Furthermore, they observed that the al-
gae Chlorella sp. and the protists Colpoda sp. and Chilomonas sp. are not accepted as
 Stylonychia 543

Table 32 Influence of food concentration (Chlamydomonas reinhardtii) on abundance, relative growth con-
stant (h-I), and generation time (h) of a species ofthe Stylonychia mytilus complex (from BENUTS-CLAUS 1974)

LlO' 0,5.10' 0, LlO'

algal cells! ml algal cells! ml algal cells/ ml

light dark light dark light dark

Number of Sty{onychia/ml
Oh 5 5 5 5 5
24 h 20 20 20 20 20 20
48 h 35 45 35 25 60 30
72h 0 80 330

Relative growth conSlant r

24 h 0,058 0,065 0,058 0,058 0.058 0.058
48 h 0.041 0,046 0.041 0.034 0,052 0.037
72h 0,000 0,029 0.032 0.033 0.047 0.000
96 h 0,000 0.030 0.044

Generation time T
24 h 12,0 10,1 12.0 12,0 12.0 12.0
48 h 17,1 15.1 17.1 20.7 13.4 18.6
72h 24.0 21,7 20.8 14.7
96 h 23,5 15.9

prey (see, however, previous paragraph). The feeding response towards various other
food materials and the generation time at 23°C in a modified Pringsheim medium are as
follows: algae Chlorogonium, feeding response = good, average number of prey organ-
isms or particles ingested within 5 min = 25, generation time = 10 ± 0.5 h; algae Anacys-
tis: poor, 3, 480 ± 4 h; ciliate Tetrahymena: fair, 10, 20 ± 2 h; bacterium Escherichia
coli: poor, 5, 36 ± 2 h; mixed bacterial flora (in hay infusion): poor, 5, 36 ± 2 h; dried egg
yolk particles: fair, 10, 12 ± 1 h; dried egg albumin particles: fair, 10, 12 ± 1 h. On the
other hand, KAUL et al. (1978a, b) stated that feeding continuously on Tetrahymena for
several generations leads to many morphological and cytological abnormalities and fi-
nally to the death of S. mytilus; the changes ranged from cytoplasmic vacuolisation to
breakdown in division process, abnormal micronuclear number, and reduced RNA syn-
thesis. Morphologically, the changes are expressed as loss of cortical structures and al-
teration of cortical membrane resulting in loss of body form.
The water leaves the membranelles of the adoral zone with a velocity of somewhat
more than 1 mm S-I yielding a transport rate of 1.36 x 106 !lm3 S-I or 4.9 x 10-3 ml h-I, Ex-
periments with the ingestion of 5,7 !lm sized latex beads yielded a maximum clearance of
about 5.5 x 10-3 ml h-I (FENCHEL 1980a, b). Maximum clearance per cell volume per hour
about 8 x 104 ; free space between cilia of undulating membranes about 3.5 !lm (FENCHEL
I 980c).
544 SYSTEMA TIC SECTION

Table 33 The reaction of a species of the Stylonychia mytilus complex adapted to sodium chloride (3 %0
ofNaCI) and of control Stylonychia to various substances (from ROSTKOWSKA 1961)
The reaction of Stylonychia mytilus adapted to t~e sodiu~ chloride (0.305% of NaCl)
and of control Stylonychla of pOlson

I Numl,~r 01 plasmolysed protozoa

alter 10 min.
II Number of plasmolysed protozoa
after 100 min.
Kind of
substance Adapted
protozoa Ii Control
protozoa
IThe statistical
ISignificance of
Adapted
protozoa
I Control
protozoa
jThe statistical
signIficance of
I
concentration
in % Abso- \ AbSO-I the difference Ab50-1 I AbSO-I the difference
lute .~ I lute % after 10 min. lute % lute % alter 100 min.
number lfiumber nurrber number

1__A_<t_et_01_in__ I__0_ _0_'_0_ _0_ dirf~~nce ~ 6.591~'53.7~1 p > 99%

1_r.;_f_~~_)_6_~g_le_I_8- ~ 13.18 P = 95% ~ 42.70 ~ 65.91

----,-------.------------I
7.01 p >99% I

Chinosol
0.02
0
I 0 i
I
0 0 d'fr no
1 erence
20 11.1 165 100
1
P > 99%

1_ _
Phenol 104 58I 5:1
o_.2_ _ _ _ _ _ 1_ _
34
1__ P >99% 191 1100
.__ 15il
1_ _ _ _ _ _ _
100 d'ffDO
1 erence

I di1f~;'n"
Mercuric
chloride
0.0002
o I 0 I 0 I 0 I 4" 136084 161 100 I p > 9"%
GRAY (1951) found that S. mytilus always exhibits maximum abundances in a chalk
stream about 4-7 d before and after full moon, with slight evidence for a drop at full
moon itself. He assumed that, in the particular stream examined, the Stylonychia were
following a lunar periodicity of diatoms on which they fed. Maximum phases of the cili-
ate might therefore follow maximum phases of the diatoms, succeeded by periods of re-
productive activity. BERECZKY (1977a) found maximum abundances in the Danube River
at 4 am and at 8 pm.
Species of the Stylonychia mytilus complex can be easily cultured. AMMERMANN et al.
(1974) used neutral Pringsheim solution and fed it with Chlorogonium grown under ster-
ile conditions. WIRNSBERGER et al. (1986) used tap water as culture medium and yeast,
Chlorogonium elongatum, and a species of the Tetrahymena pyriformis complex as food.
MACHEMER (1965b) cultured it in 10 % soil-peat-medium and 0.06-0.1 % yolk as food
supply.
TOTWEN-NoWAKOWSKA (1981) observed 1.6 ± 0.4 cell divisions per day in Pringsheim
medium at about 20°C and a species of the Tetrahymena pyriformis complex as food.
For dorsal doublets she found 1.5 ± 0.5 and for lateral doublets 0.7 ± OJ divisions per
day. CHADHA et al. (1978) and KAUL & SAPRA (1989) estimated a generation period of
about 10.5 ± 0.5 h (n = 100) at 23°C as well as at 30 °C and with Chlorogonium elonga-
tum as food. Further data on generation time (see also Table 32): 3 divisions in 24 h at 20
± 2 °C (GRoZDOV et al. 1982); 13.5 hat 23°C with Chlamydomonas reinhardtii as food
(BENUTS-CLAUS 1974). MACHEMER (l965a) stated that below 10°C nearly no more divi-
 Stylonychia 545

Table 34 Autecological data of species of the Stylonychia mytilus complex. References: column 1, from
NOLAND (1925; investigated in all 80 samples from margins oflakes, a pond, an artificial lagoon, permanent
pools, temporary pools, a river, creeks, swamp ditches, and a spring in Wisconsin, USA); column 2, from
CAIRNS (1964; in all 202 analyses from various rivers and streams, mostly in the USA); column 3, from CAIRNS
& YONGUE (1973a; data from "Station 1 and year 1970" from the Cape Fear River near Fayetteville, North
Carolina, USA, sample size not indicated); column 4, from BICK & KUNZE (1971; summary of literature data);
column 5, from DETCHEVA (1972a, 1975a-c, 1976b, 1978, 1979a, c, 1982b, 1983a, b, c; many analyses from
Bulgarian running waters); column 6, from MADONI & GHE1TI (198Ia; Torrente Stirone in Northern Italy; sam-
ple size not indicated); column 7, from FOISSNER et al. (1982a; 9 analyses from small, stagnant alpine water
bodies in Austria)

Parameterl Reference
2 3 4 5 6 7
Frequency (%) 12.5 31 1.0-9.8
Temperature COe) 4.0-23 4-40 24.5-26 2-25 0.6-31 17.5-21 2.5-20
pH 7.3-8.7 6.0-9.5 6.7-7.3 4.0-8.4 6.3-8.5 8.0-8.5 4.9-5.6
O2 (mg,·I) 2.6-15.3 3-15 6.6-8.7 0-22.5 3.0-14.3 9.5-11.3 6.7-11.3
O2 (% saturation) 23-169 30-103 86-148
BODs (mg ,.1) <0.5-202 1.4-2.4 2-25 1.3-49 7-13 4 4-76s
NH/-N (mg ,.1) <0.001-203 0.005-0.123 0-20 0.025-8.23 0.25-1.43 0-6.2
NOi-N (mg ,.1) <0.001-1.5 0-0.3 0.006-0.1 0-3
NOi-N (mg ,.1) 0.001-7 0.37-0.65 0-11.7 0.1-0.8 0-0.45
POl--p (mg ,-I) 0.033-3.3 0.09-0.135 0.13-0.69 0-0.09

I The figure "0" denotes only that the parameter in question could not be detected with the analytical method
used_
2 In CAIRNS (1964) this parameter is designated as BOD only.
3 NH3-N.
4 COD.

s KMn04-consumption.

sions could be observed. Amicronucleate cells have an increased division rate (Lu & NG
1989). For effects of alcohol on the life cycle, see WOODRUFF (1908).
Respiration rate 2.33 nl O2 cell-I hoI (condition unspecified, cell volume 900 000 ~m3,
corresponding to a biomass of 900 mg per 106 individuals, which is rather high; CHORIC &
SHUBERNETSKY 1978, FENCHEL & FINLAY 1983, KHLEBOVICH 1987, KLEKOWSKI 1981).
Data on biomass of 106 individuals rather variable: 70 mg (FOISSNER 1987a; terres-
trial population of FOISSNER 1982), 199 mg (MICIllELs 1974),277 mg (DILLON & HOBBS
1973),350 mg (FOISSNER et al. 1991; calculated from average size), 390 mg (NESTERENKO
& KOVALCHUK 1991), 402 mg (SCH<JNBORN 1981, 1982), 900 mg (FENCHEL & FINLAY
1983). As a consequence, biomass should be calculated separately for each popUlation in
question.
Some data on community production and some autecological data are shown in Ta-
bles 28, 34, 35 and Figure 155. Further autecological data: <6-2500 mg I-I CI-, I to
20 mg I-I CO2 free, <0.01-1 mg I-I Fe2+, <3-450 mg I-I Ca2+, <3-200 mg I-I Mg2+ (CAIRNS
1964); 4-28.5 DC, pH 5.8-6.9, 2-10.8 mg 1- 1 0 2 (MATIS 1975); 7-38 DC, pH 5.0-7.5 (MA-
546 SYSTEMATIC SECTION

Table 35 AutecologicaJ data of species of the Stylonychia mytilus complex (columns 1-4) and Stylonychia
putrina (columns 5-7). References: column 1, from BERECKY (I977a; ? analyses from the betamesosaprobic
Danube River in Hungary); column 2, from RU11NEN (1972) and CAIRNS & DICKSON (1972; with low abun-
dance in 4 stations in the South River, Virginia, USA, n = ?); column 3, from MuwLOwrrsCH (1989; 11-13
analyses from salt loaded running waters in Germany); column 4, from FOISSNER et a1. (1991; running waters in
Austria, n = number of analyses); column 5, from BICK (1960, 1968a, 1972a) and BICK & KUNZE (1971; sum-
mary of literature data, supplemented with data from BICK & BERTRAM 1973, BICK & DREWS 1973, GREISER
1974, PRIMC 1983, and WILBERT 1977); column 6, from DETCHEVA (1983c; several analyses from a1phameso-
saprobic areas of the Maritza River, Bulgaria); column 7, from PATRICK et a1. (1967; few analyses from the Sa-
vannah River, USA) and from RU11NEN (1972) and CAIRNS & DICKSON (1972; ? analyses from 4 stations in the
South River, Virginia, USA)

Parameterl Reference
2 3 4/n 5 6 7
Frequency 15
Temperature (0C) 2-20 4-20 0.2-12120 2-30 18-25 23-28
pH 6.7-8.1 7.6-8.2 7.4-8.0 7.1-8.7/20 4.7-9.4 7.2-7.4 7.5-9.0
Oz (mgl· l ) 7-14.4 4.5-9.3 6-13/20 0-12.6 5.5-7.7 1-7
Oz (% saturation) 55-105/20 67-80
BODs (mg I-I) 2.4-5.1z 1.1-2.3 1->7/19 0.5-9.1
NH':-N (mg ).1) 0.7-1.1 0 0.07-0.5 0-1.3/20 0-40 0.05-0.21 0-0.05)
NOz·-N (mg I-I) 0.017-0.07 0.002-1.3 0.03-0.8 0-0.02116 0.03-0.46 0-1
NO)·-N (mg 1.1) 0.47-1.6 0.004-0.39 0.3-6 0.5-2.2/19 0.23-0.34 0.03-0.4
PO/·_p (mg I-I) 0.115-2.2 0.02-0.27/13 0.016-2.2

I The figure "0" denotes only that the parameter in question could not be detected with the analytical method
used.
z "Oz-consumption".
) NH)-N.

TIS & STRAKOVA-STRIESKOvA 1991); pH 6.8-8.0 (DINGFELDER 1962); 13.5-89 mg 1.1 CO2
free, 40-265 mg 1.1 CI', 573-1,514 )lS em· I specific conductivity (MnwLoWITSCH 1989; salt

polluted running waters in Germany); 0-28 mg 1.1 CO2 free (BERECZKY 1977a; Danube
River in Hungary); 21.5-28 °C, >1-10 mg I-I CO2 free >5-9 mg 1.1 O2, 0.01-<0.9 mg 1.1
Fe2+ (PATRICK et aI. 1967); 0.36-0.74 mg I-I Fe2+, 0.05-0.08 mg 1.1 Mo2+, 32-34 mg 1.1
HC03', 7-18 mg 1.1 SOl', 15-17.5 mg 1.1 CI' (CAIRNS & YONGUE 1973a; Savannah River,
USA); 70-218 mg 1.1 loss of ignition, 93~310 mg 1.1 total solids, 124-189 mg I') total
hardness, 9-103 mg I') SOl', 1.8-34 mg 1.1 CI', 0.09-0.4 mg I') Fe2+, 0.008-0.063 mg I')
Co, 0.022-0.067 mg I') Zn, 0.007-0.036 mg 1-) Cr, 0.008-0.018 mg 1-) Pb (RUTHVEN 1972
and CAIRNS & DICKSON 1972; South River, Virginia). FOISSNER et al. (1982a) found Styla-
nychia mytilus once with high abundance in an alpine water body at lOx 106 bacteria ml-)
(direct counting), 9.7°C, pH 5.3, 10.8 mg 1.1 O2 (123 % saturation), 2.6 mg I') COlfree,
2 °dH total hardness, 76 mg I-I KMn04-consumption, 6 mg 1-) NH/-N, 0.09 mg I')
POl'-P, 1.2 mg 1.1 Fe2+, 0 mg 1.1 HzS. According to STOSSEL (1979), species of the S. myti-
Ius complex preferred a DOC range of2.6-7.2 mg 1-) (total range 0.5-7.2 mg 1.1). BICK &
KUNZE (1971), FOISSNER et al. (l982a), and FOISSNER et al. (1991) provided following bac-
 Stylonychia 547

Fig. 155 Distribution expressed as

percentage of n records of species of
the Stylonychia mytilus complex in
the spectra of some physical and
chemical factors (from HEuss 1976).
2 1 = temperature, 2 = pH, 3 = spe-
cific conductivity, 4 = O2, 5 = Or
saturation, 6 = BODs, 7 = KMn04-
consumption, 8 = Cl-, 9 = POl-, 10
=NH/.

terial counts (cells ml- I ): 6000

to 500 0000 (plate method),
500 000-10 000 000 (direct
counting with phase-contrast
microscopy), 1000-1300000
6 (plate method).
Eurythermic (Table 34,
35; BICK & BERTRAM 1973,
JAROSHENKO & CHORIK 1969,
MATIS 1975, MONcH 1970,
WILBERT 1977). However,
PRIMc (1983) found S. myti-
Ius in laboratory experiments
only at 5°C, indicating insuf-
ficient culture conditions.
WANG (1928) and LICHfEN-
BERG (1955b) mentioned
18-20°C as favourable tem-
perature. DINGFELDER (1962)
even found S. mytilus in road
pools at 47°C! KAUL & SA-
PRA (1989) reported that 30°C is the maximum temperature to which S. mytilus can adapt
without visible consequence to their cortical anatomy or generation time. However, they
found that the macronuc1ear area is significantly higher in cells grown at 30°C (1075.7 ±
43 J.lm2) than in cells grown at 23°C (781.4 ± 19 J.lm2). Heat shock induced ultrastruc-
tural alterations are described by KAUL & SAPRA (1991).
Maximum survival time in an atmosphere of pure carbon dioxide <15 min and at
very low oxygen tension about 1 h (NIKITINSKY & MUDREZOWA-WySS 1930, STOUT 1956).
FERNANDEZ-LEBORANS et al. (1985) found 1.2-1.8 J.lm sized lead deposits in speci-
mens which lived in samples treated with 500 J.lg I-I lead acetate. According to BOYKOVA
(1982), S. mytilus tolerated 20 mg 1-1 Pb (2.6 divisions per day), however, a conspicuous
physiological polymorphism occurred; after a few days with 10 mg 1-1 Zn the growth rate
548 SYSTEMA TIC SECTION

was as high as in the control (BOIKOVA 1990). Stylonychia mytilus did not tolerate more
than 100 )lg 1"1 zinc chloride, at least when 50 )lg 1"1 lead acetate was present (FERNANDEZ-
LEBORANS & ANTONIO-GARCIA 1986). Minimal inhibitory concentration and minimal lethal
concentration of tinidazole 1 mg 1"1 (incubation time 2 d) and 3 mg 1"1 (incubation time
3 d) respectively (ALONSO 1979). Presence of 0.001-0.01 % NiS04-ions in culture me-
dium did not change spiralling direction as it did in some other species (SERAYIN 1970).
MACHEMER (1966a, b) demonstrated a very weak but significant avoidance of light
(1000 Ix) over 1 h and a weak avoidance of rough over 80 min. For further ethological
data, see ALVERDES (1937, 1938), BRAMSTEDT (1935), RICCI (1994).
KITCHING (1957b) studied the effect of high hydrostatic pressure. At 28700 kPa, and
more especially at 35900 kPa, the membrane lIes and cirri, previously motionless, began
to flicker. At 43000-50200 kPa all cirri and membranelles beat wildly and the body
turned in circles, but towards the end of the period at 50200 kPa movement of cirri and
body almost ceased. There was instant recovery of movement when the pressure was re-
leased, and the animals swam with an irregular rolling or circling motion. At 57400 kPa
there was some disintegration of the posterior end of the organism, with loss of cirri, but
considerable measure of normal activity was recovered by 3 h after release of pressure.
For the effect of ultraviolet radiation, see TCHAKHOTINE (1936).
Species of the S. mytilus complex are usually classified as alphamesosaprobic indica-
tors of water quality (FOISSNER 1979b, 1988a, KOLKWITZ 1950, KOLKWITZ & MARSSON
1902, 1909, MARSSON 1903, SLADECEK 1964, 1969, 1973, SLADECEK et al. 1981, SRAMEK-
HuSEK 1958, STJERNA-POOTH 1981, WEGL 1983). PRIMc (1984) observed the highest abun-
dances under alphamesosaprobic to polysaprobic conditions. Voss (1978) obviously erro-
neously designated it as "Faulwasser-Bewohner". MAUCH (1976), MAUCH et al. (1985; Si
= 2.5), MORAVcovA (1977; b = 4, a = 6, 1= 3, SI = 2.6), and SRAMEK-HuSEK (1956a) clas-
sified S. mytilus as beta- to alphamesosaprobic, BUCK (1971) even as betamesosaprobic
with a corrected saprobic index of 1.96 ± 0.36. According to FRIEDRICH (1990; see also
BERGER et al. 1997 and DIN 38410 Teil2) the S. mytilus complex is classified as follows:
o-b = 2, b = 8, b-a = 6, a = 4, 1=4, SI = 2.3. FOISSNER et al. (1991) found it mainly in
beta- to alphamesosaprobic or alphamesosaprobic running waters with a saprobic index
of 1.9-3.2. The highest abundance was observed in an alphamesosaprobic area (6.8 mg I-I
Oz, 65 % Oz-saturation, 30 mg I-I COD) of a river in Upper Austria heavily polluted by
paper-mill effluents. Considering the autecological data given in Tables 34, 35 and the
faunistic records, the. alphamesosaprobic classification as suggested by SLADECEK et aI.
(1981) and FOISSNER et al. (1991) seems justified (a; b = 1, a = 9, 1= 5, SI = 2.9). Possi-
bly, S. mytilus and S. lemnae have somewhat different saprobic preferences, which has to
be verified by autecological and saprobiological investigations on reliably determined
material. Stylonychia mytilus does not tolerate hydrogen-sulphide (WETZEL 1928b).

Sup po sed s y non y m s: Stylonychia Yirgula FROMENTEL, 1876, Etudes micro-

zoaires, p 273, Planche XIV, fig. 3. Remarks: "Stynolychia Yirgula" in FROMENTEL (1876,
P 359) is an incorrect subsequent spelling of Stylonychia. Especially the body shape is
 Stylonychia 549

reminiscent of species of the S. mytilus complex; possibly a slightly damaged specimen.

In life about 115 !-lm long. France? The drawing in DUMAS (1929, on p 83 as "Stylonichia
virgula", in the legend to Planche XXIX, fig. 8 designated as Stylonychia auriformis) is
obviously a redrawing from the original description (Fig. 237d).

Stylonychia kahli DRAGESCO, 1966, Protistologica, 2: 92 (Fig. 152b). Remarks: Body

shape, size (about 200 !-lm long), and arrangement of transverse cirri are reminiscent of
species of the S. mytilus complex. 66-71 adoral membraneUes, 38-43 right and about 28
left marginal cirri. 2-4 micronuclei. Six dorsal kineties; two long caudal cirri, which is
rather surely an anomaly. Locus classicus is a limnetic habitat near Thonon-Ies-Bains,
France. BORROR (l972a) synonymised it with S. pusilla STERK!, 1878, which is here con-
sidered as supposed synonym of S. notophora.

Stylonychia mytilus (MULLER, 1773) EHRENBERG, 1830 (Fig. 148n, 0,

150a-f, 151j, 153d, Table 31)

1773 Trichoda mytilus MoLLER, Vermium Terrestrium et F1uviatilium, p 87.

1786 Kerona mytilus - MOLLER, Animalcu1a infusoria, p 242 (Fig. 150d-1).
1830 Sty/onychia myti/us - EHRENBERG, Abh. preuss. Akad. Wiss., year 1830: 43.
1965 Sty/onychia mytilus Varietat 2 - AMMERMANN, Arch. Protistenk., 108: III (Fig. IS0c).
1983 Sty/onychia mytilus, EHRENBERG 1838 - AMMERMANN & SCHLEGEL, 1. Protozool., 30: 292 (Fig. 148n, 0,
151j).
1983 Sty/onychia mytilus EHRENBERG, 1838 - STEINBROCK & SCHLEGEL, J. Protozool., 30: 294.
1985 Sty/onychia mytilus - AMMERMANN, Memoire Soc. tose. Sci. nat., 92: 17, Fig. I.
1986 Sty/onychia mytilus - WIRNSBERGER, FOISSNER & ADAM, Arch. Protistenk., 132: 169 (Fig. 150a, b, 153d;
1 slide of neotype specimens is deposited in the OberOsterreichische Landesmuseum in Linz, Upper
Austria).

Nom e n c I a t u rea n d t a x 0 nom y: See also relevant chapter in the S. mytilus

complex. Neither AMMERMANN & SCHLEGEL (1983) and WIRNSBERGER et al. (1986) nor
anyone else designated a clone of a true S. mytilus as neotype. Furthermore, they did not
fix a type location. To eliminate these shortcomings the following statements are made to
satisfy the qualifying conditions (1)-{6) of Article 75 (d) OfICZN (1985): (I) for the char-
acterisation of the species, see bibliographic references in the list of synonyms; (2) the
specimen from clone CI figured in WIRNSBERGER et al. (1986; Fig. 2, 4; here Fig. 150a, b)
is designated as neotype specimen; (3) a careful study ofthe post 1965 literature revealed
that no type was designated; (4) the specimen designated is consistent with that known
from the original description and many redescriptions; (5) the neotype came from a local-
ity which is not too far away from MOLLER's (1773) locality (Denmark); (6) the slide con-
taining the neotype specimen is deposited in a scientific institution. General description
see also key and S. mytilus complex. Stylonychia mytilus and S. lemnae are separated
since 45 x 106 years (SCHLEGEL & STElNBRUCK 1986).
550 SYSTEMATIC SECTION

M 0 r ph 0 log y and b i 0 log y: Length after AMMERMANN & SCIilllGEL (1983) in

life 300 ± 22 !lm after 1 day starvation and about 370 !lm before division. Left margin
with a conspicuous bulge at about level of frontal cirri. This outline is more pronounced
in starved than in well feed animals. However, the results by WIRNSBERGER et al. (1986)
proved that neither shape nor size provide criteria for separation from S. /emnae. Both
macronuc1ear nodules together in Gl contain 1004 ± 33 pg (n = 10) DNA. 3--6 (mean =
3.7; n = 25) micronuclei, each containing in Gl about 24.8 ± 0.5 pg DNA (C-value:
12.4 pg; AMMERMANN & SCHLEGEL 1983, RAIKov 1989). Intrac10nal conjugation common.
For a detailed biochemical characterisation comprising isoenzyme patterns and macronu-
clear and micronuc1ear DNA banding patterns after restriction enzyme digestion, see
STEINBROCK & SCIilllGEL (1983). Anterior portion of dorsal kineties 1-4 more curved than
in Stylonychia /emnae (cp Fig. 149b with 150b).

o c cur r e n c e and e colo g y: Common in freshwater habitats, prefers lentic wa-

ters. Type location: lake near Reutlingen-Gonningen, Germany (clone "C I" of WIRNS-
BERGER et al. 1986). AMMERMANN (1965a) and AMMERMANN & SCHLEGEL (1983) found it in
various limnetic habitats around Tuebingen and Cuxhaven, in Muenster, in Lake Feder-
see, in 4 lakes in Upper Swabia (Germany), in a lake near the Lake Neusiedlersee (Aus-
tria), in a lake south of Gdansk (Poland), in moss from West Africa, in fish ponds near
Harbin (China), and in Cashmere (India). AMMERMANN et al. (1989) isolated Sty/onychia
myti/us from a pond near Ithaca, USA. Tolerates more than 0.3 % NaCI and occurs also
in brackish water (AMMERMANN 1965a).

Slylonychia lemnae AMMERMANN & SCHLEGEL, 1983 (Fig. 3a, 4,5, lla, b,
26a-j, 148m, p, 149a-d, 153a-c, e-j, Tables 18,31)

1965 Sty/onychia mytilus Varietat 1- AMMERMANN, Arch. Protistenk., 108: III (Fig. 149d).
1983 Sty/onychia /emnae AMMERMANN & SCHLEGEL, 1. Protozool., 30: 291 (Fig. 148m, p, 149c; some slides of
the type material are deposited in the Smithsonian Institute, Washington D. C., USA).
1983 Sty/onychia /emnae STEINBROCK & SCHLEGEL, J. Protozool., 30: 294.
1985 Sty/onychia /emnae - AMMERMANN, Memoire Soc. tosc. Sci. nat, Serie B, 92: 17, Fig. 1.
1986 Sty/onychia /emnae AMMERMANN & SCill.EGEL, 1983 - WIRNSBERGER, FOISSNER & ADAM, Arch. Protis-
tenk., 132: 167 (Fig. 149a, b, 153a-c, e-j).

Nom e n c I a t u rea n d t a x 0 nom y: See key and S. myti/us complex for these
topics and a general description.

M 0 r ph 0 log y and b i 0 log y: Length in life 230 ± 12 !lm after starvation for 1
day, about 120 !lm minimal length after 3 days of starvation, and about 270 !lm before di-
vision. According to original description body margins almost in parallel (Fig. 149c, d),
the micrographs, however, show that the outlines of S. /emnae and S. myti/us are identical
(cp. Fig. 1480, p). Both macronuc1ear nodules together in G 1 contain 788 ± 28 pg DNA
 Stylonychia 551

(n = 10). 2-5 (mean = 3.4; n = 25) micronuclei, each containing in G 1 about 18.4 pg
DNA (C-value: 9.2 pg; mean of20 clones and 10 nuclei of each clone measured). Devia-
tions of micronuclear DNA content between clones about ± 15 % from mean (18.4 pg;
RAIKov 1989, STEINBRUCK et al. 1981). AMMERMANN et al. (1989) isolated clones with mi-
cronuclei without function, with small micronuclei, or with no micronuclei at all. Intra-
clonal conjugation rare and in most cases temporary (AMMERMANN 1982). Anterior por-
tion of dorsal kineties 1-4 less curved than in S. mytilus (cp. Fig. 149b with 150b).
A detailed biochemical characterisation comprising isoenzyme patterns and macro-
nuclear and micronuclear DNA banding patterns after restriction enzyme digestion is pro-
vided by STEINBRUCK & SCHLEGEL (1983).
Sty/onychia /emnae is sometimes infected by the host-specific suctorian ectoparasite,
Podophrya grelli DIECKMANN, 1985. For further details see the parasitism chapter in the
general section (Fig. 26a-j) and MATIHES (1988).
Sty/onychia /emnae is intensively used in experimental studies, especially in areas of
genetics, biochemistry, and physiology: AMMERMANN (1964, 1968, 1971b, 1987a, b, 1988,
1990, 1993), AMMERMANN et al. (1974), ASCENZIONI & LIPPS (1986), BALLANYI & DElTMER
(1984), BAROIN-TOURANCHEAU et al. (1992, 1995), BESSLER & LIPPS (1976), BIERBAUM et al.
(1991), CALLEN et al. (1994), CONZELMANN & lIELFrENBEIN (1987), DEITMER (1981b, 1982,
1983b, 1984, 1986a, b), DEITMER et al. (1983, 1984), EDER et al. (1993), ELSEVIER et al.
(1978), FLEURY et al. (1995), GAUDE (1981), GAUNITZ et al. (1992), GRIMES et al. (1981),
HARPER & JAHN (1989), HELFfENBEIN (1985), HELFfENBEIN & MOLLER (1988), lIELFrENBEIN
et al. (1989), HERRICK (1992), IVENS & DEITMER (1986), KRAUT et al. (1986), LIPPS (1975,
1977, 1980), LIPPS & BESSLER (1977), LIPPS & EDER (1996), LIPPS & ERHARDT (1981), LIPPS
& lIANTKE (1975), LIPPS & MORRIS (1977), LIPPS & STEINBRUCK (1978), LIPPS et al. (1978),
MACHEMER & SUGINO (1989), MAERCKER & LIPPS (1993, 1994a, b), MAERCKER et al.
(1997a, b), MEYERS & lIELFrENBEIN (1988), NOCK (1981), PEVER & DEITMER (1980), PEVER
& MACHEMER (1977, 1978b, 1982b, 1983), SAPRA et al. (1985, 1994), SCHLEGEL (1985),
SCHLEGEL & STEINBRUCK (1986), SCHLEGEL et al. (1989, 1990), SCHMIDT et al. (1991), STEIN-
BROCK (1983, 1990), STEINHILBER & LIPPS (1986), STOLL et al. (1991, 1993), SUGINO & MA-
CHEMER (1990), WEFES & LIPPS (1990), WEGNER et al. (1989), WEN et al. (1995), WITI'E &
KNEER (1990), WITI'E et al. (1995). Some other papers are mentioned in the last paragraph
of the relevant chapter of the S. mytilus complex.

o c cur r e n c e and e colo g y: Common in freshwater habitats, prefers lentic wa-

ters. Locus classicus is pond IV of the "Spitzberg forest" near Tuebingen, Germany. AM-
MERMANN (1965a), AMMERMANN & SCHLEGEL (1983), and AMMER.MANN et al. (1989) found
S. lemnae in 10 different water bodies in Germany, namely around Tuebingen, in
Rohrsee, in the Wutachschlucht, Black Forest, in Erbach near UIm, oxbow of the Danube
River, in Taubergiefien, oxbow of the Rhine River, in Muenster, in 6 different waters
around Cuxhaven; furthermore, they found it in Bornholm (Denmark), in China, and in
Greenbelt Lake, College Park, Maryland and near Ithaca, USA. AMMERMANN et al. (1989)
investigated 12 North-American ponds, 7 of them contained species of the S. mytilus
552 SYSTEMATIC SECTION

complex. Stylonychia myti-

Ius was found only in 1
pond, indicating that S.
lemnae is possibly more
common in North America
than S. mytilus. Attempts
by AMMERMANN et al.
(1989) to find species of
the S. mytilus complex in
other regions of the USA
failed. Stylonychia lemnae
tolerates not more than
0.3 % NaCl (AMMERMANN
1965a), suggesting that the
marine and brackish re-
c cords of a species of the S.
mytilus complex deal with
Fig. 156 Sty/onychia nodulinucleata (from SID & 11 1993). The origi-
nal description of this species is in Chinese. The meagre original Eng-
S. mytilus. TAKAHASID &
lish legend runs as follows: "Morphological comparison of three spe- SUHAMA (1991) found S.
cies of Stylonychia with regard to the ventral cirral patterns, adoral lemnae in Japan in soil
zone of membranelles and undulating membrane (drawn according to from paddy fields.
the protargol preparations)". Unfortunately, SID & 11 (1993) omitted to
Length of cell cycle
translate the species names, obviously assuming that everybody can
read Chinese. I suppose that the left illustration shows S. nodulinu- about 12 h (23 ± 2°C,
cleata. Broken lines should connect frontal-ventral-transverse cirri Pringsheim medium, Chlo-
originating from same streak, which is, however, incorrectly shown rogonium elongatum; AM-
(see Fig. 152f, 153j for correct co-ordination). Explanation of original MERMANN & MUENZ 1982).
labels: C = caudal cirri, FI-8 = frontal cirri (FI-l)' buccal cirrus (F.), and
IRATO et al. (1991) exposed
frontoventral cirri (F4-7), LMC = left marginal cirral row, TCI_s = trans-
verse cirri, uM = undulating membranes, VI-S = postoral (VI-l ) and S. lemnae to 5 and 2.5 JlM
pretransverse (V4.s) ventral cirri. Cd as CdCh.H20. The first
dose killed cells. The sec-
ond was well tolerated and
after one day cells accumulated 185 Jlg g-) Cd of dry mass; only 25 % of the metal con-
centration was in the cytosol. Chromatographic analysis did not reveal any chelating
molecule (see Table 18 for data on metal accumulation). Treatment with the DNA cross-
linking agent Cis. dichlorodiamine Platinum (cis platin) at a concentration of 150 Jlg mI-)
for 30 min resulted in amicronuclearity of cells without impairing macronuclear activity
(SAPRA et al. 1989).

Stylonychia nodulinucleata SRI & LI, 1993 (Fig. 156)

1993 Sty/onychia nodulinucleata SID & LI, Zoo!. Res., 14: II (see taxonomy).
 Sty/onychia 553

Nom e n c I at u rea n d t a x 0 nom y: This name was fIrst mentioned in FENG et al.
(1990, p 95), however, very likely without description (I did not read the work, because it
is in Chinese). Unfortunately, the original description is also in Chinese. According to the
summary, S. nodulinucleata matches S. mytilus exactly except for the macronucleus,
which is moniliform (noduliform) like in Coniculostomum. However, Coniculostomum
has two or more right marginal rows. Redescription (please in English) necessary.

M 0 r ph 0 log y: Size (in life? 270-310 x 95-110 flm), shape, and infraciliature as in
S. mytilus (see there). LI & Sm (1993) reported on amicronucleate specimens.

o c cur r e n c e: Locus classicus is Morshan Town, Heilongjiang Province, China,

where Sm & LI (1993) discovered Stylonychia nodulinucleata in marsh water. W. FOIss-
NER (pers. comm.) found it in a floodplain in Australia.

Stylonychia notophora STOKES, 1885 (Fig. 152p, 154e, 159d-f, 166c, d)

1885 Stylonychia notophora STOKES, Ann. Mag. nat. Hist., 15: 446 (Fig. 15ge).
1888 Stylonychia notophora, STOKES - STOKES, J. Trenton nat. Hist. Soc., 1: 299.
1914 Stylonychia notrophora STOKES - SMrrn, Kans. Univ. Sci. Bull., 9: 166 (Fig. 154e; incorrect subsequent
spelling).
1932 Stylonychia notophora STOKES, 1885 - KAlIL, Tierwelt Dtl., 25: 620.
1935 Stylonychia notophora STOKES 1885 - WANG & NIE, Sinensia, Shanghai, 6: 505 (Fig. 159t).
1945 Stylonychia notophora - SRAMEK-HuSEK, Veda prir., 23: 302 (Fig. 159d).
1965 Stylonichia notophora STOKES, 1885 - L6PEZ-OcHOTERENA, Revta Soc. mex. Hist. nat., 24: 161 (Fig.
152p; incorrect subsequent spelling of Stylonychia).
1971 Stylonychia putrina STOKES (?) - DRAGESCO & NJINE, AnnIs Fac. Sci. Univ. fed. Cameroun, 7-8: 130; in
part: "petite varlete", Fig. 28 (Fig. 166d); not "grande varlete", Fig. 27.
1986 Sty/onychia putrina STOKES, 1885 - DRAGESCO & DRAGESCo-KERNEIS, Faune tropicale, 26: 487; in part:
Planche 145 B (Fig. 166c); not Planche 145 A, C-F.

Nom e n c I at u rea n d t a x 0 nom y: KAHL (1932) classifIed Stylonychia as sub-

genus of Oxytricha; the correct name in his revision is thus Oxytricha (Stylonychia) noto-
phora. According to BORROR (1972a) and WIRNSBERGER et al. (1985a), S. notophora is a
synonym of S. pustulata. In my opinion, however, S. notophora is a valid species, which
differs from S. pustulata mainly in the arrangement of the transverse cirri. In S. noto-
phora they are arranged in two groups, similarly to in S. bifaria and S. mytilus (cp. Fig.
149a, 166c, 157a). Conversely, in S. pustu/ata they form a hook (Fig. 161c). Sty/onychia
bifaria has only inconspicuous caudal cirri, whereas they are distinctly elongated in S.
notophora, similarly to in S. mytilus. It differs from the species of the S. mytilus complex
mainly in body shape and size, the narrowly spaced caudal cirri, and the arrangement of
the postoral ventral cirri (Fig. 159d-f, 166d). Detailed redescription necessary for fmal
elucidation of validity. The populations by CONN (1905; Fig. 229b), EDMONDSON (1906;
Fig. 226h), and WANG (1925; Fig. 242c) are insufficiently redescribed. The populations
554 SYSTEMATIC SECTION

described by BARY (1950; Fig. 161y) and SAPRA & DASS (1970a; Fig. 162i) are classified
as S. pustulata.

M 0 r ph 0 log y and b i 0 log y: In life about 120-140 x 50 11m, after BUCHAR

(1957) only 90 11m long, after MORAvcovA (1962) 100-115 x 34-50 11m. Elliptical or
with more or less parallel margins. Posterior end rounded and sometimes slightly emargi-
nate on right-hand side, anterior end obliquely truncated at left-hand side.
About 33-36 adoral membranelles. Postoral ventral cirrus V/4 arranged behind level
of cirrus IV12 (Fig. 166c, d), which is a conspicuous difference to species of Stylonychia
mytilus complex (for example, Fig. 149a). According to original description, the two left
transverse cirri do not protrude beyond posterior end, whereas the three right ones project
slightly to distinctly. About 12-15 left and 18-22 right marginal cirri. Caudal cirri about
25 % of body length.

o c cur r e n c e and e col 0 g y: Throughout the year in lotic and lentic waters;
much more seldom than Stylonychia pustulata and species of the S. mytilus complex.
STOKES (1885b) discovered S. notophora in lentic water with dead leaves and various
aquatic plants. STOKES did not give the specific locality where he found it; however, he
lived and worked in Trenton, New Jersey, USA, and it is assumed that the locus classicus
is near there.
Further records substantiated by illustrations: polysaprobic running water in Czecho-
slovakia (SRAMEK-HUSEK 1945, 1953); in old standing cultures from ponds and pools with
dead leaves and filamentous algae from Nanking, China (WANG & NIE 1935); Kansas,
USA (SMITH 1914); mesosaprobic habitats in Chapultepec, Mexico (LOPEZ-OCHOTERENA
1965, LOPEZ-OCHOTERENA & ROURE-CANE 1970); ponds near Yaounde, Cameroon (DRA-
GESCO & NJINE 1971, NJINE 1977).
Records not substantiated by illustrations: pelagial of clear lakes in Styria, Austria
(KIwNER 1988); limnocrene Slatna Panega and rivers in Bulgaria (DETCHEVA 1981, DET-
SCHEWA 1972, RUSSEV 1988, RUSSEV et a1. 1976, 1984); pond, saprobic brook near
Prague, and alphamesosaprobic area of the Moldavia River, Czechoslovakia (BUCHAR
1957, HASSDENTEUFELovA-MoRAvcovA 1955, MORAvcovA 1962); on submerged macro-
phytes in a mesotrophic to eutrophic lake (Heiliges Meer) in Germany (MOCKE 1979);
limnetic habitats near Ferrara and paddy field with water from the Po River, Italy (CAN-
ElLA 1954, MAnoNI 1987, 1988b; further record from Italy: GRISPINI 1938); Turiec River
in Slovakia (TIRJAKovA 1993); China (SHEN & GONG 1989, SHEN & JIANG 1979, YANG
1989; Su et a1. 1988, as S. notophra); India (JACOB et at. 1993); Japan (EDMONDSON &
KINGMAN 1913); Lake Corona (3400 m above sea level) and Boulder Creek, Colorado,
USA (EDMONDSON 1912, HAMILTON 1943); Iowa, USA (SHAWHAN et al. 1947); among al-
gae in Creel Bay, Devil's Lake Complex, North Dakota, USA (EDMONDSON 1920); Nor-
man, Oklahoma, USA (TROWBRIDGE & STAMBAUGH 1935); Conestoga Basin and from
May to July 0.02-1.3 indo ml-I in a pond at University of Pennsylvania, USA (CAIRNS
1965a, WANG 1928); Venezuela (SCORZA & NuNEz MONTIEL 1954); Lake Chad, Chad
 Sty/onychia 555

(DRAGESCO 1972b; as S. putrina "petite fonne" of DRAGESCO & NJINE 1971). STAMM
(1984) recorded S. notophora from activated sludge plants in Switzerland.
Feeds on bacteria, unicellular algae, including diatoms (L6PEZ-OCHOTERENA 1965,
PRATT & CAIRNS 1985, PRATT et al. 1989, WANG & N!E 1935). Food vacuoles usually
crowded in posterior half of well-fed specimens (WANG & N!E 1935). Biomass of 106 in-
dividuals about 85 mg.
DETCHEVA (1978, 1979a, c, 1983c) found S. notophora in various rivers in Bulgaria
under following conditions: 0, b, a, frequency 1.3-2.l %, 10--24 DC, pH 6.3-7.6,
3.0--9.8 mg 1.1 O2 (30--87 % saturation), 1.2-12.4 mg 1.1 BOD5, 22-92 mg 1.1 Ca2+, 0 to
44 mg 1.1 Mg2+, 28-287 mg 1.1 HC03', 38-167 mg 1.1 SO/, 8-29 mg 1-1 CI-, 0.04 to
7.8 mg 1-1 NH/-N, 0--0.l8 mg 1-1 N02--N, 0--1.54 mg I-I N03--N, 0.02-1.5 mg I-I Fe2+.
PATRICK et aI. (1967) recorded it from the Savannah River, USA, under following condi-
tions (n = ?): pH 6.5-7.0,23-28 DC, 7-9 mg 1-1 0 2, 1-<5 mg I-I BOD5, ~llIg I-I NH/-N,
~0.007 mg I-I NOi-N, 0.2-<0.7 mg I-I N0 3--N, 0.1-0.5 mg 1-1 POlo, 5 to <10 mg {-I CI-.
According to Ax & Ax (1960), S. notophora tolerates up to 0.3 % salinity.
Saprobic classification: b; b = 7, a = 3, 1=4, SI = 2.3 (SLADECEK et al. 1981, WEGL
1983); b; b = 10, a = +, I = 5, SI = 2.3 (MoRAvcovA 1977; SI obviously incorrectly calcu-
lated); alphamesosaprobic (MoRAvcovA 1962; see also MAUCH 1976). Taking into con-
sideration the autecological and faunistic data, the classifications proposed by SLADECEK
et al. (1981) and MORAvcovA (1977) are possibly too good; further studies on reliably de-
termined populations recommended. ForssNER et al. (1991) excluded S. notophora from
the list of indicators of water quality.

Sup p 0 sed s y non y m: Stylonychia myti/us var. pusilla STERKI, 1878, Z. wiss.
Zool., 31: 56. Remarks: Incorrect subsequent spellings: Sty/onychia pusilus (BICK &
SCHOLTYSECK 1960, p 201); Sty/onychia pussila KAHL (KRAvCHENKO 1969, P 73). KAHL
(1932, P 618, Fig. 167e) considered it as valid species and mentioned it as "Stylonychia
pusi/la spec. n." expressly for the variety by STERK!. However, BORROR (1972a, P 15)
stated correctly that STERK! (1878) is the author of this taxon. Since KAHL (1932) classi-
fied Sty/onychia as subgenus of Oxytricha, the correct name in his revision is Oxytricha
(Stylonychia) pusilla. KAHL is the only one who gives an illustration of this fonn (Fig.
167e), which strongly resembles S. notophora. After STERK! (1878) 90--120 lim, accord-
ing to KAHL (1932) in life 75-90 lim long and about 2.5 times as long as broad Body
margins distinctly converging posteriorly, left one slightly convex or straight. Both ends
rounded. Moves hastily to and fro. 12-18 right and 10--14 left marginal cirri (STERKI
1878). Caudal cirri rather likely widely separated and fringed, similarly to in species of
the S. mytilus complex. SCHLEGEL (1985) and SCHLEGEL & STEINBRUCK (1986) studied the
macronuclear DNA and the aIlozyme variation, however, the identification is not substan-
tiated by an illustration. LYNN & CORLISS (1991, P 355) provided a scanning electron mi-
crograph of the dorsal surface of the SCHLEGEL population. The ecological data are rather
fragmentary. STERK! (1878) found it somewhere in Switzerland, and KARL (1932) ob-
served it together with S. mytilus. Records not substantiated by illustrations: Main River
 VI
VI
0\

'#
/ (> ~
• •/1 ~
Q 0 ~
• II ~ ~
~ •• \1 ~ 0° ~ ~
\\ ~ ~
~~ ~
~,;J~ :
~ CIl
• .• .- 00 -<
CIl
/I ~' ~~ ()
• .-•
/o () ~
()
•• • ~
# • o () >-1
~~': ..... (S
#. \ .- o o CIl
()
•••
/ o d\, o tTl
o ()
f • oy o >-1
()
. .. •• <> o o
00
~ o
T6~. • •• '
o ~
.../
-, b °0 o
a
.. --
c 0.0 o o
o d

Fig. 157a-d Sty/onychia bi/aria (from WIRNSBERGER et aI. 1985a a, from life; b-d, protargol impregnation). a) Ventral view, 135 Ilm. Note the inconspicuous caudal
cirri. b, c) Ventral and dorsal infraciliature, b, c = 80 Ilm. Note that the transverse cirri are separated into two distinct groups as in species of the S. my/ilus complex. Ar-
row = pretransverse ventral cirrus VIl2. d) Ventral infraciliature of a very early morphogenetic stage, 95 Ilm. Arrowhead marks oral primordium. TC = transverse cirrus
VVI; p 557.
 Stylonychia 557

and sewage ponds in Gennany (BERNERTH 1982, BICK & SCHOLTYSECK 1960, KAUFMANN
1958); Lake Dong Hu, Wuhan, China (SHEN 1980). Does not feed on algae (KAHL 1932).

Stylonychia hifaria (STOKES, 1887) comb. nov. (Fig. 157a-p, 159a,

Tables 3, 4, 31)

1887 Oxytricha bifaria STOKES, Ann. Mag. nat. Hist., 20: 110 (Fig. 157i).
1888 Oxytricha bifaria, STOKES - STOKES, J. Trenton nat. Hist. Soc., I: 292.
1931 Oxytricha bifaria STOKES - SAMANO & SOKOLOFF, Monografias Inst. BioI. Univ. naco Mex., I: 35, Fig.
71.
1932 Oxytricha bifaria STOKES, 1887 - KAHL, Tierwelt DtI., 25: 602.
1945 Oxytricha bifaria STOKES - KAy, Trans. Am. microsc. Soc., 64: 91 (Fig. 157j, I).
1953 Oxytricha bifaria STOKES (1887) - JiROVEC, WENIG, FOTT, BARTOS, WEISER & SRAMEK-HuSEK, Protozoo-
logie, p 511, 512 (Fig. I 57p).
1961 Histrio muscorum KAHL, 1930/35 - REUTER, Acta zool. fenn., 99: 21 (Fig. 159a; misidentification).
1962 Oxytricha bifaria STOKES - MORAVCOVA, Sb. vys. Sk. chem.-technol. Praze, 6: 380 (Fig. I 57n, 0).
1963 Oxytricha bifaria STOKES - LUNDIN & WEST, Free-living protozoa, p 67 (Fig. 157k).
1985 Sty[onychia vorax STOKES, 1885 - WIRNSBERGER, FOiSSNER & ADAM, J. Protozool., 32: 261 (Fig. 157a-h,
m; misidentification; authoritative redescription).

Nom e n c I at u rea n d t a x 0 nom y: The correct name in KAHL (1932) is Oxytri-

cha (Oxytricha) bi/aria because he divided Oxytricha into several subgenera. "Diophrys
bi/aria" and "Oxytricha scutum" in CHADHA et al. (1978, p 9) are erroneously combina-
tions; the correct names should probably be "Oxytricha bi/arid' and "Diophrys scutum".
STOKES (1887b) classified the present species in Oxytricha, very likely because of the
inconspicuous caudal cirri. However, the arrangement of the frontal, ventral, and trans-
verse cirri and the straight, parallel undulating membranes indicate that it is a
Stylonychia. According to the original description, S. bi/aria is about 11160 inch (=
159 !lm) long. Conversely, STOKES (1888) and subsequently KAHL (1932) gave a length of
1/100 inch (= 254 !lm; par lapsus?). Probably this last value is too large, although two re-
descriptions exist, both giving a length of 250 !lm (KAy 1945, LUNDIN & WEST 1963).
According to other redescriptions, S. bi/aria is 100-216 !lm long.
I have no doubt that S. vorax sensu WIRNSBERGER et al. (1985a) is in fact S. hi/aria.
This is indicated by the very good agreement in body shape, the inconspicuous caudal
cirri, and the arrangement of the transverse cirri (cp. Fig. 157a, b, i). Thus, the detailed
redescription by WIRNSBERGER et al. (1985a) should be considered as authoritative. "Oxy-
tricha bi/aria" of the Italian protozoologists (for example, ROSATI et al. 1988) is (very
likely) identical with Stylonychia pustulata (see there). The populations by CONN (1905;
Fig. 228g, 23li, j), SMITH (1914; Fig. 233t), SOKOLOFF (1931; Fig. 228b), and WANG
(1925; Fig. 242h) are insufficiently redescribed.

M 0 r p hoI 0 g y and b i 0 log y: Size in life obviously rather variable (see, how-
ever, taxonomy): about 160 !lm (REUTER 1961, STOKES 1887b), 140!lm (SAMANO & SOK-
 VI
VI
00

, ,
o ~
~

o /::=='"
,
0° ;r~~ ,
Q\ 1/~ §

o
<'
'0 • r/:l

<' ~~ -<
r/:l
...,
(> ":'.
00 ~J1 :
l.~·f(f: ... J ~
...,
':;~'.~ ~ .... ~
, ~:.,!:
n
-
o r/:l
0": tIl
t : 1. n
...,
0° . " 'J
() 5
() o o z
o c
o Q o.
D ,,0.
°0 '00 0 ' o Q Q ~'DO
C!0 e f 9 h
-
Fig. 157e-h Sty/onychia bi/aria (from WIRNSBERGER et al. 1985a; protargol impregnation). e) Early morphogenetic stage in ventral view, 95 Jlm. f-h) Morphogenesis of
dorsal infraciliature proceeds in Oxytricha pattern, f= 100 Jlm, g = 90 Jlffi, h"" 110 Jlm. Arrows mark new dorsomarginal kineties; dorsomarginal kineties of opisthe are
still on ventral surface; p 557.
 Sty/onychia 559

Fig. 157i-m Sty/onychia hi/aria (i, after STOKES 1887b; j, from KAy 1945a; k, from LuNDIN & WEST 1963; I,
after KAy 1945b; m, from WIRNSBERGER et aI. 1985a i-m, from life). i-k) Ventral views, i = 156 ~m, j =
216 ~m, k = 250 ~m (length from WEST & LUNDIN 1963).1) Resting cyst, 71 ~m. m) Lateral view; p 557.

OLOFF 1931), 100-250 Ilm (KAy 1945a), 140-180 x 70-120 Ilm (MORAvcovA 1962),
about 250 Ilm (WEST & LUNDIN 1963),93-137 x 48-55 Ilm (WIRNSBERGER et al. 1985a).
Right margin convex, left straight (Fig. 157a) or even slightly concave (Fig. 157i) and
with striking bend in posterior part, thus posterior end somewhat pointed. Flattened about
3: 1 dorsoventrally, first and last quarter of cell very thin, anterior part distinctly erected
(Fig. 157m, 0). Macronuclear nodules in life about 17 x 12 Ilm, arranged along median
or even slightly right of it, filled with about 2 Ilm sized chromatin bodies. Two micronu-
clei, in life about 4 Jlm across. Rapid and erratic movement.
Frontal cirri, buccal cirrus, frontoventral cirri III12 and IV/3, and the 2 right trans-
verse cirri distinctly enlarged. For arrangement of cirri, see Figure 157b. Right pretrans-
verse ventral cirrus at level of middle transverse cirrus. Transverse cirri in life 17-21 Ilm
long, characteristically arranged in a group of 3 left cirri forming an oblique row and
hardly reaching posterior end of cell and a detached group of2 cirri protruding distinctly
beyond posterior end of cell. Left marginal row terminates rather exactly at posterior end,
right one terminates slightly behind level of posterior-most transverse cirrus. Marginal
and frontal cirri in life 17-23 !lm long. Invariably 6 dorsal kineties. Rows 1 and 2 of
 560 SYSTEMATIC SECTION

body length, rows 3 and 4

slightly shortened posteriorly
and anteriorly respectively.
Caudal cirri not fringed, ra-
ther closely arranged, in life
only about 19-22 Jlm long
and thus hardly discernible
from marginal cirri.
Cyst globular, 60 to
80 !lm across. Ectocyst
brownish, with 6-8 rows of
long spines (Fig. 1571). Fur-
ther details on cyst
formation, see KAy (1945b).
Excystment can usually be
induced by washing the cysts
n o clean of old medium and de-
bris with several changes of
Fig. 157n-p Sty/onychia hi/aria (n, 0, after MORAvcovA 1962; p, tap-water and placing them
from JIROVEC et aI. 1953. n--p, from life). D, p) Ventral views, n =
in fresh nutritive solution.
170 J.lm, p = 200 J.lm. 0) Lateral view, 155 J.lm; p 557.
Two solutions have proved
suitable: 0.01 % tryptone in
tap water exposed to air for
8-12 h before use, or tap-water in which few grains of polished rice have stood for sev-
eral days. In either of these solutions a bacterial flora develops which is readily ingested
by the young excystants. The entire process of excystment usually takes about 18 h (KAy
1945b).
Conjugation can frequently be observed (WIRNSBERGER et al. 1985a). To induce this
part of life cycle, diverse cultures were provided with sufficient nutrient to allow 0.5 to 1
fission per day, and were mixed in a depression slide when they had exhausted most of
the available bacteria. SIEGEL (1956) found 9 mating types (see also MIYAKE 1981). Nu-
clear behaviour during conjugation see KAy (1946); she found that the diploid number of
chromosomes is 24.
M 0 r p hog e n e sis of cell division commences with the formation of the oral
primordium close to the intact one or two leftmost transverse cirri (Fig. 157d). Somewhat
later this primordium extends close to the cytostome (Fig. 157e). The next stages are
identical with those of S. pustulata (see Fig. 162f-n; WIRNSBERGER et al. 1985a), indicat-
ing a very close relationship of these species. The origin of some primordia is rather dif-
ferent from that of S. mytilus (Table 4, cpo Fig. 23a with 23f), suggesting the StylofT)'chia
is not monophyletic (see Fig. 25a). Morphogenesis of dorsal infraciliature shows no pe-
culiarities and follows the Oxytricha pattern (Fig. 157f-h). For details on macronuclear
division, see SESHACHAR & PADMAVATHI (1956) and SOKOLOFF (1930).
 Sty/onychia 561

SmGENAKA et al. (1973) tested different fixation techniques for electron-microscopic

studies on S. hi/aria. Furthermore, they show cross-sections of cilia and mitochondria.
VAN WAGTENDONK & SOLDO (1965) stated that S. hi/aria contains self-duplicating, rod-
like particles which take up nuclear stains such as aceto-orcein.

o c cur r e n c e and e colo g y: In freshwater, obviously prefers stagnant waters;

according to my experience not as common as S. pustu/ata. Probably cosmopolitan.
STOKES (1887b) discovered S. hi/aria in an infusion of hay; STOKES did not give the spe-
cific locality where he found it, however, he lived and worked in Trenton, New Jersey,
USA, and it is thus assumed that the locus classicus is near there.
Further records substantiated by illustrations: pasture ponds in the Schlossalm area,
Bad Hofgastein, Austria (WIRNSBERGER et al. 1985a); diverse cultures (bioclein, culture
with grains of wheat or barley, Pringsheim biphasic culture), Czechoslovakia (MO-
RAvcovA 1962, JiROVEC et al. 1953); rock pools in Finland (REUTER 1961); debris col-
lected in a pool in the Drug Garden of the University of Washington, USA (KAy 1945a);
various freshwater habitats from the Upper Peninsula of Michigan, USA (LUNDIN & WEST
1963, WEST 1953, WEST & LUNDIN 1963); ponds in Mexico (SAMANO & SOKOLOFF 1931).
Records from freshwater habitats not substantiated by illustrations: Armenia (AM-
BARTZUMYAN 1962, ZHARrKov 1982); Czechoslovakia (SRAMEK-Hu~EK 1953); eutrophic
pond (poppelsdorfer Weiher) in Bonn, Germany, from summer to early December (SoNG
& Wn..BERT 1989, as "S. vorax which agrees with the redescription ofWIRNSBERGER et al.
1985a"); Hiroshima, Japan (MATSUOKA et al. 1983); in freshwater bodies of the rocky
supra-littoral zone of the White Sea (BURKOVSKY 1971a); Conestoga River Basin, Penn-
sylvania, USA (CAIRNS 1965a); ponds and lake on Mount Desert Island, USA (McCASH-
LAND 1956); bottom debris of a pond in Sandy Point area on Abaco Island, Bahamas
(CAIRNS & RUTHVEN 1972); abundant in Creel and Whipple Bay ofDevil's Lake complex,
North Dakota, USA (EDMONDSON 1920, HAMME,l 1986); soft water Escambia River, Flor-
ida, USA (PATRICK 1961); rarely in a pond in the botanical garden of the University of
Pennsylvania, USA (WANG 1928); rare in clear springs and streams and clear small pools
with abundant decomposing organic sediment, New York, USA (HAUSMAN 1917, 1934);
Mexico (LoPEZ-OCHOTERENA & ROURE-CANE 1970); brook and artificial pool in Colombia
(BOVEE 1957); Venezuela (SCORZA & NuNEZ MONTIEL 1954); soil from flood plain pri-
mary (?) rain forest and from blackwater inundation primary (?) rain forest near Manaus,
Brazil (FOISSNER 1997b); <10 indo ml-l in mud surface (PH 5.8) of a volcanic lake on
Raoul Island, New Zealand (BROWN & PEART 1973).
Records from brackish, marine, or terrestrial habitats not substantiated by illustra-
tions: freshened site of the White Sea (salinity about 1 %0; BURKOVSKY 1976); rarely in
the Logy Bay, Newfoundland, Canada, a marine habitat (LACKEY & LACKEY 1970); soils
in Sweden and New Jersey, USA (KOFFMAN 1926, FELLERS & AllISON 1920); soil from
flood plains in Brazil, see previous paragraph.
PATRICK et al. (1967) recorded S. hi/aria, rarely, from the Savannah River, USA, un-
der following conditions: 23-28 DC, >5-7 mg I-I O2 , 1-<5 mg I-I BODs, 0.05-<1 mg 1-1
 562 SYSTEMATIC SECTION

NH3-N, 0.001-<0.007 mg 1-1 N02--N,

0.2-<0.7 mg 1-1 N03--N, 0.1-0.5 mg I-I
P04-. CAIRNS & YONGUE (1973) found
Stylonychia bifaria in the Cape Fear
River near Fayetteville, North Carolina,
USA, at 23-26°C, pH 6.2-7.1, 5.8 to
8.5 mg 1-1 O2, 0.9-3.8 mg I-I BOD (proba-
bly BODs), 5.7-14.7mgI-1 Cl-, 0.17 to
0.68 mg I-I NH/-N, 0.05-0.8 mg 1-1 N03--
N, 0.07-0.3 mg I-I P04--P (summruy of the
extreme values of3 sites).
KAy (1945a) established cultures on
tryptone solution and rice infusion with
bacteria as food. As a culture medium
tap water can be used, with yeast and a
species of the Tetrahymena pyriformis
Fig. 158a-c Oxytricha phytophaga, a supposed syno- complex grown on dry egg yolk added
nym of Sty[onychia hi/aria (from GELEI & SZABADOS as the food supply (WIRNSBERGER et al.
1950. a, sublimate fixation; b, c, opal-blue stain after
BRESSLAU)_ a) Ventral view, 105 J.1m. b, c) Ventral and
1985a). SIEGEL (1956) maintained mass
dorsal infraciliature, b = 60 J.1m, c = 50 J.1m. cultures and isolation lines in lettuce or
Cerophyl infusions inoculated with
Aerobacter cloacae. Biomass of 106 in-
dividuals about 54 mg (popUlation by
WIRNSBERGER et al. 1985a). Swims about 1.2 mm S-I (BULLINGTON 1925, identification un-
certain, because cells 235-329 J.1m long).
Stylonychia bifaria is not included in the lists of indicators of water quality by SLAoE-
CEK et al. (1981), WEGL (1983), and FOISSNER (1988a). According to MORAvcovA (1962,
1977) it is confmed to alphamesosaprobic areas (s = a; b = +, a = 10, I = 5, SI = 2.8). The
faunistic and ecological data mentioned above suggest the following classification: a-b; 0
= 1, b = 5, a = 4, I = 3, SI = 2.3; however, this has to be verified by further saprobiologi-
cal and ecological studies on reliably determined populations.

Sup p 0 sed s y non y m: Oxytricha phytophaga GELEI & SZABADOS, 1950, AnnIs
bioI. Univ. szeged., 1: 261, 282 (Fig. 158a-c). Remarks: BORROR (1972a) already pro-
posed synonymy with S. bifaria, however, since there are some minor differences, I pro-
vide a short characterisation of this Hungarian population. Large specimens in life about

Fig. 159a Sty[onychia hi/aria (after REUTER 1961). Ventral view from life, about 150 J.1m; p 557. b, c Sty[a- ~
nychia vorax (from DRAGESCO 1966b. b, c, protargol impregnation). b) Ventral view, 125 J.1m. Caudal cirri not
shown in full length. c) Posterior end with caudal cirri in dorsal view; p 591. d-f Sty[onychia notophora (d,
after SRAMEK-HuSEK 1945; e, after STOKES 1885b; f, after WANG & NIE 1935. d-f, from life). d) Ventral cilia-
ture in dorsal view, size not indicated. e, 1) Ventral views, e = 125 J.1m, f= 105 J.1m; p 553. g Sty[onychia myti-
[us complex (from DRAGESCO & NJINE 1971). Ventral infraciliature, protargol impregnation, 125 J.1m; p 504.
Stylonychia 563
564 SYSTEMATIC SECTION

140 x 90 !lm, starved specimens about 110 x 50-55 !lm. Especially starved specimens
often triangular in shape. Anterior part of cell distinctly erected. Two macronuclear nod-
ules connected by a conspicuous fibre; very likely 2 micronuclei. Contractile vacuole
about in mid-body. Feeds mainly on phytoplankton, especially on volvocales. Vivacious
and persistently swimming, obviously a typical planktonic form. Arrangement of frontal,
ventral, and transverse cirri, see Figure 15Sb; looks very similar to in S. bi/aria. Invaria-
bly 5 transverse cirri. A distinct separation of the transverse cirri into 2 groups is only
discernible in Figure 15Sb (thus not synonymous with Pseudostrombidium planctonti-
cum). 15-21 right and 12-lSleft marginal cirri. Caudal cirri obviously rather inconspicu-
ous. Six dorsal kineties, dorsal cilia 2--4 !lm long (Fig. 15Sc). Figure 15Sb probably
shows a young postdivider because the dorsomarginal kineties are still visible on the ven-
tral surface. Locus classicus is a rain puddle in Szeged, Hungary. "Oxytricha phytofaga"
in RAMlREz-MoNTESINOS & PEREZ-SILVA (1966) is an incorrect subsequent spelling.

Stylonychia pseudograndis WANG & NIE, 1935 (Fig. 167a)

1935 Sty/onychia pseudograndis WANG & NIE, Sinensia, Shanghai, 6: 507.

T a x 0 nom y: The increased number of cirri indicates misobservation or observation of

atypical specimens of another species. However, since WANG & NIE (1935) found this
pattern in numerous individuals, I would rather not, at this point, classify it as species in-
determinata. Possibly a giant of Stylonychia pustulata (cp Fig. 164c). Detailed redescrip-
tion necessary. BORROR (1972a) did not mention S. pseudograndis.

M 0 r p hoI 0 g y: In life 140-200 x 60-100 !lm. Elliptical, broadest at or slightly be-

hind mid-body, ± obliquely truncated anteriorly, rounded posteriorly. Macronuclear nod-
ules ovoid or elongate ellipsoidal, often connected by a thread. Cytoplasm appears
slightly darkish.
Usually 10 or 12 frontal, frontoventral, and buccal cirri. Frontal cirri distinctly en-
larged. As in IS-cirri oxytrichids invariably 5 postoral and pretransverse ventral and 5
transverse cirri, the latter not conspicuously enlarged and scarcely projecting beyond pos-
terior end of cell. Left marginal row J-shaped. Caudal cirri in life about 30 !lm long and
conspicuously shifted to the right.

o c cur r e n c e and e colo g y: Locus classicus is Nanking, China, where WANG &
NIE (1935) discovered S. pseudograndis in an old standing culture of a sample from a
freshwater habitat, together with high numbers of Glaucoma sp., Ophryoglena sp., and
Paramecium sp., indicating that S. pseudograndis prefers saprobic conditions. Not found
since. Voracious feeder (bacteria, Glaucoma sp., other ciliates). Biomass of 106 individu-
als about 215 mg.
 Sty/onychia 565

Stylonychia pustulata (MOLLER, 1786) EHRENBERG, 1835 (Fig. 3b, 20d,

22a, 154a-d, 160a-j, 161a-z, 162a-j, 163a-d, g-l, 164a-e, 165e-g,
Tables 3, 4, 18,31,36)

1786 Kerona pustulata MOLLER, Animalcula Infusoria, p 246 (Fig. l61r).

1830 Kerona pustulata MOLLER - EHRENBERG, Abh. preuss. Akad. Wiss., year 1830: 43, Tafel VI, Fig. 11- 13
(see also EHRENBERG 1831, P 119).
1835 Stylonychia pustulata - EHRENBERG, Abh. preuss. Akad. Wiss., year 1835: 164.
1838 Stylonychia pustulata - EHRENBERG, Infusionsthierchen, p 371 (Fig. 161 s).
1852 Cerona pustulata - PERlY, Kenntniss kleinster Lebensfonnen, p 154.
1858 Stylonychia pustulata EHR. - CLAPAREDE & LACHMANN, Mem. lust. natn. genev., 5: 161 (Fig. 16lt).
1858 Stylonychiafissiseta CLAPAREDE & LACHMANN, Mem. Inst. natn. genev., 5: 163 (Fig. 16Iu).
1859 Stylonychia pustulata. EHRBG. - STEIN, Organismus der Infusionsthiere I, p 161 (Fig. 161x, 162b).
1862 Stylonychia pustulata - ENGELMANN, Z. wiss. Zool., II: 354 (Fig. 162d).
1865 Stylonychia pustulata EHRBG. - QUENNERSTEDT, Acta Univ. lund., 2: 57 (Fig. 16lp, 162c).
1882 Stylonychia pustulata, EHll. - KENT, Manual infusoria II, p 791.
1882 Stylonychiafissiseta, C. & L. - KENT, Manual infusoria II, p 791.
1888 Stylonichia pustulata - MAUPAS, Archs Zool. expo gen., 6: 196 (Fig. 161v; incorrect subsequent spelling
of Stylonychia).
1888 Stylonichia mytilus - MAUPAS, Archs Zool. expo gen., 6: 212 (Fig. 162a; misidentification; incorrect
subsequent spelling of Stylonychia).
1899 Stylonychiapustulata EHRB. - PROWAZEK, Arb. zool. lust. Univ. Wien, II: 225 (Fig. 163a).
1901 Stylonychiapustulata (O.-F. MOLL.) - Roux, Mem. Inst. natn. genev., 19: 104 (Fig. 162e).
1905 Stylonychiapustulata (O.F. MOLL.)- BORGER, An. Univ. Chile, 117: 435 (Fig. I 63b).
1906 Stylonychiapustulata EHR. - EDMONDSON, Proc. Davenport Acad. Sci., II: 102 (Fig. 234c).
1914 Stylonychia pustulata EHR. - SMrrn, Kans. Univ. Sci. Bull., 9: 166 (Fig. 154d).
1931 Stylonychia pustulata EHRBG. - TAI, Sci. Rep. natn. Tsing Hua Univ., 1: 52 (Fig. 154a).
1932 Stylonychia pustuiata EHRENBERG, 1838-STEIN, 1859 - KAHL, Tierwelt Dtl., 25: 619 (Fig. 161 q, 162h).
1932 Stylonychiafissiseta CLAPAREDE u. L., 1958 - KAHL, Tierwelt Dtl., 25: 618.
1934 Stylonychia pustulata- CALKINS, BioI. Bull. mar. bioI. Lab., 67: 4Il (Fig. 163k, I).
1935 Stylonychia pustulata - SUMMERS, Arch. Protistenk., 85: 190 (Fig. 161z; in legend to plates on p 206
obviously erroneously designated as Oxytrichafallax).
1942 Stylonychia pustulata - LILLY, Physiol ZoOl., 15: 147 (Fig. 162j; possibly S. vorax!).
1950 Stylonychia notophora STOKES - BARY, Trans. R. Soc. N. Z., 78: 317 (Fig. 161y; misidentification).
1950 Stylonychia putrina STOKES - BARY, Trans. R. Soc. N. Z., 78: 317 (Fig. 163d; misidentification?).
1952 Stylonychia pustulata EHRBG - HASHIMOTO, Zool. Mag., Tokyo, 61: 332 (Fig. 163c).
1961 Stylonychia pustulata EHRENBERG, 1838-STEIN, 1859 - RElJ1"ER, Acta zool. fenn., 99: 21 (Fig. 162t).
1968 Oxytricha (Stylonychia) pustulata EHRENBERG, 1838 - CHORIK, Free-living ciliates, p 139 (Fig. 162w).
1969 Stylonychia pustulata EHRENBERG, 1838 - CURDS, Wat. Po\lut. Res., No. 12: 70 (Fig. 162g).
1970 Stylonychia notophora STOKES - SAPRA & DASs, Acta Protozool., 7: 193 (Fig. 162i; misidentification).
1972 Stylonychia pustulata EHRENBERG, 1838 - CELA, Physis, 31: 576 (Fig. I 54b).
1974 Stylonychia putrina STOKES - PATSCH, Arb. Inst. landw. Zool. Bienenkd., I: 63 (Fig. 165g; misidentifi-
cation).
1976 Stylonychia (Oxytricha) pustulata EHRB. - KORNIYENKo, Hydrobiol. 1., 12: 62.
1985 Stylonychia pustulata (MOLLER, 1786) EHRENBERG, 1838 - WIRNSBERGER, FOISSNER & ADAM, J. Proto-
zool., 32: 262 (Fig. 161a-o; authoritative redescription).
1986 Stylonychia putrina STOKES, 1885 - DRAGESCO & DRAGESCO-KERNEIS, Faune tropica1e, 26: 487; in part,
Planche 145 C-E (Fig. 165e, t); not Planche 145 A, B, F.
1988 Oxytricha hi/aria - ROSATI, GIARI & RICCI, Europ. J. Protistol., 23: 343 (Fig. 164a-c; misidentification).
566 SYSTEMATIC SECTION

1988 Stylonychia pustulata (0. F. MULLER, 1786) - SHIN & KIM, Korean 1. syst. Zool., Special Issue No.2:
73, Fig. 4A-F.
1991 Stylonychia pustulata (MUELLER, 1786) EHRENBERG, 1835 - FOISSNER, BLAITERER, BERGER & KOHMANN,
Infonnationsberichte des Bayer. Landesamtes flIr Wasserwirtschaft, 1191: 323 (Fig.l60a-j).
1991 Stylonychia pustulata - FOISSNER, Europ. 1. Protistol., 27: 323, Fig. 32, 33 (Fig. 160c).
1991 Stylonychiapustulata (MuLLER, 1786) EHRENBERG, 1838 -GSCHWIND, Dip1omarbeit, p 48 (Fig. 163g-j).
1991 Stylonychiapustulata - YANO & SUHAMA, J. Protozool., 38: 111 (Fig. 154c).
1997 Clara pustulata (MULLER, 1786) - EIGNER, 1. Euk. Microbiol., 44: 568 (new combination with Clara
EIGNER, 1997, ajunior homonym).

Nom e n c I a t u rea n d t a x 0 nom y: Stylonychia pustulata was originally de-

scribed by MOLLER (1786) from a marine habitat. EHRENBERG (1838), the frrst reviser, re-
corded S. pustu/ata from various freshwater habitats and from the sea near Copenhagen.
He mentioned some older synonyms (for example, Trichoda silurus MOLLER, 1773, Tri-
choda cyc/idium MOLLER, 1773) in his comprehensive list of synonyms, and it is unclear
why he did not use one of these senior names for the designation of the present taxon.
Somewhat later, STEIN (1859b) even questioned the identity of MOLLER'S (1786) marine
species and the freshwater form. According to STEIN (1859b), the freshwater species
Kerona silurus in MOLLER (1786, P 244) - originally described by MOLLER (1773, P 88)
as Trichoda silurus - agrees better with the form figured by EHRENBERG (1838). But de-
spite his doubts, STEIN (1859b) finally accepted EHRENBERG'S (1838) unfounded nomen-
clatural act of ignoring senior synonyms. The experiments by FINLEY (1930; identification
not substantiated by illustration!) on the tolerance of freshwater protozoa to increased sa-
linity indicated that S. pus/u/ata can live in the sea. Thus, the identification of the com-
mon freshwater form with the marine species, S. pustula/a, as done by EHRENBERG (1838)
should be accepted. This is a typical case where Article 23 (b) of the IcZN (1985) has to
be applied, that is, the principle of priority is not intended to be used to upset a long ac-
cepted name in its accustomed meaning through the introduction of an unused name that
is its senior synonym. Thus, the following senior synonyms of EHRENBERG'S (1838) list of
synonyms are nominated to be suppressed ("nomina oblita"): Trichoda cyclidium
MOLLER, 1773, T. pulex MOLLER, 1773, T. calvitium MOLLER, 1773, T. acarus MOLLER,
1773, Volvox oniscus ELLIS, 1769, Cyc/idium radians HERMANN, 1784 (the papers ofEL-
LIS and HERMANN are not cited in my reference list!). Another very detailed list of old
synonyms is given by DIESING (1850, P 163).
KAlIL (1932) classified Stylonychia as subgenus of Oxytricha; the correct names in his
revision are thus Oxytricha (Stylonychia) pustulata and O. (Stylonychia) fissiseta. Incor-
rect subsequent spellings: Stylonichia postulata (ENRIQUES 1905b, P 391; PATIL et al. 1993,
p 316); Stylonichia pustulata (BHATIA 1924, P 2; BRODSKY 1928, P 135); Stylonychia pos-
tulata (DILLER 1965a, P 231; GRACIA et al. 1987a, p 26; KOWALEwsKlEGO 1882, p 397,
402); Stylonychia pustula (GONKEL 1997, P 222); Stylonychia pusulata (HUNTER 1958,
p 23); Stylonychia pustalata (CLAPAREDE & LACHMANN 1858, P 158); Stylonychia pustutata
(SUMMERS 1935, P 175); Stylonychia pastulata (DUTIA 1961, P 212); Stylonychia pustula-
tus (BODENHEIMER 1937, P 270); Stylonychnia pustulatus (BALDENSPERGER 1927, P 205).
 Sty/onychia 567

.
..... lMR

e
. . ".
' . '-CC f

Fig. 160a-f Stylonychia pustulata (from FOlsSNER et aI. 1991. a, b, d, scanning electron micrographs; c, f, pro-
targol impregnation; e, from life, bright field micrograph). a, c) Ventral infraciliature. Arrows in (a) mark left
frontoventral cirrus (= cirrus 1III2) and posterior postoral ventral cirrus (V/3), respectively. b, 1) Dorsal infra-
ciliature; arrows in (b) mark caudal cirri which are arranged at the posterior end of dorsal kineties 1,2, and 4.
d) Resting cyst e) Ventral view of a specimen containing many food vacuoles. BC = buccal cirrus, CC = cau-
dal cirri, FC = right frontal cirrus, LMR = left marginal row, Ma = posterior macronuclear nodules, RMR =
right marginal row, TC = left transverse cirrus, 1, 6 = dorsal kineties 1 and 6; p 565.
568 SYSTEMATIC SECTION

The fIrst detailed redescription of S. pustulata was provided by STEIN (l859b, Fig.
161x) and included a description and illustrations of the morphogenesis and the conjuga-
tion. He stated that CLAPAREDE & LACHMANN (1858) obviously overlooked a frontoventral
cirrus and the right transverse cirrus (Fig. 161t). STEIN (1859b) and KAHL (1932) consid-
ered S. jissiseta CLAPAREDE & LACHMANN as valid species (Fig. 161 u); conversely, I agree
with BORROR (1972a) that it is very likely identical with S. pustulata, although the fringed
caudal cirri remind somewhat on the species of the S. mytilus complex. I<AHL (1932)
stated that in S. pustulata no transverse cirrus protrudes beyond the posterior end of the
cell, indicating a misobservation since this is in contradiction to almost all other descrip-
tions. Stylonychia pustulata sensu TATEM (1870) is rather surely identical with Ancystro-
podium maupasi (Fig. 198h, 0.
I agree with WIRNSBERGER et al. (1985a) that S. notophora sensu SAPRA & DASS
(1970; Fig. 162i) is as. pustulata and therefore all other papers concerning the "s. noto-
phora" of the Indian group are mentioned here.
The scanning electron micrographs of the "Oxytricha bi/aria" by some Italian work-
ers (for example, BANCHETII & RICCI 1986, ROSATI et al. 1988; see also AMMERMANN
1985) strongly suggest that this intensively investigated form is also a Stylonychia pustu-
lata (Fig. 164b, 231 t). This is mainly indicated by the arrangement of the transverse cirri,
which are obviously not separated into two distinct groups as in S. bi/aria; however, a de-
tailed description of this Italian population is needed before fmal assignment, especially
since RICCI et al. (1991c, p 1184) found both Stylonychia pustulata and "Oxytricha bi-
farid' in fIeld samples.
I suggest the detailed redescription ofWIRNSBERGER et al. (l985a) as authoritative be-
cause (i) the description agrees very well with that by STEIN (1859b), (ii) the population is
characterised both by live observations and after protargol impregnation, (iii) the popula-
tion is well characterised morphometric ally, (iv) the morphogenesis is studied in great de-
tail, and (v) slides ofprotargol-impregnated specimens are deposited in a museum.
The redescriptions of CARTER (1859; Fig. 220d), CHEN (1944; Fig. 242i), ClENKOWSKY
(1855; Fig. 222k, I), CONN (1905; Fig. 228t), DUJARDIN (1838, 1841), DUMAS (1929; Fig.
237a), EcKHARD (1846; Fig. 232h), FRoMENTEL (1876), KOFFMAN (1926; Fig. 231e),
LUNDIN & WEST (1963; Fig. 230e), and NAlDU (1965; Fig. 2241) are insufficient. Histricu-
Ius similis sensu CHARDEZ (1986, Fig. 178u-w) is possibly identical with S. pustulata
since the body shape, the cirral pattern, and the resting cyst are very similar.
Stylonychia pustulata is rather difficult to distinguish from S. putrina (see key); how-
ever, synonymy is very unlikely because S. pustulata forms spiny cysts (Fig. 160d,
161o-q, 162b, 163k), whereas smooth ones are described for S. putrina (STOKES 1885a).
Stylonychia vorax and S. curvata are also rather similar.

Fig. 160g-j Stylonychia pustulata (from FOISSNER et a1. 1991. g-j, scanning electron micrographs). g, h) Ven- ~
tral view of a broad and a slender specimen. Arrow in (g) marks a caudal cirrus. i) Buccal area j) Posterior end
in dorsal view to show attachment of caudal cirri (arrows) at posterior end of dorsal kineties 1,2, and 4. AZM
 Sty/onychia 569

= adoral zone of membranelles, BC = buccal cirrus, e = endoral, p = paroral, TC = transverse cirri, VC = pre-
transverse ventral cirri, 1-4 = dorsal kineties 1-4; p 565.
 570 SYSTEMATIC SECTION

'# •
'. d

""
o"
o

o"
o"
o
o o
o o
o
o D
o D
o Cl
o
o CJ
o CJ
o o
o o
o o
o o
o o
o o
o
~o o
~()
e

Fig. 161a-f Stylonychia pustulata (from WIRNSBERGER et a1. 1985a. a, b, from life; c-f, protargol impregna-
tion). a, b) Ventral and lateral view, a = 80 Jlm, b = 105 Jlm. c, d) Ventral and dorsal infraciliature, c, d =
80 Jlm. Arrow marks pretransverse ventral cirrus VI/2. e) Ventral infraciliature of a very early morphogenetic
stage (70 Jlm) showing de novo formation of oral primordium (arrow). 1) Ventral infraciliature of a middle
 Stylonychia 571

BERGER & FOISSNER (1997) stated that Stylonychia is very likely not monophyletic,
that is, Stylonychia pustulata and S. bi/aria are probably not the sister group of the type
species, S. mytilus (Fig. 25a). Stylonychia mytilus is obviously very closely related to
Coniculostomum and Steinia. We suggested that molecular biological data on Coniculo-
stomum and Steinia should be awaited before splitting Stylonychia. Later, EIGNER (1997)
transferred Stylonychia pustulata and S. vorax, but not S. mytilus, to the Parakahliellidae
EIGNER, 1997. Thus, he had to establish a new taxon, Clara EIGNER, 1997, which is, how-
ever, a junior homonym that has to be replaced. I preliminary prefer my conservative
classification, that is, I do not accept Clara EIGNER (further comments on the classifica-
tion proposed by EIGNER 1997, see p 929).

M 0 r p hoi 0 g y and b i 0 log y: Size rather variable: 86-208 !lm long, on average
148 x 69 !lm (PROWAZEK l899a, STEIN l859b), 230 x 96 !lm (BDRGER 1905), 180-220
times 85-105!lm (Roux 1901), l50-230!lm (BHATIA 1936), 100-120!lm (REuTER
1961), 48-124 x 26-83 !lm (WIRNSBERGER et al. 1985a); according to ZINGHER & NAR-
Burr (1931) and ZINGHER et al. (1932) size varies between 74 x 34!lm in starved and
166 x 81!lm in well-feed specimens. ZHINGHER (1935) found that S. pustulata is some-
what longer at low temperatures (7 DC: mean = 99 !lm; n = 925) than at high ones (25 DC;
mean = 90 !lm; n = 308), possibly due to lower generation time. RICCI & RIGGIO (1984)
stated that linear dimensions of giants (200-220 x 90-100 !lm) are almost twice those of
normal cells (100 x 50 !lm); for detailed morphometric comparison of normal cells and
giants of the Italian "Oxytricha bi/aria", see ROSATI et al. (1988). Body margins usually
slightly convex, that is, outline normally elliptical, sometimes, however, almost in
parallel, as in S. putrina; rarely left margin slightly concave. Anterior end broadly, poste-
rior usually narrowly rounded. Flattened about 2:1 dorsoventrally, central dorsal portion
often strongly bulging (Fig. l6lb). Macronuclear nodules in life about 16 x 7 !lm, lie
about in median or slightly left of it. Usually 2, in life 4 x 3 ~m sized micronuclei (DEN-
NAKER & HUNTER 1965, WIRNSBERGER et al. 1985a). Contractile vacuole pulsates about
every 8-11 s at 20 DC (GELEI 1935, KANITZ 1907, PROWAZEK l899a, SMYTH 1942); takes
20 min at 24 DC to excrete the amount of water which corresponding to cell volume
(BRAND 1935, MAUPAS 1883). Cytoplasm usually densely filled with cytoplasmic crystals.
Movement gliding and often standing still for some time, left rotation about longer axis
(BULLINGTON 1925, VERWORN 1889). RICCI (1981b, 1990) measured an apparent velocity
(= velocity of the ciliate along the central axis of the helicoid) of750 ± 300!lm S·I. RICCI
(1989) found that his "Oxytricha bi/aria" (see taxonomy) is a species whose behaviour is
strongly affected by the dimensional quality of the substrate - namely by its granulo-
metry. Singlets avoid sands with grains ranging from 50-100 !lm in diameter, while con-
jugating pairs also avoid those with grains 100-200 ~m in diameter. Moreover, in sand

~ morphogenetic stage, 75 Jim. Arrowhead marks disintegrated posterior-most frontoventral cirrus (cirrus !v13),
arrow denotes primordium of frontal, ventral, and transverse cirri streaks of opisthe originating from oral pri-
mordium; p 565.
 572 SYSTEMATIC SECTION

Fig. 161g-k Stylonychia pustulata (from WIRNSBERGER et a1. 1985a g-k, ventral infraciliature of morphoge-
netic stages after protargol impregnation). g, b) Middle stages, g, h = 75 f.lm. Arrow marks primordium of
frontal, ventral, transverse cirri streaks of proter originating from parental cirrus IV13. i-k) Late stages, i-k =
 Sty/onychia 573

Fig. 16U--q Stylonychia pustulata (1-0, from WIRNSBERGER et al. 1985a; p, after QUENNERSTEDT 1865; q, from
KAHL 1932. I-n, protargol impregnation; o-q, from life). I) Ventral infraciliature of a very late morphogenetic
stage, 85 11m. Cirri originating from individual primordium are connected by a broken line; parental structures
white, new ones black. m, n) Ventral infraciliature ofproter and opisthe, m = 55 11m, n = 50 11m. o--q) Resting
cysts, 0 = 42 11m, p, q = size not indicated; p 565.

with grains of diameter <50 J.lm, singlets behave in much the same way as on flat sub-
strates; the effect appears to be due to body length.
Frontal and transverse cirri distinctly enlarged, in life about 19-24 J.lm long. Left
frontoventral cirrus (111/2) arranged immediately beside other frontoventral cirri (Fig.
161 c), which is very likely an important difference to the rather similar S. putrina. All
transverse cirri protrude distinctly beyond posterior end of cell. Right pretransverse ven-
tral cirrus arranged immediately in front of right transverse cirrus (Fig. 161c, arrow).
Marginal cirri in life about 15 J.lm long. Six dorsal kineties, rows 1-4 of about body
length, composed of 19-29 pairs of basal bodies each. Dorsomarginal rows (kineties 5
and 6) rather short. Caudal cirri in life 16-25 J.lm long, not fringed, very stiff, indistinctly

+- 80 11m. Cirri originating from individual primordium are connected by a broken line; parental structures white,
new ones black. DP = primordium of dorsomarginal kineties, MP = primordium of marginal row; p 565.
 574 SYSTEMATIC SECTION

Fig. 161r-z, 162a, b Sty/onychia pustulata (r, after MOLLER 1786; s, after EHRENBERG 1838; t, II, after CLA-
PAREDE & LACHMANN 1858; V, a, after MAUPAS 1888; w, from CHORIK 1968; X, b, after STEIN 1859b; y, after
BARY 1950; z, from SUMMERS 1935. r-z, a, b, from life). 1 am not quite sure if all these figures in fact represent
S. pustulata. Thus, identification should be according to the authoritative redescription by WIRNSBERGER et aI.
(1985a). r) Dorsal view, size not indicated. s-z, a) Ventral views, s-t, x = size not indicated, u = about
100 ~m, v, a= 145 ~m, w= 140~m,y= 130~m,z= 105 ~m. b) Resting cyst, size not indicated; p 565.

separate; distance between caudal cirri of kineties 1 and 2 always smaller than that be-
tween cirri of kineties 2 and 4 (Fig. 161d).
Cyst with spiky surface and fme bright cytoplasm, in life about 40--55 )lm across
(Fig. 160d, 161o-q, 162b, 163k, 164d, e; BALBIANI 1861, CALKINS 1934, FERMOR 1913,
FOissNER et al. 1991, KARL 1932, PROWAZEK 1899a, QUENNERSTEDT 1865, RICCI et al.
 Stylonychia 575

,,
,,
,"," ....
,
, I ,\
9

Fig. 162c-j Stylonychia pustulata (c, after QuENNERSTEDT 1865; d, from ENGELMANN 1862; e, after Roux
1901; f, after REmER 1961; g, from CURDS 1969; h, from KAHL 1932; i, from SAPRA & DASS 1970a; j, from
LILLY 1942. c-h,j, from life; i, wet silver impregnation). c, e-h) Ventral views, c, f=? /lm, e = 95/lm, g, h =
150 /lm. d) Conjugation, 190 /lm. i) Very early morphogenetic stage in ventral view showing oral primordium
(arrow), ? Ilm. j) This population could also be as. vorax, as indicated by the tapered posterior end; p 565.

1985, STEIN 1859b, VERNI et al. 1984, WIRNSBERGER et al. 1985a). The cyst figured by
STEIN (1854) has no spines, indicating that it is a very young one. PROWAZEK (l899a) ob-
served cysts infected with flagellates and amoebas. Cysts fonned in aged cultures are not
576 SYSTEMATIC SECTION

#'

-
I
I
t
""
""
';
I • II.
""
""
", "~.:

h
 Sty/onychia 577

viable (KAUSHAL & SAXENA 1978). 15 and 30 %

NaOH destroyed ecto- and endocyst (ILOWAISKY
1926b), and continuous reduction of oxygen content
induced encystment (MICHELSON 1928). According to
LACHMANN (1856, P 380, footnote 3), Discodella
multipes WEISSE, 1845 is only the cyst of S. pustu-
lata.
M 0 r p hog e n e sis of cell division is de-
scribed by HAslllMOTO (1952), SAPRA & DASS
(1970a), STEIN (l859b), WALLENGREN (1902, P 50),
and in great detail by WIRNSBERGER et al. (1985a)
and YANO & SUHAMA (1991). The oral primordium
originates de novo between the left marginal row
Fig. 163k, 1 Stylonychia pustulata
and the postoral ventral cirri (Fig. 154c, 161 e, 162i, (from CALKINS 1934). k) Resting cyst
arrows). This is the unique difference in cortical from life. I) Ventral view, size not indi-
morphogenesis between S. pustulata and S. bi/aria cated; p 565.
(WIRNSBERGER et al. 1985a). Therefore, Figures
161 f-n are valid for both species. Only the anterior
third of the parental undulating membranes is reorganised (Fig. 161g-k). The primordium
of the undulating membranes of the opisthe is formed by a group of basal bodies extend-
ing toward to the anterior right of the oral primordium (Fig. 16lf, arrow). The pharyngeal
fibres of the proter are not visible with protargol in later stages of morphogenesis (Fig.
161L n); probably they are reorganised. The frontal-ventral-transverse cirri streaks de-
velop in chronological order as follows: 3 streaks in the proter originate from the
posterior-most of the frontoventral cirri (cirrus IV/3, Fig. 161f, g, arrowhead and arrow).
At the same time, 3 anlagen in the opisthe separate from the oral primordium (Fig. 161f).
Two streaks evolve from the dispersion of the right and one from the left postoral ventral
cirrus (Fig. 161g, h). In the proter, additional basal bodies from the undulating mem-
branes, the buccal cirrus, and the left frontoventral cirrus (IIII2) participate in the forma-
tion of the primordia (Fig. 161h). Uniformly, 6 streaks are produced in each filial product
(Fig. 16li, j). See Table 4 and Figure 23a for a summary of the morphogenesis. The de-
velopment of the marginal primordia always starts within the right row and proceeds
more rapidly than in the left row in the earlier stages (Fig. 161h, i). Morphogenesis of
dorsal kineties proceeds identically to in S. bi/aria (Fig. 16lf-h), that is, in Oxytricha

~ Fig. 163a-d, g-j Stylonychia pustulata (a, after PRowAZEK 1899a; b, after BORGER 1905; c, after liAslllMOTO
1952; d, after BARY 1950; g-j, from GsCHWIND 1991. a, after alum or borax carmine stain; b, d, g, h, from life;
c, method?; i, j, protargol impregnation). a-d, g) Ventral views, a, c = size not indicated, b = 230 11m, d =
100 11m, g = 90 11m. b) Lateral view. i, j) Ventral and dorsal infraciliature, 60 11m. Arrow marks anteriorly
slightly shortened dorsal kinety 4; p 565.
Fig. 163e, r Stylonychia mytilus complex (e, after DEMBOWSKA 1925; f, after WALLENGREN 1902. e, f, from
life). e) Arrangement of cirri in dorsal view, size not indicated. 1) Ventral view, size not indicated. Caudal cirri
not shown; p 504.
578 SYSTEMATIC SECTION

pattern. Scanning electron micrographs of these events are provided in GRIMES & ADLER
(1976).
Pattern formation in mirror-image doublets is described by YANO & SUHAMA (1991,
see also FRANKEL 1991) and division and reorganisation of the nuclear apparatus was in-
vestigated by IVANIC (1931a), SUMMERS (1935), and TCHANG (1931). Replication bands
move synchronously at a uniform rate and take about 6 hours (SHARMA et al. 1986). Mac-
ronuclear S phase occupies 1/3-112 of cellular cycle (DINI et al. 1975). Further papers
concerning nuclear apparatus: ENRIQUEs (1912), PROWAZEK (1899b).
GRIMES & ADLER (1978) studied the regeneration of the ciliary pattern in longitudinal
fragments. Conjugation was studied by BOTsCHLI (1876, P 324), ENGELMANN (1862; Fig.
162d), PATSCH (1974), PIERI (1966), and PROWAZEK (1899a). ENRIQUEs (1905a, b) and
MAUPAS (1888, see also JENNINGS 1939) described senile degeneration.
WATANABE (1959) performed some experiments on the reversal action of cirri in-
duced by treatment with calcium and potassium ions, and on the relation of the reversal
action of the cirri to the macronucleus.
The complete small subunite rRNA sequence for S. pustulata is 1771 bases long,
which is identical to Oxytricha nova (identical with Sterkiella histriomuscorum) and
Onychodromus quadricornutus; G + C content 45 % (LEIPE et al. 1994). There are many
other new and old papers on genetics, sexual phenomena, including mating type of deter-
mination, biochemistry, ultrastructure: BALBIANI (1858, 1862, 1882), BOTscHLI (1875),
CANELLA (1959), DAWSON et al. (1983), DENNAKER & HUNTER (1965; see also DUTTA
1974), ELWOOD et al. (1985), ENGELMANN (1875), ENTZ (1901, 1904b), FERREIRA & PEREZ
(1936), GAJADHAR et al. (1991), GREENWOOD et al. (199Ib), HALL (1931), IVANIC (1931a,
b), KAzANCEV (1928), KLOBUTCHER et al. (1981), KRAUT et al. (1986), KUZNICKI & WALNE
(1993), LYNN & SooIN (1988), MAUPAS (1887, 1888, 1889), OKA & HONJO (1983), PIERI
(1965), PRESCOTT et al. (1979, 1982), RAo (1958), SCHLEGEL (1985, 1991), SCHLEGEL &
STEINBROCK (1986), SCHLEGEL et al. (1991), SIMPSON (1901), STEINBROCK (1990), SWANTON
et al. (1980b, 1982), YANO (1979, 1985a-d, 1986, 1989), YANO & SUHAMA (1990a, b,
1992).
As mentioned above, WIRNSBERGER et al. (1985a) synonymised S. notophora sensu
SAPRA & DASS (1970a) with S. pustulata. There are various other papers by these Indian
protozoologists using "S. notophora" as model organism: DASS & SAPRA (1966, 1967,
1968), KAuSHAL & SAXENA (1978), Rup LAL & SAXENA (1980), Rup LAL et al. (1981), SA-
PRA & DASS (1969, 1970b, 1971), SAXENA & Rup LAL (1981), SHIVAJI et al. (1975,1976,
1978).
The "Oxytricha bi/aria" of some protozoologists in Pisa, Italy (Fig. 164a--e; see tax-
onomy) is the subject of many genetical, biochemical, physiological, and ultrastructural
investigations (culturing technique: RICCI et al. 1980b; biochemistry: KRAMER 1988,
SCHLEGEL & STEINBROCK 1986; nuclear phenomena: RICCI & BANCHETTI 1981; homopolar
doublets: BANCHETTI & RICCI 1986, BANCHETTI et al. 1985, 1996, RICCI et al. 1996; canni-
balism, giants: BANCHETTI & RICCI 1987, BRAY! et al. 1991, GRANDINI et al. 1991, RICCI &
BANCHETTI 1989, 1993, RIcCI et al. 1979a, b, 1989a, 1991b, RIGGIO et al. 1987a, b, ROSATI
 Stylonychia 579

Fig. 164a, b Stylonychia pustuiata (from ROSATI et a1. 1988. a, b, scanning electron micrographs). Normal
cells in dorsal and ventral view, scale bars = 20 11m. This is the "Oxytricha hifaria" of the Italian protozoolo-
gists; however, the transverse cirri are not arranged in two separate groups, strongly suggesting that their iden-
tification is incorrect. Further data are needed for final decision. Explanation of original labels: FC = frontal
and frontoventral cirri and buccal cirrus, IC = transverse cirri including pretransverse ventral cirri, VC = post-
oral ventral cirri, 1-6 = dorsal kineties 1-6; p 565.

et al. 1987, 1988; conjugation: BANCHETII et al. 1978a, 1982b, DINI & LUPORINI 1974,
ESPOSITO & NOBILl 1982, ESPOSITO & RICCI 1975, ESPOSITO et al. 1976b, LUPORINI & DINI
1975, RICCI 1981a, 1982, RICCI & BANCHETII 1982, RICCI & CETERA 1978, RICCI et al.
1975a, b, 1977, 1980a, c; cyst: RICCI et al. 1985, ROSATI et al. 1984, VERNI et al. 1984;
motile behaviour and cell-substrate interactions in locomotion: BARBANERA et al. 1997,
ERRA et al. 1989, 1990, 1996, PINTo et al. 1991, RICCI 1981b, 1985, 1987, 1989, 1990,
1992, RICCI & ERRA 1996, RICCI et al. 1989b, c, 1991c, 1992a, b, Russo et al. 1988,
1989a-<:, 1990, 1992). According to RICCI et al. (I991a) the heteromorphic life cycle of
this hypotrich comprises at least 3 phases: conjugating pairs, resting cysts, and carnivo-
rous giants. The gigantic forms represent unique adaptive devices which enable the spe-
cies to survive in a certain environment when the normal bacterial food is exhausted.
 580 SYSTEMATIC SECTION

Fig. 164c-e Sty/onychia pustu/ata (c, from ROSATI et aI. 1988; d, e, from RIccI et aI. 1985. c,scanning elec-
tron micrograph; d, bright field; e, interference contrast). c) Giant cell in ventral view. The shape and the su-
pernumerary frontoventral cirri (arrows) are reminiscent of Sty/onychia pseudograndis (cp. Fig. 167a). Scale
bar = 20 ~m. d, e) Resting cysts, scale bars = 45 ~m, 10 ~m. Explanation of original labels: FC = frontal and
frontoventral cirri and buccal cirrus, M= macronucleus, m = micronucleus; TC = transverse cirri including
pretransverse ventral cirri, VC = postoral ventral cirri; p 565.

o c cur r e n c e and e colo g y: One of the most common freshwater oxytrichids,

often rather abundant in the aufwuchs and detritus; prefers mesosaprobic, slowly running
and eutrophic, stagnant waters; sometimes pelagic and in the air-water interface (RICCI et
al. 1991c), and also marine (see locus classicus and PATIERSON et al. 1989). Avoids hy-
drogen sulphide (WETZEL 1928b).
Locus classicus is the sea near Denmark, where MOLLER (1786) found Sty/onychia
pustulata together with Uronychia transfuga. EHRENBERG (1838) recorded it from infu-
sions and from freshwater habitats with Oscillatoria at Berlin, Germany, but also from
the sea near Copenhagen, Denmark. Locus classicus of the synonym, Sty/onychia [IS-
 Stylonychia 581

siseta, is a peat-bog (Jungfernheide) near Berlin, Germany (CLAPAREDE & LACHMANN

1858).
Records from freshwater habitats substantiated by illustrations: in putrefying, stag-
nant water bodies, especially in shallow ponds with decaying leaves, and once in the
drinking water of Vienna, Austria (PROWAZEK 1899a); lakes (Lunzer-See, Mondsee) in
Austria (GSCHWIND 1991, WIRNSBERGER et al. 1985a); putrescent infusions, dung-pools,
and other stagnant waters in Czechoslovakia (STEIN 1859b); rock pools (salinity
<0.01-1 %) in Finland (REUTER 1961); France (MAUPAS 1888); aufwuchs of freshwater
habitats in Bonn, Germany (PATSCH 1974); Leipzig, Germany (ENGELMANN 1862); oli-
gosaprobic to betamesosaprobic brooks and beta- to alphamesosaprobic brooks and riv-
ers in Bavaria, Germany (ForssNER 1997a, ForssNER et al. 1991, 1992a, b); Sweden
(QUENNERSTEDT 1865); common in clear and polluted lentic waters near Geneva, Switzer-
land (Roux 1901); USSR (CHORIK 1968, LoKOT 1987); pond in Chieh-Pei-Tze Yuan,
China (TAl 1931); Han River, Korea (SHIN & KIM 1988, 1993a); India (BHATIA 1936, SA-
PRA & DASS 1970a); Japan (HAslllMOTO 1952); vegetable infusions, Kansas, USA (SMITH
1914); New York (SUMMERS 1935); common in infusions of lentic water, USA (EDMOND-
SON 1906); freshwater pond at Woods Hole (Massachusetts) and from various freshwater
habitats near Providence, Rhode Island, USA (LILLY 1942); Argentina (CARBONELL 1935,
CELA 1972); Chile (BORGER 1905, 1908); Cotonou, Benin (DRAGESCO & DRAGESCo-
KERNEls 1986); stream at Beatrix Bay, Pelorus Sound, and from the Wainui-o-mata Res-
ervoir, New Zealand (BARY 1950).
The "Oxytricha bi/aria" of the Italian protozoologists was originally collected from a
freshwater canal in San Rossore, Pisa, Italy (DINI et al. 1975, RIccI et al. 1975a).
Numerous records from various freshwater habitats not substantiated by illustrations.
Europe and Asia: drinking water, Linz, Upper Austria (SCHIEDERMAYR 1882); Danube
River in Vienna, Austria (KALTENBACH 1960); mesosaprobic Traun River in Upper Aus-
tria (ForssNER & MOOG 1992); ponds, rivers, and many other habitats in Austria (BLAT-
TERER 1989, DALLA TORRE 1891, ForssNER & ForssNER 1988, SCHMARDA 1846, SPANDL
1926a, own observations); Belgium (CHARDEZ 1987); rivers, brooks, lakes, ponds, bogs,
ditches in Bulgaria (DETCHEVA 1979b, 1981, 1991, 1993, DETSCHEWA 1972); common in
the Moldau River, Czechoslovakia (KALMUS 1928); together with S. mytilus and among
putrefying material in a pond, Czechoslovakia (SVEC 1897; further record from Czecho-
slovakia: SRAMEK-HuSEK 1953); with 0.6 % frequency in moderate (10-100 ind. mI·l ) and
small numbers «10 indo mI- l ) in sites with sewage fungus outbreaks in England (CuRTIS
& CURDS 1971); backwaters of River Earn and other freshwater sites near Perth, England
(CRAIGIE 1921); pond in England (DIXON 1920); puddle and pond in Estonia (JACOBSON
1928); lentic small water bodies on an island in Finland (LEVANDER 1900); France (OR-
MANCEY 1852); caves in Germany (GRIEPENBURG 1939, see also GITILESON & HOOVER
1969); polluted Hamburg harbour, Germany (HECKMAN et al. 1990); small water bodies,
peat-bog, and dystrophic lakes in Germany (SCHMIDT 1913, 1916, STRODER-KypKE 1996);
hay infusion (?; probably with water from a pond or brook) in Germany (BRAND 1923;
further record from Germany: HENDERSON 1905); moraine lake and other sites in Green-
582 SYSTEMATIC SECTION

land (ELLINGER 1914, LARSEN 1992); plankton of the mesosaprobic Danube River in Hun-
gary (BERECZKY 1969, 1977c, 1979, BERECZKY et al. 1983, NOSEK & BERECZKY 1981); dis-
tinctly polluted Tisza River in Hungary (J6SA 1974, STILLER 1942); Lake Balaton, Hun-
gary (ENTZ 1897, FRANCE 1897, GELLERT & TAMAs 1958; further records from Hungary:
HORVATH 1933, KREpUSKA 1917); on surface (max. 226 indo cm-2) and bottom (max.
16 indo cm-2) of a productive Lemna pond in Italy mainly in summer (MAnoNI 1991 a, b);
Lake Como, Italy (CATTANEO 1882; further records from Italy: CANELLA 1954, CUNEO
1891, ENRIQUES 1913, GRISPINI 1938, LONGHI 1894, MOLA 1928, 1935, STELLA 1947);
common in the Netherlands (VERSCHAFFELT 1930, WIERSMA-VERSCHAFFELT 1936); Poland
(JAWOROWSKI 1893, WRZESNIOWSKI 1861, WRZESNIOWSKIEGO 1866); very abundant in mud
samples from the Danube River in Romania (SPANDL 1926b; further record from
Romania: LEPSI 1932); Lobregat River, Spain (GRACIA et al. 1987a); Sweden (QUEN-
NERSTEDT 1869); alpine lakes and brook about 2065 m above sea level in the Swiss (7)
Alps (BAUMANN 1910, ZSCHOKKE 1900); ponds and peat-bogs in Switzerland (MERMOD
1914, MESSIKOMMER 1948, 1954, PERTY 1852b); littoral of Lake Geneva, Switzerland
(ANDRE 1916, FOREL 1904, Roux 1900, SCHNETZLER 1854); in September up to
10 indo ml- l in the botanical garden of Basle, Switzerland (RIGGENBACH 1922; further re-
cords from Switzerland: ANDRE 1912, 1915, BOURQUIN-LINDT 1919, SAKOWSKy-CAMPIONI
1906, STEINMANN & SURBECK 1918); sandy sediment from Oka River, Russia (SASSUCHIN
et al. 1927; further record from Russia: AVERINTZEV 1899); very common in the littoral
area of Lake Baikal at 5-19 °C (GAJEVSKAJA 1927, GAJEWSKAJA 1933); benthal and pela-
gial of reservoirs in Azerbaijan (ALEKPEROV 1980, 1982b, 1983, 1984b, c); Volga River,
USSR (MAMAEVA 1979a, ZYKOFF 1903); plankton of pond in Kiev, Ukraine, during win-
ter (KOVALCHUK & SVINTSOVA 1994); on silt in small rivers in Latvia (VEYLANDE & LlYEPA
1985); Turkestan (DADAY 1903, 1904); mesotrophic reservoir and eutrophic lake in Ku-
ban Region, USSR (KORNIYENKO 1972; further records from the USSR: BELOVA 1990,
CHORIK & VIKOL 1973, EICHWALD 1844, 1852, MAMAEVA 1979b, MiNKEWITSCH 1898,
WEISSE 1845, 1848a); in December in lentic water from a burrow close to a garden well
in the Gol Bagh, India, together with Bursaria truncatella (BHATIA 1922; further records
from India: BHATIA 1924, BHATIA & MULLICK 1930, l<HAmRIA 1951); Palestine (BODEN-
HEIMER 1937, LEPSI 1929a); Kandy, Sri Lanka (DADAY 1898).
America: Costa Rica (RUIZ 1961); tap water (22°C, pH 7) from Mexico City (RIVERA
et al. 1979; further record from Mexico: L6pEZ-OcHoTERENA & ROURE-CANE 1970); abun-
dant at 3-6 °C and pH 6.0-6.4 in pools near Point Barrow, Alaska (SULLIVAN 1957); Iowa
(SHAWHAN et al. 1947); Kansas (CRAGIN 1889); at 20-24°C and pH 6.5-7.0 in upper
Black River and Douglas Lake, Michigan (CAIRNS & YONGUE 1966); natural waters in
Upper Peninsula of Michigan (WEST 1953); Brush Lake, Ohio (RIDDLE 1905); abundant
in polluted zones of the Scioto River in Ohio (LACKEY 1938b); tree holes in Ohio and
North Alabama (LACKEY 1940); Oklahoma (GABEL 1927); common in the Conestoga
drainage basin, Pennsylvania (CAIRNS 1965a); pond in the botanical garden of the Univer-
sity of Pennsylvania (WANG 1925); in a metal tank on hillside farm with much cyanobac-
teria and sulphur bacteria at 22°C and in a rapid, turbid mountain stream over rocks with
 Stylonychia 583

high Orcontent in Virginia (BOVEE 1960); shallow streams with mediwn hard water in
eastern USA (PATRICK 1961); ponds, pools, still areas along stream margins, and polluted
streams in USA (LACKEY 1938a); marsh pools, probably in USA (HAUSMAN 1917); Ar-
gentina (BUSTOS 1933, SECKT 1924); cistern in Rio de Janeiro, Brazil (OLIVEIRA et a1.
1954; further records from Brazil: CUNHA 1913, PRowAZEK 1910); Rio Rondos, Peru
(CAIRNS 1966a); Venezuela (SCORZA & NuNEz MONTIEL 1954).
Africa: basin with Chlorogonium in Egypt (SCHMARDA 1854); pond, lake, rice field,
and ditch in East Africa and Madagascar (SONDHEIM 1929).
Records from saline inland waters and marine habitats (see also locus classicus):
shallow brackish pools at Leasowe, Chesire, England (GALLIFORD & WILLIAMS 1948); in
salt marshes of the Dee Estuary at Parkgate, Cheshire, England, in April (WEBB 1956);
salt-pit in Cagliari, Italy (PARONA 1883); saline lakes in Germany, Transylvania
(Romania), and Central Asia (ZACHARIAS 1888, ENTZ 1900, 1904a, GURWITSCH 1934);
Baltic Sea (EICHWALD 1849, 1852); White Sea (MERESCHKOWSKY 1877, CIENKOWSKY
1881); Logy Bay, Newfoundland (LACKEY & LACKEY 1970); Gulf of Mexico (SMITH
1904; see also BORROR 1962).
Not very common in sewage treatment plants like activated sludge (BANINA 1983,
CURDS 1969, 1975, KUTIKOWA 1984, SIMAKOV 1986, VERSCHAFFELT 1930) and trickling fil-
ters (CURDS 1975, RICHARDS & CUTLER 1933). OBERSCHMIDLEITNER & AEscHT (1996) found
S. pustulata 4-30 d after sampling in the activated sludge plant of the town of Linz, Up-
per Austria. LACKEY (1938a) counted 100 to 2000 indo ml-! in trickling filters and about
200 indo m!"! in activated sludge samples. SCHERB & BRAUN (1971) observed Stylonychia
pustulata in activated sludge supplied with water from intensive fish culture. PATIL et a1.
(1993) found it in stabilisation ponds in India with influent BODs of up to 300 mg l-!.
Records from terrestrial habitats unreliable although resting cysts are formed (BROD-
SKY 1928, EHRENBERG 1869, FELLERS & ALLISON 1920, NIKOUUK & GELTZER 1972,
TROITZKY & ZEREN 1926, VARGA 1936, 1961, YAKIMOFF & ZEREN 1924, 1926); very likely
confused with Sterkiella histriomuscorum, which is rather common in soil. According to
VARGA (1961), cysts of Stylonychia pustulata survived 8 years of dryness. WEISSE (1856)
reported encystment from l-year-old dry mud samples.
Feeds on bacteria, algae (Chlamydomonas reinhardii, Chlorogonium elongatum,
diatoms), amoebas, heterotrophic flagellates, and ciliates such as Colpoda, Cyclidium,
species of the Tetrahymena pyriformis complex, and zooids of small Vorticella (BEERS
1933, CURns & CURDS 1971, DAWSON et a1. 1983, GRIMES & ADLER 1976, LILLY & CEVAL-
LOS 1956, LILLY et at 1953, NOLAND 1925, PRATT & CAIRNS 1985, PROWAZEK 1899a,
WEBB 1956, YANO & SUHAMA 1990b). Stylonychia pustulata also attacks Paramecium
caudatum, and the digestion of flagellates takes about 1.5 h. PROWAZCEK (1899a) ob-
served that 7-15 specimens ofCyclidium sp. were ingested successively. Cannibalism oc-
curs under culture conditions at 340 cells ml-! and more (RICCI et a1. 1989a). Detailed in-
formation on the nutritional and supplementary growth factors is given by LILLY (1942)
and LILLY et a1. (1949, 1952, see also JOHNSON 1956, P 201). For culture they used a solu-
tion of Brewers' yeast; accordingly, living yeast cells, Euglena gracilis, Astasia klebsii,
584 SYSTEMATIC SECTION

Tetrahymena pyriformis, T. vorax, and Dexiostoma campylum, are adequate as food.

Glaucoma scintillans, Chilomonas paramaecium, and the algae Stichococcus bacillaris
and Chiarella sp. did not support growth of S. pustulata. REILLY & LILLY (1957) found
some differences between S. pustulata and S. putrina in the preferred food organisms;
both species were able to utilise the flagellates Polytom a uvella and P. agills. Stylonychia
pustulata also utilised Euglena gracilis and Dexiostoma campylum, whereas S. putrina
did not feed on these organisms, but did use Chilomonas paramaecium.
Sty/onychia pustu/ata is able to oxidise xanthine and it was found that it contains an
amylase system capable of the hydrolysis of starches from rice, potatoes, com, and glyco-
gens. The adenosinetriphosphatase was most active at a pH of about 7.6 at 30°C; phos-
phorylase was more active at pH 6.2 than at pH 7.2, 8.2, or 9.2 (HUNTER 1958, 1959,
1960, 1963a, b, 1964, 1965). SHAPIRO (1927) found that pH in the food vacuoles ranges
from pH 4.8 to 7.0. The time of complete digestion in giants of "Oxytricha bifaria" (see
taxonomy) is about 4 h (VERN! & ROSATI 1992). Sty/onychia pustulata is captured and
killed by bladders of Utricularia (HEGNER 1926) and by the suctorian Podophrya collini
(HULL 1961). Larvae of the fish Hypotha/michthys malitrix (silver carp), Aristichthys no-
bills (big head), and Ctenopharyngodon idella (grass carp) feed on S. pustulata (KORNIY-
ENKO 1971).
Rather easily to culture (BICK & KUNZE 1971, SUMMERS 1935). WIRNSBERGER et al.
(1985a) used tap water as a culture medium with yeast and a species of the Tetrahymena
pyriformis complex grown on dry egg yolk added as food. Others used tap water and a
few drops of milk (ZINGHER 1935, ZINGHER et al. 1932) or alfalfa hay infusion (ALEXANDER
1985, HUNTER 1958) as culture medium. OKA et al. (1980) found a generation time of24
h under the following conditions: 0.1 mM sodium phosphate (PH 7.1) at 25°C supple-
mented with the algae Chlorogonium e/ongatum as food; a 500 ml culture of S. pustuiata
(about 5 x 1()l cells mI·l ) was fed with about 5 x 109 cells of C. eiongatum per day, and
the medium was changed daily. Further culture methods and notes on nutrition and diges-
tion, see LILLY (1953, 1967), LILLY & CEVALLOS (1956), LILLY et al. (1949), LITTLE et al.
(1951), LUCCHESI & LILLY (1957), PROVASOU et al. (1958), RICCI et al. (1980b), RICE &
LILLY (1960), VAN WAGTENDONK (1955).
GREENLEAF (1926), DARBY (1930), and BEERS (1933) studied the influence of the vol-
ume of the culture medium and cell proximity on the rate of reproduction (see also KrrCH-
ING 1957a and ZAIKA 1970). DARBY (1929) found, in hay infusions, a higher division rate
at pH 8.0 (44 divisions per 10 days) than at pH 6.1-6.5 (31-33 division per 10 days).
Conversely, GAUSE (1936a, b) found no conspicuous difference in growth at pH 6.3 or
7.9. DASS & SAPRA (1965), DINI et al. (1975), RICCI (1981c), and Rup LAL & SAXENA
(1980) reported a generation time of about 8-9 h at 22°C and 28 DC; according to LILLY
(1942), divisions occurred at intervals from 8-12 h at about 26°C. SUMMERS (1935)
found a mean daily fission rate of 3.6 for 2 successive lO-day periods; the maximum
daily fission rate, using 3-day old media, was 5.0. BAITSELL'S (1911, 1912; see also RICH-
ARDS 1964) cultures of S. pustuiata reached a maximum of about 3.1 (beef extract
culture) and 2.4 (beef extract and hay infusion culture) divisions per day for a single 10-
 Stylonychia 585

Table 36 Autecological data of Sty/onychia pustu/ata. References: column 1, from BICK & KUNZE (1971;
summary of literature data, including FINLEY 1930, NOLAND 1925, REUTER 1961, and some dissertations); col-
umn 2, from DETCHEVA (1972a, 1975b, 1978, 1979a, c, 1982b, 1983a-c; many analyses from beta- and aI-
phamesosaprobic Bulgarian running waters); column 3, from PATRICK et aI. (1967; ? analyses from the Savan-
nah River, USA); column 4, from RlffiIVEN (1972) and CAIRNS & DICKSON (1972; ? analyses from the South
River in Virginia, USA); column 5, from CAIRNS & YONGUE (1973; unknown number of analyses in June and
September from the Cape Fear River near Fayetteville, North Carolina, USA); column 6, from MnwWWlTSCH
(1989; 29-33 analyses from salt polluted running ditches in Germany)

Parameterl Reference
2 3 4 5 6
Frequency (%) 0.9-3.6
Temperature (DC) 1-23 4-28 >8-23 23-26 3-20
pH 6.9-8.3 6.5-7.0 8.2 6.7-7.3 6.7-7.9
O2 (mg 1.1) 0.1-11.8 3.5-12 7-10 6.6-8.7 1.2-19.6
O2 (% saturation) 45-100
BODs (mg 1.1) 1.2-28 1.1 1-2.4
NH/-N (mg 1.1) 0.15-0.39 0.05-0.8 0.009-<1 0 0.005-0.68 0.Q7-1.3
NOi-N (mg 1.1) ()"'0.13 <0.007 0.002 0.01-0.8
NOi-N (mg 1.1) ()"'1.6 0.07-0.2 0.39 0.35-0.84 2.2-6.2
PO/-p (mg 1. 1) 0.01-0.033 0.08 0.09-0.28

I The figure "0" denotes only that the parameter in question could not be detected with the analytical method
used.

day period. MIDDLETON (1915a, b, 1918) recorded a maximum of about 3.2 divisions per
day for one lO-day period at 28-32 DC (further data on generation time, see CALKINS 1923
and MAUPAS 1888). Specimens which have been treated for 15 s with 5 rnA altemating-
current (i) increase temporarily in volume, (ii) often spin under treatment, and (iii) tend to
orient between the electrodes so that the current passes across the short body axis rather
than anteriorly-posteriorly (HUTclllNGS 1937,1946); furthermore, HUTClllNGS found no
statistically significant difference between experimental and control groups as regards the
total number of animals produced after such treatment.
Biomass of 106 individuals: 400-2000 mg (KORNIYENKO 1971, 1976; obviously
wrongly calculated or misidentification), about 80 mg (FOISSNER et al. 1991), 40 mg (NES-
TERENKO & KOVALCHUK 1991).
Autecological data are summarised in Table 36. Some further data: 0-22 mg I-I CO2
free, salinity up to 3.09 % (HICK & KUNZE 1971); 16-100 mg I-I Ca2+, 4.8-46 mg I-I Mgl+'

110-235 mg I-I HC03-, 35-116 mg 1.1 SOlo, 10-23 mg I-I Cl", 0.02-1 mg I-I Fe2+ (DETCHE-
VA 1975b, 1978, 1979a, c, 1982b, 1983a-c); 11-89 mg I-I COHee, 37-939 mg I-I CI-, 56
to 381 mS m-I specific conductivity (MnwLoWITSCH 1989); at 19.5 DC, pH 7.3, 51 ppm
total hardness, and 8 mg I-I O2 in the New River, Virginia, USA (YONGUE & CAIRNS
1979). GITTLESON & FERGUSON (1971) recorded S. pustulata from a slow-moving stream
with leaf-type debris at a temperature of 2-3 DC. Low temperature (10 ± 1 DC) for 7 days
caused high mortality; the survivors exhibited erratic binary fission during the course of
586 SYSTEMA TIC SECTION

treatment resulting in daughter cells with abnonnal body shape and nuclear apparatus;
some of them remained as doublets (SHIVAJI et al. 1976). MATONlcKIN (1957) found S.
pustulata in thennal waters in Yugoslavia at 42.2 °C. The same temperature was used by
YANO & SUHAMA (1991) to obtain doublets by a heat shock treatment consisting of incu-
bation at 42°C for 10 min and at room temperature for 1 min.
STROM et al. (1974) recorded S. pustulata in an experiment on the purification of ef-
fluents from the production of alkylphenol additives. 50 and 100 ppm DDT inhibited
growth and cell division (sterilised hay infusion, 22°C, Klebsiella aerogenes; Rup LAL &
SAXENA 1980); in specimens exposed for 6 h to 100 ppm DDT, synthesis of RNA and
protein was inhibited throughout the cell cycle (SAXENA & Rup LAL 1981); maximum con-
centration of total DDT residue was obtained within I h and accumulation factor varied
from 97 to 295 (RUp LAL et al. 1981). For cadmium accumulation, see Table 18. Of 14
acids tested, salicylic and hydrochloric were most toxic, while valeric and butyric are
among the least toxic (COLLETT 1919). For the toxicity of dyes and arsenic acid, see FER-
REIRA & PEREZ (1937) and ZACHARIAS (1902b). Supersonic waves with a frequency of
3 MHz and an intensity of 2.18 W cm-2 killed S. pustulata within I min (ZrvANoVIc
1959).
The saprobic classification is not unifonn. Originally, S. pustulata was classified as
oligosaprobic indicator (KOLKWITZ & MARSSON 1909, KOLKWITZ 1950, see also MAUCH
1976). KARL (1932) designated it as katharobic to moderately saprobic. This classifica-
tion is supported, for example by the record of KOHN (1950), who found S. pustulata in
springs near Vienna, Austria. However, STEIN (1859b) observed it mainly in distinctly or-
ganically polluted habitats. According to SLADECEK et al. (1981) and WEGL (1983), S. pus-
tulata is confmed to betamesosaprobity: b; b = 10, I = 5, SI = 2.0. ForssNER (1988) classi-
fied it at as follows: b; b = 9, a = 1, I = 5, SI = 2.1. Considering the rather different habi-
tats mentioned above, ForsSNER et al. (1991) proposed the following classification: b; 0 =
1, b = 7, a = 2, 1=4, SI = 2.1.

Slylonychia putrina STOKES, 1885 (Fig. 165a-d, h, Table 3)

1885 Sty/onychia putrina STOKES, Am. mono microsc. 1., 6: 187 (Fig. 165a).
1888 Sty/onychia putrina, STOKES - STOKES, 1. Trenton nat Hist Soc., 1: 298 (Fig. 165a).
1932 Sty/onychia putrina STOKES, 1885 - KAlIL, Tierwelt Dtl., 25: 619.
1963 Sty/onychia putrina STOKES - LUNDIN & WEST, Free-living protozoa, p 68 (Fig. 165h; see also WEST &
LUNDIN 1963).
1969 Stylonychia putrina - GROLIERE, Annis Stn limnol. Besse, 4: 347 (Fig. 165c, d).

Fig. 165a-d Sty/onychia putrina (a, from STOKES 1885a; b, from CURDS 1969; c, d, from GROLIERE 1969. a, b, ~
from life; c, d, protargol impregnation). a, b) Ventral views, a = 125 ~m, b = 150 ~m. Note distinctly separate
caudal cirri; probably an important difference to S. pustutata. c) Ventral infiaciliature, size not indicated. Ar-
row marks posterior postoral ventral cirrus. d) Ventral infraciliature of middle morphogenetic stage, size not
indicated. Arrow denotes left frontoventral cirrus which is inconspicuously separate from the other frontoven-
tral cirri both in the protargol preparations by GROLIERE and in the type population (see Fig. a).
 Sty/onychia 587

.....
'l'Il'I/

It. ~
d~ .. Q

~
~
~
~
~

;;::::

~
~
§!§

f)
~..
a

',_
"
..I. '
Q 0 0
.-
.-
.-
.-

..
•
•

-9

••
•.'•• ..,
..
"... .
" ~

c 9

Fig. 165e-g Sty/onychia pustulata (e, f, from DRAGESCO & DRAGESCO-KERNEIS 1986; g, after PATSCH 1974
from DRAGESCO & DRAGESCO-KERNEIS 1986. e-g, protargol impregnation). e-g) Ventral infraciliature, e =
80 jlm, f= 100 jlm, g = size not indicated; p 565.
588 SYSTEMA TIC SECTION

1969 Sty/onychia putrina STOKES, 1885 - CURDS, Wat. Poll. Res., No. 12: 70 (Fig. 165b).
1991 Sty/onychia putrina STOKES, 1885 - FOISSNER, BLATTERER, BERGER & KOHMANN, Infonnationsberichte
des Bayer. Landesamtes filr Wasserwirtschaft, 1/91: 329.

Nom e n c I at u rea n d t a x 0 nom y: KAHL (1932) classified Sty/onychia as sub-

genus of Oxytricha; the correct name in his revision is thus Oxytricha (Sty/onychia) pu-
trina. "Sty/onychia putrinum" in PROVASOLI et al. (1958) is an incorrect subsequent spell-
ing.
Non-dividing specimens of S. putrina are very similar to S. pustu/ata; however, the
validity of S. putrina is beyond doubt because the cyst wall is smooth and not spiky as in
S. pustu/ata; furthermore, morphogenesis commences with the proliferation of basal bod-
ies close to the left transverse cirrus (Fig. 165d), and not in the area left of the postoral
ventral cirri as in S. pustulata (Fig. 161e). Stylonychia vorax and S. curvata are also
rather similar (see key).
Although the illustration by LUNDIN & WEST (1963; Fig. 165h) is somewhat superfi-
cial, the identification can be accepted. Sty/onychia putrina sensu BARY (1950) does not
match the original description very well, which was already stated by BARY (1950) him-
self; it is classified as Sty/onychia pustu/ata (Fig. 163d). The populations described by
DRAGESCO & NJINE (1971) are classified as species of the Sty/onychia mytilus complex
("grande variete"; Fig. 159g) and Sty/onychia notophora (''petite variete"; Fig. 166d), re-
spectively, because the transverse cirri are arranged in two distinctly separated groups.
Sty/onychia putrina sensu DRAGESCO & DRAGESCo-KERNEIS (1986; Fig.
165e, f) and PArsCH (1974; Fig. l65g) are classified as Sty/onychia
pustu/ata, because the ventral infraciliature is much more similar to
that of Sty/onychia pustu/ata sensu WIRNSBERGER et al. (1985a) than to
that of Sty/onychia putrina sensu GROLIERE (1969). The redescriptions
by BARWICK et al. (1955; Fig. 223h), BURKOVSKY (1984; Fig. 233k),
CONN (1905; Fig. 228e), SAMANO & SOKOLOFF (1931; Fig. 24li), and
WANG (1925; Fig. 242f) are insufficient. Detailed redescription neces-
sary.

Morphology and biology:InlifeaboutI20-150~mlong

and three times as long as broad. Outline usually parallel sided; how-
ever, sometimes anterior end slightly broader than posterior one; both
ends usually broadly rounded. Cytoplasm often filled with dark gran-
ules. Rapid and erratic movement. Adoral zone of membrane lIes
40-45 % of body length, 50-55 membranelles. The 4 right transverse
cirri protrude slightly to distinctly beyond posterior end of cell. 45-53
Fig. 165h Stylo-
nychia putrina right and 28-32 left marginal cirri (GROLIERE 1969). Caudal cirri
(from LUNDIN & about 15 % of body length, rather distinctly separate. Formation of
WEST 1963). Ven- oral primordium commences adjacent to left transverse cirrus (GRO-
tral view from LIERE 1969; Fig. 165d). Cyst wall smooth (STOKES l885a). DNA con-
life, 125-150 J.Im;
p 586.
 Stylonychia 589

tent of nuclei and cellular protein content, see AMMERMANN & MUENz (1982; identifica-
tion not substantiated by an illustration; see also RAIKov 1989).

o c cur r e n c e and e colo g y: In the aufwuchs and detritus oflentic and lotic wa-
ters with decomposing plant material; much less frequent than S. pustulata. Rare in sew-
age treatment plants.
Locus classicus is a stale vegetable infusion; STOKES (1885a) did not give the specific
locality where he found S. putrina, however, he lived and worked in Trenton, New
Jersey, USA, and it is thus assumed that the locus classicus is nearby. LUNDIN & WEST
(1963) collected it from various freshwater habitats in the Upper Peninsula of Michigan,
USA. GROLIERE (1969) found S. putrina between decaying aquatic plants in Lake Cham-
bou, France.
Records not substantiated by illustrations: beta- to alphamesosaprobic Traun River in
Upper Austria (FOISSNER & MOOG 1992); brook in Upper Austria (BLATIERER 1994);
Maritza River, Bulgaria (DETCHEVA 1981); benthal of the Bulgarian Ossam River during
fall (RUSSEV et al. 1984); in small numbers «10 indo ml-I ) in 2.5 % of 178 sites of sewage
fungus outbreaks in England (CuRTIS & CURDS 1971); mud of Loch Leven, England
(BRYANT & LAYBOURN 1974); rare in Neckar River, Germany (BUCK 1959, 1961); rare in
an alphamesosaprobic brook in the southern Black Forest, Germany (BAUER 1987); small
lakes near MUnster, Germany (NUSCH 1970); aufwuchs in mesotrophic lakes in Germany
(MOCKE 1979, PACKROFF 1992, PACKROFF & WILBERT 1991); cooling system of a power
station at the beta- to alphamesosaprobic Main River, Germany (BERNERTII 1982); about
400 indo ml- I in experiments with decomposition of cellulose (BICK 1967b, 1973, Voss
1978); unpolluted foothill stream (Breitenbach) near the village of Schlitz, Germany
(PACKROFF & ZWICK 1996); Tisza River in Hungary (STILLER 1942); well in Italy (GRISPINI
1938); 8 indo cm-2 in the mesosaprobic Baganza River and in a shallow macrophyte cov-
ered basin near Reggio Emilio, Italy, fed with sewage from a pig farm (MAnoN! & GHETTI
1977, MAnoN! & VIAROLI 1985); alphamesosaprobic areas of the Save River and al-
phameso- to polysaprobic zone of a mountain brook in Yugoslavia (PRIMc 1981, 1984);
up to 160 indo }-I in the neuston of a eutrophic reservoir near Barcelona, Spain (SALVADO
CABRE 1993, SALVADO & GRACIA 1991; further records from Spain: FERNANDEZ-LEBORANS
et al. 1990, FINLAY et al. 1993, MARGALEF-li>PEZ 1945); Azerbaijan (Al.EKPERov 1986);
polysaprobic to alphamesosaprobic puddles and in rivers in India (KAUR & MEHRA 1998,
KruSHNAMOORTIll 1969, RANGANATHAN & SHREEDHARAN 1986); rare in small clear pools
with much decomposing organic sediment, USA (HAUSMAN 1917; determination uncer-
tain because only 50 ~m long); Iowa, USA (SHAWHAN et al. 1947); rivers in eastern
United States with soft and medium hard water (PATRICK 1961); Conestoga drainage ba-
sin, Pennsylvania, USA (CAIRNS 1965a); Mexico (LOPEZ-OcHOTERENA & ROURE-CANE
1970); tributary of the Amazon River, Colombia (BOVEE 1957); with a frequency of9 %
in a pond with leaf litter in Cameroon (NJINE 1977).
GRAY (1952) found Stylonychia putrina in a soil surrounding a brook. No further ter-
restrial records available.
590 SYSTEMATIC SECTION

Sty/onychia putrina is an oIigo- to meso-euryhaline species with freshwater optimum,

that is, tolerates 5 g I-I Cl- and 14 g 1-1 salt (ALBRECHT 1983, 1984, 1986, BICK 1967a,
1972a). BICK (1968b) counted about 750 indo ml- I during the decomposition of cellulose
in brackish water with 0.7 % salinity. DZHURTUBAYEV (1977) found 3 indo cm-2 in the up-
per sediment layer (0-1 cm) of the Odessa Bay (Black Sea), and RAMAN et al. (1993) and
KALAVATI et al. (1995) recorded S. putrina with a frequency of 10 % in a polluted harbour
on the east coast of India.
Sty/onychia putrina occurs also in sewage treatment plants such as activated sludge
(ANrONIETII et al. 1982, BANINA 1983, CURDS 1969, CURDS & COCKBURN 1970a, Ku-
TIKOWA 1984, MACEK 1987, 1989, 1991, MAnoNI 1981, 1988a, MAnoNI & GHETIl 1981b,
STAMM 1979, 1984), rotating biological contactors (MAnoNI & GHETIl 1981b), stabilisa-
tion ponds with 50-80 mg I-I BODs (KruSHNAMOORTHI & SARKAR 1981), and other sys-
tems (BEGER 1935). RIVERA et al. (1987) counted 37 000 indo mI-1 bottom sample from a
waste water stabilisation pond in Mexico at 11-17 DC and pH 7.1-8.7. Not recorded from
percolating filters and Imhoff tanks (CURDS 1975). VARGA (1953) found S. putrina in fer-
mented farmyard manure.
Feeds on bacteria, diatoms, filamentous growths, heterotrophic flagellates (Chi/omo-
nas paramaecium), and ciliates (BICK 1972a, CURTIS & CURDS 1971, MACEK 1989, REll..LY
& LILLY 1957). Sty/onychia putrina does well in a monoxenic culture of sterile lettuce ex-
tract plus K2HP04 and KH2P04 (PH 6.8-7.2) and a species of the Tetrahymena pyri-
formis complex as food (PRovASOLI et al. 1958). DOWNS (1952, 1956, 1959), studying the
mating types in S. putrina, used a species of Chlamydomonas and Aerobacter cloacae as
food. REll..LY & Lll..LY (1957) found some differences between S. putrina and S. pustu/ata
concerning the food organisms (see S. pustufata). Duration of cell cycle about 12 h at
23 DC in Pringsheim medium and with Ch/orogonium e/ongatum as food (AMMERMANN &
MUENZ 1982). Biomass of 106 individuals about 68 mg (FOISSNER et al. 1991). SCHMEREN-
BECK (1975) found S. putrina at flowing velocities of up to 0.6 m S-I.
Data on the autecology and saprobiology of S. putrina have to be used with some
care because it cannot be excluded that it was sometimes confused with the rather similar
and more common S. pustulata (Table 35). Further autecological data: pH 6.5-7.0 (opti-
mal range), 0.1-1 mg 1-1 O2 (optimal range), 0-56 mg I-I CO2 free, 0-2 mg I-I free NH3,
1000-20000000 bacteria mI-1 (plate counts on peptone agar; BICK & KUNZE 1971, BICK
1972a); 0.9 % frequency, 41-60 mg I-I Ca2+, 7-38 mg 1.1 Mgl+, 116-214 mg I-I HC03-,
19-113 mg I-I S042-, 10-20 mg I-I Cl-, 0.03-0.05 mg I-I Fe2+ (DETCHEVA 1983c). REiN-
NARTH (1979) found S. putrina in a eutrophic pond in the upper layer of a mud-water in-
terface at pH 7.1, +181 mV redox potential, 0.13 mg I-I NH/-N, and 1.2 mg I-I N03--N.
Cysts are killed at temperatures between 40 and 45 DC (MOL1MANN 1978). NUSCH (1970)
counted about 40 indo cm-z at 10 DC in an aquarium three weeks after addition of
100 mg I-I peptone. GREISER (1974) found S. putrina in laboratory ecosystems in areas
with low oxygen content and high ~+-concentrations together with Dexiostoma campy-
fum, Glaucoma scintillans, Tetrahymena pyriformis, Trimyema compressum, and Vorti-
cella microstoma.
 Stylonychia 591

In agreement with the faunistic and ecological data, S. putrina is classified as indica-
tor of alphamesosaprobic conditions: a; b = 2, a = 7, P = 1, I = 3, Sl = 2.9 (FOISSNER
1988a, FOISSNER et al. 1991, SLADECEK et al. 1981, WEGL 1983). However, this has to be
verified on reliably determined populations.

Stylonychia vorax STOKES, 1885 (Fig. 159b, C, 166a, b, e)

1885 Stylonychia vorax STOKES, Am. mono micros. 1., 6: 188 (Fig. 166a).
1888 Stylonychia vorax, STOKES - STOKES, J. Trenton nat. Hist. Soc., 1: 298 (Fig. 166a).
1932 Stylonychia vorax STOKES, 1885 -KAHL, TielWe1t Dt!., 25: 619.
1957 Stylonychia vorax STOKES 1885 - SRAMEK-HuSEK, V~st. ~S!. zoo!. Spo!., 21: 23 (Fig. 166b).
1966 Stylonychia ovalis DRAGESCO, Protisto1ogica, 2: 93 (Fig. 159b, c).
1973 Stylonichia vorax STOKES, 1885 - MADRAZo-GARIBAY & L6PEZ-OcHOTERENA, Revta Soc. mex. Hist. nat.,
34: 66 (Fig. 166e; incorrect subsequent spelling of Stylonychia).
1991 Stylonychia vorax STOKES, 1885 - FOISSNER, BLATTERER, BERGER & KOHMANN, lnfonnationsberichte des
Bayer. Landesamtes filrWasselWirtschaft, 1191: 334.

Nom e n c I at u rea n d t a x 0 nom y: KAHL (1932) classified Stylonychia as sub-

genus of Oxytricha; the correct name in his revision is thus Oxytricha (Stylonychia) vo-
rax, which is a secondary homonym of Oxytricha (Histrio) vorax in KAHL (1932, p 617).
Clara vorax (STOKES, 1885) in EIGNER (1997, P 568) is a new combination with the junior
homonym Clara EIGNER, 1997 (details see Stylonychia pustulata). This transference is
based on the morphogenetic data of S. vorax sensu WIRNSBERGER et al. (1985a), which is,
however, a Stylonychia hi/aria.
The populations described by STOKES (1885a), SRAMEK-HUSEK (1957), and DRAGESCO
(1966b) agree very well in the slightly tapered posterior end, the long, rather narrowly ar-
ranged caudal cirri, and the transverse cirri which are not separated in two groups. Espe-
cially the more or less pointed posterior end seems to exclude synonymisation with the
rather similar Stylonychia pustulata. Thus, S. pustulata sensu LILLY (1942, Fig. 162j)
could be S. vorax. Synonymy of Stylonychia. vorax and S. ovalis was suggested by BOR-
ROR (1972a). Detailed redescription necessary. Difficult to distinguish from Stylonychia
pustulata, S. putrina, and S. curvata!

M 0 r p hoi 0 g y: In life about 71-85 x 43 ~m, according to DRAGESCO (1966b) after

protargol impregnation 11 0 ~m long on average, about 2.3 times as long as broad. Ellip-
tical or obovoid, posterior end rather narrowly rounded or even pointed (Fig. 159b, c,
166a, b). Two micronuclei.
41-60 adoral membranelIes. Frontal and transverse cirri distinctly enlarged; all trans-
verse cirri protrude beyond posterior end of cell. According to DRAGESCO (1966b) 30-36
left and only 23-25 right marginal cirri; however, in his illustration (Fig. 159b) 36 right
marginal cirri are shown. Marginal rows only slightly separated posteriorly. Caudal cirri
stout, about 28 % of body length, rather narrowly spaced and, after STOKES (1885a), they
 592 SYSTEMATIC SECTION

-
,
,
,
:~ ~.~ .... ~
~
, \1.$ ,
,
, .-:.IVl2 '
:
'.. . ,
V/4u~

.
~
~

I
I

,
,
,
,
,

Fig. 166a, b, e Sty/onychia vorax (a, from STOKES 1885a; b, after SRAMEK-HuSEK 1957; e, from MADRAZo-
GARIBAY & L6PEZ-OcHOTERENA 1973). a, b, e) Ventral views from life, a = 125 Ilm, b = size not indicated, e =
71 Ilm; p 591.
Fig. 166c, d Sty/onychia notophora (c, after DRAGESCO & NJINE 1971 from DRAGESCO & DRAGESCo-KERNEIS
1986; d, from DRAGESCO & NJINE 1971). c, d) Ventral infraciliature, protargol impregnation, c = 80 Ilrn, d =
100 Ilm. In S. notophora postoral ventral cirrus IVI2 is placed more anteriorly than postoral ventral cirrus V/4;
a conspicuous difference to species of the S. mytilus complex, where cirrus V/4 is placed more anteriorly (see
Fig. 149a); p 553.

arise directly from posterior end and not from the dorsal surface, as for instance in S.
myti!us, S. notophora, and S. putrina.

o c cur r e n c e and e colo g y: Probably confined to freshwater, obviously prefer-

ring lentic waters. Locus classicus is a shallow pond, where STOKES (1885a) discovered S.
vorax in early spring. STOKES did not give the specific locality where he found it;
 Stylonychia 593

however, he lived and worked in Trenton, New Jersey, USA, and it is thus assumed that
the locus classicus is nearby. Locus classicus of the synonym, S. ovalis, is a freshwater
habitat near Thonon-Ies-Bains, Lake Geneva, France (DRAGESCO 1966b). KAlIL (1932)
did not fmd S. vorax. SRAMEK-HuSEK (1956b, 1957, 1958) isolated S. vorax from a beta-
mesosaprobic habitat of the Moravice River system, Czechoslovakia. MADRAZO-GARlBAY
& L6pEZ-OCHOTERENA (1973) found it in the San Anton waterfall in Morelos State, Mex-
ico.
Records not substantiated by illustrations: spring area of the Melbbach «1 mg 1-1
BODs), a small brook near Bonn, Germany (JUTRCZENKI 1982); brook in Hungary
(VOROSVMY 1950); urban runoff of Hickey Run, District of Columbia, USA (SAYRE et al.
1986, as "Stylonychia ovale"); soil of the Apsheron peninsula, Azerbaijan (AGAMALIEV &
MUSAYEV 1988).
Voracious predator on flagellates (STOKES 1885a; the remarkable appetite suggested
the species name), but also feeds on bacteria and diatoms (VOROSVMY 1950); also canni-
balistic. Biomass of 106 individuals about 57 mg (FOISSNER et al. 1991). WENRICH (1918)
reported S. vorax impaled on a fungal filament; many specimens were located at points
basal to branches of the filaments, indicating that they had been held prisoner for some
time.
SLAoECEK et al. (1981), WEGL (1983), FOISSNER (l988a), and FOISSNER et al. (1991)
classified S. vorax as betamesosaprobic indicator of water quality: b; b = 10, I = 5, SI =
2.0. This strict classification has to be verified by further autecological and saprobiologi-
cal data on reliably determined populations.

Stylonychia curvata (KARL, 1932) BORROR, 1972 (Fig. 167f, 234m)

1932 Sty/onychia curvata KAHL, Tierwelt OtI., 25: 619 (Fig. 167t).
1971 Sty/onichia curvata KARL - CHARDEl, Revue verviet. Hist. nat., 28: 38 (Fig. 234m; incorrect subsequent
spelling of Sty/onychia).
1972 Sty/onychia curvata KARL, 1932 - BORROR, J. ProtozooI., 19: 15.

Nom e n cIa t u rea n d t a x 0 nom y: KAlIL (1932) classified Stylonychia as sub-

genus of Oxytricha; the correct name in the original description is thus Oxytricha (Styla-
nychia) curvata. Although BORROR (l972a) did not use the phrase "comb nov." as he did
in other cases, I consider him as combining author. Stylonychia curvata sensu GIESE &
ALDEN (1938) does not have the characteristic oval outline and has only inconspicuous
caudal cirri; it is classified as Sterkiella histriomuscorum (Fig. 186t). The illustration in
CHARDEZ (1971) looks like a redrawing from KAlIL (1932; cpo Fig. 167f, 234m). Detailed
redescription necessary. Difficult to distinguish from S. pustulata, S. putrina, and S. va-
rax.

M 0 r p hoi 0 g y: In life about 60-80 11m long, about 2.5 times as long as broad. Dis-
tinctly ovoid, that is, posteriorly more broadly rounded than anteriorly, anterior portion
 594 SYSTEMATIC SECTION

Fig. 167a Stylonychia pseudograndis (after WANG & NIE 1935). Ventral view from life, 170 )lm; p 564.
Fig. 1671HJ Stylonychia stylomuscorum (b, after SRAMEK-Hu~EK 1957; c, d, from KAm. 1932). Ventral views
from life, b = size not indicated, c, d = 80 )lm. Note single micronucleus between macronuclear nodules;
p 595.
Fig. 167e Stylonychia pusilla, a supposed synonym of S. notophora (from KAm. 1932). Ventral view from
life, 85 )lm; p 555.
Fig. 167f Stylonychia curvata (from KAm. 1932). Ventral view from life, 80 )lm; p 593.

slightly bent to the left. Adoral zone of membranelles about 40 % of body length. Ante-
rior part of undulating membranes possibly curved to the left, indicating that it possible
belongs to another genus. Transverse cirri protrude distinctly beyond posterior end of
cell. Caudal cirri about 30 % of body length.

o c cur r e n c e and e colo g y: Probably confmed to freshwater habitats. Locus

typicus not fIXed; KAliL (1932) discovered Sty/onychia curvata with high abundance in a
trench containing decaying plants, probably somewhere in Germany. CHARDEZ (1971)
found it in the Ardennes, Belgium. Further records: lentic waters of the Tibetan Plateau
(WANG 1977); Yuelushan Area, China (YANG 1989); soil from paddy field in Japan
(TAKAHAsm & SUHAMA 1991).
PATRICK et al. (1967) found Sty/onychia curvata, rarely, in the Savannah River, USA,
at 20-23 DC, pH 6.5-7.0, 7-9 mg 1- 1 0 2 , 1-<5 mg I-I BOD5 , 5-<10 mg I-I Cl-, 0.001 mg I-I
 Stylonychia 595

NH3-N, <0.007 mg I-I N02--N, 0.2-<0.7 mg I-I N03--N, 0.017-0.033 mg I-I POl--P,
>1-10 mg I-I S043-.
Temporary fasting can result in the formation of giant cannibalistic individuals (TUF-
FRAU 1965b). SEN et al. (1967) cultivated Stylonychia curvata in a bacterised proteose-
peptone-cerophyl-wheat broth diluted 1:18 (v/v) with distilled water; synchronous cell di-
vision was induced by exposure to low temperature.

Stylonychia stylomuscorum (FOISSNER, BLATIERER, BERGER & KOHMANN,

1991) FOISSNER, BLATIERER, BERGER & KOHMANN, 1991 (Fig. 167b-d)

1932 Sty/onychia muscorum KAHL, Tierwelt DtI_, 25: 619 (Fig. 167c, d).
1957 Sty/onychia muscorum KAHL 1932- SRAMEK-HuSEK, vtst. ~I. zool. Spol_, 21: 23 (Fig. 167b).
1972 Sty/onychia muscorum KAHL, 1932 - BORROR, 1. Protozool., 19: 15_
1991 Oxytricha sty/omuscorum FOISSNER, BLATIERER, BERGER & KOHMANN, Informationsberichte des Bayer_
Landesamtes ftlr Wasserwirtschaft, 1/91: 332.
1991 Sty/onychia sty/omuscorum (FOISSNER, BLATIERER, BERGER & KOHMANN, 1991) - FOISSNER, BLATIERER,
BERGER & KOHMANN, Informationsberichte des Bayer. Landesamtes ftlr Wasserwirtschaft, 1/91: 332.

Nom e n c I at u rea n d t a x 0 nom y: I<AHL (1932) classified Stylonychia as sub-

genus of Oxytricha; the correct name in the original description is thus Oxytricha (Stylo-
nychia) muscorum, which is a primary homonym of 3 other Oxytricha muscorum estab-
lished by I<AHL (1932). For further details on this nomenclatural problem, see the relevant
chapter in Sterkiella histriomuscorum.
The most important species character of Stylonychia stylomuscorum is the single mi-
cronucleus between the 2 macronuclear nodules. Thus, identity with the rather similar
and very common Sterkiella histriomuscorum can be excluded because this species al-
ways has more than 1 micronucleus. ALONSO & PEREZ-SILVA (1963a, b) described 2-4 mi-
cronuclei and CHARDEZ (1967; Fig. 233j) and MATIS & DANISKOvA (1972; Fig. 230f, g)
did not mention the number; consequently, these identifications can not accepted. Latter
records by MATIS, however, are listed here. Detailed redescription necessary.

M 0 r p hoi 0 g y: In life 60-100 J-lm long, about 2.1 times as long as broad. Elliptical,
slightly converging posteriorly, or even with parallel margins, posterior end rounded.
Contractile vacuole in about mid-body or slightly in front of it. Transverse cirri not (Fig.
167d) to distinctly enlarged (Fig. 167b, c) and possibly arranged in 2 groups (Fig. 167c),
indicating that perhaps 2 closely related species exist. Transverse cirri protrude slightly
(left) to distinctly (right) beyond posterior end of cell. Caudal cirri only slightly longer
than marginal cirri (Fig. 167c, d).

o c cur r e n c e and e colo g y: In terrestrial habitats (mosses, soil) and in the auf-
wuchs and detritus of freshwater. Locus classicus not fIXed. I<AHL (1932) discovered S.
stylomuscorum in mosses from Europe and North America. SRAMEK-RuSEK (1953, 1956b,
596 SYSTEMATIC SECTION

1957) observed it in one alphamesosaprobic and seven betamesosaprobic waters in

Czechoslovakia. Records not substantiated by illustrations: with a frequency of 19 % in
submerged and 12 % in wet mosses in Czechoslovakia (TIRJAKOVA & MATIS 1987a; see
taxonomy); pond and fishponds in Czechoslovakia and Poland (HAsSDENTEUFELOvA-
MORAvcovA 1955; MATIS 1973, see taxonomy; SIEMINSKA & SIEMINSKA 1967); agricultural
soils in Czechoslovakia (TIRJAKovA 1988; further records from Czechoslovakia: MATIS &
TIRJAKovA 1994, TIRJAKOVA 1992b, 1993); various, mostly heavily polluted rivers in
NordwUrttemberg, Germany, once highly abundant in the Fils River (BuCK 1959, 1961);
among detritus and in the surface film of ditches regularly contaminated with pesticide
sprays and in unsprayed ditches in the Elbe flood plain, Germany (CASPERS & HECKMAN
1982); cooling towers of power stations in Germany (SCHARF 1961); aufwuchs and
detritus-drifts in the littoral of Lake Balaton, Hungary (GELLERT & TAMAs 1958, 1959a,
1960, TAMAs & GElLERT 1958); soil under mosses in Hungary (GELLERT 1956); freshwa-
ter in Italian caves (BoscoLO 1968, GITILESON & HOOVER 1969, STEllA & SALVADORI
1953); in the bottom ofa fish pond in Poland (KWIATKOWSKA-GRABACKA 1965); freshwa-
ter habitats in the Tibetan Plateau (WANG 1977); freshwater in Yuelushan Area, China
(YANG 1989); soil from paddy fields in Japan (TAKAHASID & SUHAMA 1991).
Feeds on algae (including diatoms) and heterotrophic flagellates (GELLERT 1956,
GELLERT & TAMAs 1958, TAMAs & GELLERT 1958). Biomass of 106 individuals about
30 mg (FOISSNER et al. 1991). According to the experiments ofPRIMc (1983) on the influ-
ence on water temperature, S. stylomuscorum was present only at about 20°C. SCHARF
(1961), however, reported an upper temperature limit of 36 °C. SRAMEK-HuSEK (1954)
found S. stylomuscorum scattered in grey, moderately saprobic depositions at 5.4 °C and
pH 7.1, and sporadically in the littoral sediments of the March River, Czechoslovakia.
MATIS (1975; identification uncertain, see taxonomy) observed it in road pools and run-
ning waters near Bratislava at 3-28.5 °C, pH 5.8-6.9, and 1.2-12.4 mg 1·\ O2• MATIS &
STRAKOVA-STRIESKOvA (1991) recorded Stylonychia stylomuscorum from a brook and two
thermal lakes in Czechoslovakia at 7-40.5 °C and pH 6.0-7.3.
According to SRAMEK-HuSEK (1958, see also MAUCH 1976) typical for betamesosa-
probity. SLAnECEK et al. (1981), WEGL (1983), FOISSNER (1988a), and FOISSNER et al.
(1991) give the following values: b; b = 10, 1= 5, SI = 2.0. The restriction to betamesosa-
probity has to be verified by further autecological and saprobiological data on reliably
determined specimens, because BUCK (1959, 1961) found S. stylomuscorum in various
heavily polluted rivers and thus classified it as alphamesosaprobic.

Species transferred to a genus not treated in this book

Stylonychia appendiculata EHRENBERG, 1838, Infusionsthierchen, p 373, Tafl. XLII, Fig.

III. Remarks: Now Diophrys appendiculata. Marine.
 Sty/onychia 597

Species indeterminata

DUMAS (1929, 1930, 1937), a French amateur protistologist, described about 550 new
species, mainly ciliates from various habitats of the French Massif Central. Very likely,
most of the about 80 hypotrichous species, mainly assigned to Kerona, Oxytricha, and
Sty/onychia, are species indeterminata. A list of DUMAS' names is provided by FOISSNER
(1995a).

Ceratidium cuneatum EHRENBERG, 1838, Infusionsthierchen, p 367, Tafel XLI, Fig. VI.
Remarks: Ceratidium EHRENBERG, 1838, was established for a small (50--60 flm) organ-
ism, probably a fragment of a hypotrichous ciliate. FOISSNER & FOISSNER (1988) consid-
ered it a synonym of Stylonychia mytilus. EHRENBERG (1838) found it only once in 1820
in ponds near Berlin. EICHWALD (1844) and WEISSE (1848c) apparently observed this form
in Russia. However, these records are not substantiated by illustrations. CORLISS (1979;
P 208) considered Ceratidium as nomen oblitum.

Hypotrichen (nov. spec.?) - FORTNER, 1934, Arch. Protistenk., 81: 284 (Fig. 240h). Re-
marks: In life 225-270 flm long and only one macronucleus 50-60 flm across; possible a
post-conjugant of a large Stylonychia. In the surface layer of a saprobic infusion (Prague,
Czechoslovakia).

Stylonichea bistyllus DUMAS, 1930, Microzoaires, Planche XXX, fig. 4 (Fig. 238n). Re-
marks: Incorrect subsequent spelling of Stylonychia. Without description.

Stylonichia anfracta DuMAs, 1929, Microzoaires, p 84 (Fig. 238a). Remarks: Incorrect

subsequent spelling of Stylonychia.

Stylonichia auriformis DUMAS, 1937, Microzoaires, p 25 (Fig. 240a). Remarks: Incorrect

subsequent spelling of Stylonychia. See also S. virgula.

Stylonichia bicaudatus DUMAS, 1937, Microzoaires, p 27 (Fig. 240c, d). Remarks: Incor-
rect subsequent spelling of Stylonychia.

Stylonichia capucinus DuMAs, 1930, Microzoaires, Planche XXIX, fig. 1 (Fig. 238m).
Remarks: Incorrect subsequent spelling of Stylonychia. Without description.

Stylonichia corniculata DUMAS, 1929, Microzoaires, Planche XXXI, fig. 5 (Fig. 238d).
Remarks: Incorrect subsequent spelling of Stylonychia. Without description.

Stylonichia cornifrons DUMAS, 1929, Microzoaires, p 85 (Fig. 238b). Remarks: Incorrect

subsequent spelling of Stylonychia.
598 SYSTEMATIC SECTION

Sty/onichia dupla DUMAS, 1929, Microzoaires, p 82 (Fig. 237g). Remarks: Incorrect sub-
sequent spelling of Stylonychia dupla (legend to Planche XXIX, fig. 3). A middle mor-
phogenetic stage.

Stylonichia ellipsoides DUMAS, 1929, Microzoaires, p 84 (Fig. 237j). Remarks: Incorrect

subsequent spelling of Stylonychia (in the legend to Planche XXX, fig. 7 designated as
"Stylonichia pustulata"). The shape is reminiscent of S. pustulata, whereas the widely
spaced and very long caudal cirri are typical for species of the S. mytilus complex.

Stylonichia excavata DUMAS, 1930, Microzoaires, p 61 (Fig. 2381). Remarks: Incorrect

subsequent spelling of Stylonychia. The outline is reminiscent of Psilotricha (see FOISS-
NER 1983b).

Stylonichia jastigiata DUMAS, 1929, Microzoaires, p 83. Remarks: Incorrect subsequent

spelling of Sty/onychia jastigiata (legend to Planche XXIX, fig. 9). Two elongated cau-
dal cirri. Anterior end with a small process; possible an opisthe of a Stylonychia.

Stylonichiafronte-a/bens DUMAS, 1937, Microzoaires, p 26 (Fig. 239v). Remarks: Incor-

rect subsequent spelling of Stylonychia.

Stylonichia frontosus DUMAS, 1937, Microzoaires, p 25 (Fig. 240b). Remarks: Incorrect

subsequent spelling of Stylonychia. Possible a somewhat deformed S. mytilus.

Stylonichia globifrons DUMAS, 1930, Microzoaires, p 61 (Fig. 238e). Remarks: Incorrect

subsequent spelling of Stylonychia.

Stylonichia gyriniformis DUMAS, 1930, Microzoaires, Planche XXIX, fig. 4 (Fig. 238p).
Remarks: Incorrect spelling of Stylonychia. Without description.

Stylonichia labiata DUMAS, 1930, Microzoaires, p 61 (Fig. 238k). Remarks: Incorrect

subsequent spelling of Stylonychia.

Stylonichia limbiformis DUMAS, 1930, Microzoaires, p 60 (Fig. 238i, q). Remarks: Incor-
rect subsequent spelling of Stylonychia. "Stylonichia limbifrons" in the legend to the fig-
ures (Planche XIV, fig. 6) is an incorrect original spelling. Possibly synonymous with
Stylonychia mytilus.

Stylonichia mamillata DUMAS, 1929, Microzoaires, p 85 (Fig. 238c). Remarks: Incorrect

subsequent spelling of Stylonychia.

Stylonichia massula DUMAS, 1929, Microzoaires, p 84, Planche XXXVII, fig. 1 and not
Planche XXXI, fig. 1 (Fig. 237e). Remarks: Incorrect subsequent spelling of Sty/onychia.
 Sty/onychia 599

Stylonichia membranaceus DUMAS, 1937, Microzoaires, p 27 (Fig. 239w). Remarks: In-

correct subsequent spelling of Sty/onychia. Possibly a S. mytilus.

Sty/onichia mutabunda DUMAS, 1930, Microzoaires, p 59, Planche XIV, fig. 1 and not
Planche XVI, fig. 1 (Fig. 238f-h). Remarks: Incorrect subsequent spelling of Sty/ony-
chia.

Stylonichia nUm. 1 - IzQUIERDO, 1906, Protozoos, P 191 (Fig. 229a). Remarks: Incorrect
subsequent spelling of Sty/onychia. 190 ).lm long. Freshwater in Chile.

Sty/onichia octonistylus DUMAS, 1937, Microzoaires, p 28 (Fig. 240e). Remarks: incor-

rect subsequent spelling of Sty/onychia. 8 (4 + 4) prominent (transverse) cirri at posterior
end.

Stylonichia reclinis DUMAS, 1930, Microzoaires, p 60 (Fig. 238j). Remarks: Incorrect

subsequent spelling of Sty/onychia.

Sty/onichia ros/rata DUMAS, 1929, Microzoaires, p 83 (Fig. 237c). Remarks: Incorrect

subsequent spelling of Sty/onychia rostrata (legend to Planche XXIX, fig. 6).

Stylonichia trochiformis DUMAS, 1929, Microzoaires, p 81 (Fig. 237b). Remarks: Incor-

rect subsequent spelling of Sty/onychia trochiformis (legend to Planche XXIX, fig. 1).
One macronucleus, 2 contractile vacuoles.

Stylonichia truncata DUMAS, 1930, Microzoaires, Planche XXX, fig. 6 (Fig. 2380). Re-
marks: Incorrect subsequent spelling of Sty/onychia. Without description.

Sty/onichia unguiculus DUMAS, 1930, Microzoaires, Planche XXVI, fig. 23 (Fig. 238r).
Remarks: Incorrect subsequent spelling of Sty/onychia. Without description.

Sty/onichia viridis DUMAS, 1937, Microzoaires, p 26 (Fig. 239p). Remarks: Incorrect sub-
sequent spelling of Stylonychia. Possible a morphogenetic stage of a Stylonychia. Green
due to ingested algae.

Stylonychia calva FRoMENTEL, 1876, Etudes microzoaires, p 273, Planche XIV, fig. 2.
Remarks: Probably the posterior part of a species of the Sty/onychia mytilus complex.
France.

Sty/onychia harbinensis FENG, Gu & ZHANG, 1990, Zoo1. Res., 11: 95. Remarks: This
name is mentioned (as "S. harbin ens is n. sp.") without description and is thus a nomen
nudum. I do not know the description.
600 SYSTEMATIC SECTION

Sty/onychia makrostyla STERKI, 1878, Z. wiss. Zool., 31: 36, 56. Remarks: Stylonichia
macrostyla WRZES. in MERMOD (1914) is an incorrect subsequent spelling of both the ge-
nus and the species name. Furthennore, MERMOD obviously erroneously mentioned WR-
ZESNIOWSKl as author of this species (or did he mean Oxytricha macrostyla WRZESNI-
OWSKl?).KARL (1932) considered S. makrostyla as valid species (subsequently incorrectly
spelled as Stylonychia macrostyla) in the subgenus Oxytricha (Stylonychia); the correct
name in KARL (1932, P 618) is thus Oxytricha (Stylonychia) macrostyla.KARL (1932) did
not realise the secondary homonymy with "Oxytricha (Urosoma) macrostyla WRZESN-
IOWSKl, 1870" (KARL 1932, P 607), now classified in Urosoma. STERK! (1878) provided
only a short description, but no figure of his species: resembles Stylonychia mytilus, how-
ever, has a metabolic (!) body, anteriorly regularly rounded and posteriorly widely trun-
cated; all cirri extremely long, frontal cirri located more posteriorly; caudal cirri about
112 of body length, loose, posteriorly not fringed; cytopharynx long, obliquely arranged.
REID (1969) recorded S. makrostyla in activated sludge.

Stylonychia monostylus FROMENTEL, 1876, Etudes microzoaires, p 275, Planche XN, fig.
12. Remarks: Probably a fragment of a hypotrich. France.

Stylonychia regularis FROMENTEL, 1876, Etudes microzoaires, p 274, Planche XIV, fig. 6.
Remarks: According to arrangement and number of ventral cirri surely not a Stylonychia.
The drawing in DUMAS (1929, P 82; as Stylonichia regularis; Fig. 237h) is a redrawing
from the original description; he found it in a bog in France. LoNGill (1895), MOLA (1928,
1935), and PARONA (1883) found this rather doubtful form in freshwater habitats in Italy.

Stylonychia sphaerica FROMENTEL, 1876, Etudes microzoaires, p 274, Planche XN, fig.
5. Remarks: Rather certainly a fragment ofa hypotrich. France.

Insufficient redescriptions

Kerona pustulata - DuJARDIN, 1841, Zoophytes, p 423, Planche VI, fig. 10, 11,14, 18,
Planche XIII, fig. 7. Remarks: 180 J.1m long!

Kerona pustufata, MOLL. Stylonychia pustulata, EHR. - CARTER, 1859, Ann. Mag. nat.
Hist., 3: 247 (Fig. 220d). Remarks: Certainly not a Sty/onychia because it is "soft in con-
sistence". About 130 J.1m long. Macronucleus consists of a single cylindrical part, or often
divided into 2 parts. Cyst without spines. Salt and freshwater habitats in Bombay, India.

Kerona silurus - DUJARDIN, 1841, Zoophytes, p 427, Planche XIII, fig. 4. Remarks: In
life 120 J.1m long. Reminiscent of a species of the Stylonychia mytilus complex.
 Stylonychia 601

Kerona silurus - FRoMENTEL, 1876, Etudes microzoaires, p 273, Planche XIV, fig. 8. Re-
marks: The outline of this superficially described, in life about 77 !lm long fonn is remi-
niscent of species of the Stylonychia mytilus complex.

Kerone pustulata - DUJARDIN, 1838, AnnIs Sci. nat., Zoologie, 10: 313, Planche 14, fig.
C, D. Remarks: Very poorly described.

Oxytricha hi/aria - SOKOLOFF, 1931, An. Inst. BioI. Univ. Mex., 2: 167 (Fig. 228b). Re-
marks: The figure does not show the characteristic arrangement of the transverse cirri.
Mexico.

Oxytricha hi/aria STOKES - CONN, 1905, Bull. Conn. St. geol nat. Hist. Surv., 2: 60 (Fig.
228g, 231 i, j). Remarks: The larger fonn (Fig. 228g) is reminiscent of Gastrostyla since
it has many postoral ventral cirri. Freshwater habitats in Connecticut, USA.

Oxytricha hi/aria STOKES - SMITH, 1914, Kans. Univ. Sci. Bull., 9: 165 (Fig. 233f). Re-
marks: The flexible body and the 4 macronuclear nodules indicate that the identification
is incorrect. Freshwater habitats in Kansas, USA.

Oxytricha hi/aria STOKES - WANG, 1925, Contr. bioI. Lab. Sci. Soc. China, 1: 51 (Fig.
242h). Remarks: The five transverse cirri are arranged in an oblique row and not in two
distinctly separated groups, as is typical for Stylonychia hi/aria. Small pond in Pe Ji Keh,
Nanking, China.

Stylonichia muscorum KAHL - CHARDEZ, 1967, Revue Ecoi. BioI. Sol, 4: 294 (Fig. 233j).
Remarks: Incorrect subsequent spelling of Sty/onychia. Nothing indicates that the identi-
fication is correct. The rather long caudal cirri and the body shape suggest that it is possi-
ble a post-divider or an abnonnal specimen of the S. mytilus complex. Soils in Belgium
(CHARDEZ 1987).

Sty/onichia mytilus, EHR. - DuMAs, 1929, Microzoaires, p 82 (Fig. 237f, i). Remarks: In-
correct subsequent spelling of Sty/onychia.

Stylonichia notophora STOKES - WANG, 1925, Contr. bioI. Lab. Sci. Soc. China, 1: 51
(Fig. 242c). Remarks: Incorrect subsequent spelling of Sty/onychia. The body shape and
the rather distinct separate caudal cirri are reminiscent of species of the S. mytilus com-
plex. Lake Ho Hu and small ponds in Nanking, China.

Stylonichia pustulata, EHR. - DUMAS, 1929, Microzoaires, p 81 (Fig. 237a). Remarks: In-
correct subsequent spelling of Stylonychia. Rather surely not a Stylonychia because ven-
tral rows are present.
602 SYSTEMATIC SECTION

Stylonichia putrina STOKES - WANG, 1925, Contr. bioI. Lab. Sci. Soc. China, 1: 52 (Fig.
242f). Remarks: Incorrect subsequent spelling of Stylonychia. The tapered posterior end
and the size (95 /lm) are reminiscent of S. vorax. Lake Ho Hu near Nanking, China.

Stylonichia sp. - OUVEIRA, KRAu & NASCIMENTO, 1954, Mems Inst. Oswaldo Cruz, 51: 409
(Fig. 234p). Remarks: Incorrect subsequent spelling. Cistern in Rio de Janeiro, Brazil.

Stylonichia sp. - WANG, 1925, Contr. bioI. Lab. Sci. Soc. China, 1: 52 (Fig. 242e). Re-
marks: Incorrect subsequent spelling. About 150 /lm. Lake Mo Chu Hu, Nanking, China.

Stylonichia tricornis (TAGLIANI, 1922) - MADRAZO-GARIBAY & L6PEZ-OCHOTERENA, 1973,

Revta Soc. mex. Hist. nat., 34: 67 (Fig. 240j). Remarks: Incorrect subsequent spelling of
Stylonychia. I could not fmd the basionym; possibly mixed up with Oxytricha tricornis
MILNE, 1886. In life 128 x 57 /lm. Certainly a deformed specimen of another species. San
Anton waterfall in Morelos State, Mexico.

Stylonychia muscorum KAHL, 1932 - MATIS & DANISKOVA, 1972, Acta Fac. Rerum nat.
Univ. comen. Bratisl., 17: 51 (Fig. 230f, g). Remarks: The identification cannot be ac-
cepted because the characteristic single micronucleus between the two macronuclear nod-
ules is not described. Furthermore, it is questionable if the 2 specimens figured are con-
specific because the transverse and caudal cirri are somewhat differently arranged. In life
75-130 /lm long. Macronuclear nodules hardly discernible because numerous dark gran-
ules (food vacuoles?) present. 5 transverse cirri arranged in 2 groups (Fig. 230g). Caudal
cirri only slightly longer than marginal cirri, reminiscent of Stylonychia bi/aria. Pools on
roads and meadows and in an artificial gravellake near Bratislava, Czechoslovakia.

Stylonychia mytilus EHRENBERG, 1838 - AGAMALIEv, 1978, Acta Protozool., 17: 440 and
AGAMALIEv, 1983, Ciliates of Caspian Sea, p III (Fig. 230c, d). Remarks: Identification
with a species of the Stylonychia mytilus complex should be avoided because the trans-
verse cirri are not arranged in 2 groups. AGAMALIEV (1978) provided following data: size
after wet silver nitrate impregnation 90--120 x 40 /lm. 2 micronuclei. Adoral zone of
membranelles almost half of body length with about 36-38 adoral membranelles. Ar-
rangement and number of frontal, ventral, and transverse cirri, see Figure 230c. Each
23-37 left and right marginal cirri. Only 3 dorsal kineties, which is rather surely a misob-
servation possibly due to the inadequate impregnation method; each kinety with 13-19
basal body pairs only. Periphyton and pelagial oflow salinity regions of the Caspian Sea.

Stylonychia mytilus - L6PEZ-OCHOTERENA, 1964, Trans. Am. microsc. Soc., 83: 430, 431,
Fig. 10, 18. Remarks: In my opinion, these two micrographs do not show a species of the
S. mytilus complex; possibly S. pustulata.

Stylonychia mytilus - LUNDIN & WEST, 1963, Free-living protozoa, p 68 (Fig. 234a). Re-
marks: Freshwater from the Upper Peninsula of Michigan, USA.
 Stylonychia 603

Stylonychia mytilus EHRENBERG, 1838 - RAMIREz DE GUERRERO, 1970, Revta Soc. mex.
Hist. nat., 31: 85 (Fig. 2261). Remarks: 150 x 60 11m. Mexico.

Stylonychia notophora STOKES - CONN, 1905, Bull. Conn. St. geol. nat. Hist. Surv., 2: 61
(Fig. 229b). Remarks: The body shape and the very long caudal cirri are reminiscent of
the S. mytilus complex. Freshwater in Connecticut, USA.

Stylonychia notophora STOKES - EDMONDSON, 1906, Proc. Davenport Acad. Sci., 11: 102
(Fig. 226h). Remarks: Probably not a S. notophora because transverse cirri not distinctly
separate and caudal cirri not narrowly spaced. Freshwater in Iowa, USA.

Stylonychia patula, EHR. - LoNGIll, 1892, Atti Soc. ligust. Sci. nat. geogr., 3: 155. Re-
marks: I could not find the basionym; probably he meant S. pustulata.

Stylonychia pustulata - BAmsH, 1992, Freshwater zooplankton of India, p 58, Fig. 55.
Remarks: Very superficially illustrated. Planktonic. India.

Stylonychia pustulata - CHEN, 1944, J. Morph., 75: 341 (Fig. 242i). Remarks: The ar-
rangement of the frontoventral, ventral, and transverse cirri is not in accordance with the
authoritative descriptions of S. pustulata. China.

Stylonychia pustulata E. - CIENKOWSKY, 1855, Z. wiss. Zool., 6: 304 (Fig. 222k, 1). Re-
marks: The body shape and the obviously rather inconspicuous caudal cirri suggest that
he did not observe S. pustulata, although CIENKOWSKY described a "star-shaped" cyst.

Sty/onychia pustulata EHRBG. - CONN, 1905, Bull. Conn. St. geol. nat. Hist. Surv., 2: 60
(Fig. 228f, 229c). Remarks: Freshwater in Connecticut, USA.

Stylonychia pustulata - ECKHARD, 1846, Arch. Naturgesch., 12: 231 (Fig. 232h). Re-
marks: Probably a Euplotes species.

Stylonychia pustulata - FROMENTEL, 1876, Etudes microzoaires, p 274, Planche XIV, fig.
9. Remarks: Stylonychia pustullata (p 359) is an incorrect subsequent spelling. The out-
line, the very prominent transverse cirri, and the conspicuously widely separated caudal
cirri are reminiscent of the S. mytilus complex.

Stylonychia pustulata, EHRBG. - KOFFMAN, 1926, Acta Zoo!., Stockh., 7: 312 (Fig. 231e).
Remarks: The parallel body margins and the rather inconspicuous caudal cirri are remi-
niscent of Sterkiella histriomuscorum, which is common in terrestrial habitats, whereas S.
pustulata is not reliably recorded from soil. Garden soil near Stockholm, Sweden.
 604 SYSTEMATIC SECTION

Stylonychia pustulata EHRENBERG - LUNDIN & WEST, 1963, Free-living protozoa, p 68

(Fig. 230e). Remarks: See also WEST & LUNDIN (1963, P 108). Freshwater from the Up-
per Peninsula of Michigan, USA.

Stylonychia pustulata EHRENBERG - NAlDu, 1965, Hydrobiologia, 25: 562 (Fig. 2241). Re-
marks: In life 130--169 x 60--70 Jim. Body shape and arrangement of ventral cirri unreli-
able. Gutter water in Vijayawad, India.

Sty/onychia putrina STOKES - BARWICK, BEVERIDGE, BRAZIER, CLOSE, HIRsCHFELD, PILLAI,

RAMSAY, ROBINSON, STEVENS & TODD, 1955, Tuatara, 5: 96 (Fig. 223h). Remarks: Fish-
pond in Wellington area, New Zealand.

Sty/onychia putrina STOKES - BURKOVSKY, 1984, Ecology of free-living ciliates, p 167

(Fig. 233k). Remarks: The size (200 Jim), the arrangement of the cirri, and the lack of
caudal cirri indicate misidentification.

Sty/onychia putrina STOKES - CONN, 1905, Bull. Conn. St. geoi. nat. Hist. Surv., 2: 61
(Fig. 228e). Remarks: The tapered posterior end of the cell is reminiscent of S. vorax,
whereas the somewhat broadened anterior portion indicates similarity with a species of
the S. myti/us complex. Freshwater in Connecticut, USA.

Sty/onychia putrina STOKES - SAMANO & SOKOLOFF, 1931, Monografias Inst. BioI. Univ.
naco Mex., 1: 36 (Fig. 24li). Remarks: The body shape does not match the original de-
scription. Mexico.

Sty/onychia sp. - ALoNSO & PEREz-SILVA, 1963, Microbiologia esp., 16: 56 (Fig. 231g).
Remarks: ALoNSO & PEREZ-SILVA (1963b) mention that their population is very similar to
Sty/onychia stylomuscorum. But, according to ALONSO & PEREz-SILVA (1963a, 1965b),
their population has 2-4 micronuclei, whereas S. stylomuscorum is characterised by a sin-
gle micronucleus (see Fig. 167b-d). The general morphology, the body size, and the in-
fraciliature are reminiscent of Sterkiella histriomuscorum; however, in this species the 3
caudal cirri are not as long as described by the Spanish authors. ALoNSO & PEREz-SILVA
(1963b) population rather variable in size; nonnal specimens in life 80--100 x 40--60 JIm;
cannibal giants measure 175 x 98 JIm, smallest specimens 45 x 15 Jim. Arrangement of
cirri, see Figure 231g. About 20--30 right and 15-20 left marginal cirri. Stagnant water
near Charmatin, Madrid, Spain. ALONSO (1965), ALONSO & PEREZ-SILVA (1965a, 1966a, b,
1967, 1969), JARENO et al. (1969b, 1970), and PEREZ-SILVA & ALONSO (1966) studied
mainly the polytene chromosomes in the macronuclear anlage of this population. Minimal
inhibitory concentration of tinidazole is 0.5 mg ml· l , minimal lethal concentration
1 mg mI· 1 (ALONSO 1979).
 Stylonychia 605

Sty/onychia sp. (n. sp.) - LEPSI, 1951, Buletin sti. Acad. Repub. pop. rom., 3: 521 (Fig.
220f). Remarks: LEPSI (1965, p 218, legend of his Fig. 97m) figured a "Stylonychia n.
sp." which is identical with the LEPSI (l951) form. A conspicuous cirral row right of the
undulating membranes. Soil from Temesvar, Romania.

Stylonychia sp. - NIKOUUK & GELTZER, 1972, Pocvennye prostejsie SSSR, p 133 (Fig.
240f). Remarks: Soil in the USSR.

Stylonychia sp. - VUXANOVICI, 1963, Studii cerc. Biol., 15: 212 (Fig. 220g). Remarks: In
life about 100--120 !lm long; adoral zone of membranelles 1/3 of body length; 5 trans-
verse cirri protruding about 2/3 beyond posterior end of cell; 3 conspicuous caudal cirri;
18-20, about 9 !lm long marginal cirri; contractile vacuole near mid-body; cytoplasm
very transparent, filled with 6-8 !lm sized symbiotic algae! Freshwater in Bucharest, Ro-
mania. The illustration shows an anteriorly distinctly curved undulating membrane, indi-
cating that it perhaps belongs to Cyrtohymena or Steinia. Differs from Oxytricha chlorel-
ligera in that it has longer caudal cirri and transverse cirri which protrude beyond poste-
rior end of cell.

Stylonychia sp. - VUXANOVICI, 1963, Studii. Cerc. Biol., 15: 212 (Fig. 220h-j). Remarks:
In life 120--180 !lm long; number and arrangement of ventral cirri unreliable; body elastic
but not contractile and not metabolic; cytoplasm transparent, with 3-10 !lm sized symbi-
otic algae(?). Freshwater habitat in Bucharest, Romania.

Stylonychia sp. - WEST & LUNDIN, 1963, Pap. Mich. Acad. Sci., 48: 109 (Fig. 220a). Re-
marks: See also LUNDIN & WEST (l963, P 166). Perhaps a Gastrostyla because about 26
frontal, ventral, and transverse cirri present. Limnetic habitats in Michigan, USA.

Stylonychia (?) sp. - NANsEN, 1906, Sci. Res. Norw. N. Polar Exped., 5: 16, Plate IV,
Fig. 1-5. Remarks: In accumulations of algae from ponds on the ice of the North Polar
Sea

Stylonychia steini - HAYASHI, 1959, Zool. Mag., Tokyo, 68: 239 and MURAMATSU, 1957,
Misc. Rep. Yamashina Inst. Om. ZOOl., 11: 467. Remarks: MURAMATSU mentioned this
name in a paper on the protozoa of the city ofHakodate and Oonuma Park, Japan, where
he found it in foul water, together with Glaucoma scintillans and Colpidium colpoda. I
do not know which species they meant, possibly Histrio stein;; STERKI, 1878. Stylonychia
steinii STILLER was only established in 1974.
606 SYSTEMATIC SECTION

Coniculostomum NJINE, 1979

1977 Coniculostomum monilata - NJINE, AnnIs Stn Homo!. Besse, 11: 24 (see nomenclature).
1978 Coniculostomum monilata- NJINE, Dissertation, p 167 (see nomenclature).
1979 Coniculostomum NJINE, Protistologica, 15: 353 - Type (by monotypy): Laurentia monilata DRAGESCO
& NJlNE, 1971.
1979 Laurentiella (Laurosticha) JANKOWSKI, Trudy zoo!. Inst., Leningr., 86: 57 - Type (original designation):
Laurentia monilata DRAGESCO & NJINE, 1971.
1986 Coniculostomum NJINE, 1978 - DRAGESCO & DRAGESCO-KERNEIS, Faune tropicale, 26: 455.
1990 Coniculostomum NJINE, 1978 - KAMRA & SAPRA, Europ. J. Protistol., 25: 264.
1994 Coniculostomum - KAMRA, SAPRA & AMMERMANN, Europ. 1. Protisto!., 30: 63.
1997 Coniculostomum NJINE, 1979 - BERGER & FOISSNER, Arch. Protistenk., 148: 149.

C h a r act e r i sat ion: Undulating membranes in Stylonychia pattern. Two or more

right and 1 or more left rows of marginal cirri, distinctly separate posteriorly. Six or more
dorsal kineties. Caudal cirri present and usually distinctly elongated. Primordium II of
proter originates from oral primordium. Primordia V and VI of proter, and primordium
IV of opisthe originate from postoral ventral cirrus V/4. Parental marginal cirri partially
retained after division. Dorsal morphogenesis in Coniculostomum pattern.

Add i t ion a 1 c h a r act e r s: Large to very large. Outline Stylonychia mytilus-like,

that is, anteriorly distinctly broader than posteriorly. Flattened about 2: 1 dorso-ventrally.
Body rigid, not contractile. Macronucleus moniliform, anteriorly usually curved to the
right (in ventral view) and thus often formed like a question mark. Contractile vacuole
about in mid-body. Cortical granules lacking. Cytoplasm colourless. Adoral zone of
membranelIes about 55 % of body length. Buccal area very large, Stylonychia mytilus-
like! Frontal and transverse cirri and buccal cirrus enlarged. Cirri of left and outer right
marginal row rather closely arranged, inner right marginal row(s) (= retained parental
rows) distinctly more widely spaced. Dorsal kineties 1-3 distinctly curved anteriorly. In-
variably 3 very long caudal cirri on dorsal kineties 1,2, and 4. Dorsal cilia about 2-4 J.1m
long.

Nom e n c I a t u rea n d t a x 0 nom y: The name Coniculostomum was mentioned

for the first time in NJINE (1977). Because it was published without sufficient descriptive
information to satisfy the criteria of availability, it has to be considered a nomen nudum
(lCZN 1985). Coniculostomum was mentioned, for the second time, in the thesis by NJINE
(1978) and, finally, NJINE (1979) published it in the Protistologica Therefore I assume
that his thesis is unpublished and does not constitute a publication according to Article 9
of the IcZN (1985). Thus, 1979, and not 1978, as supposed by TUFFRAU (1979, 1987), is
the correct date of publication. The problem is complicated somewhat further since JAN-
KOWSKI (1979) created the subgenus Laurentiella (Laurosticha) to include the same spe-
cies. JANKOWSKI informed me that his paper was published on September 17, 1979. Both
JANKOWSKI (pers. comm.) and I wrote to the publishers of the "Protistologica" asking for
the exact date of publication of vol. 15, fasc. 3 of this journal. Regrettably, we never re-
 Coniculostomum 607

ceived a reply, so that I could not fmally clarify priority. However, in agreement with
other authors, I (preliminarily) consider Coniculostomum as the senior synonym. Incor-
rect subsequent spellings: Conicyc/ostomum NJINE, 1979 (HEMBERGER 1982, P 275); Cu-
niculostomum monilata (DRAGESCO 1985); Lautosticha (DRAGESCO & DRAGESCo-KERNEIS
1986, P 455).
The characterisation is according to BERGER & FOISSNER (1997). For a detailed de-
scription of patterns, see chapter 2 in the general section. Previously, Coniculostomum
monilata was classified in the Holostichidae, especially because of the longitudinal cirral
rows (DRAGESCO & DRAGESCO-KERNEIS 1986, DRAGESCO & NJINE 1971, TUFFRAU 1979,
1987). HEMBERGER (1982, P 223; unpublished thesis) proposed a classification within the
oxytrichid taxon Pleurotricha (P. monilata is thus an unpublished combination), which
has, however, no caudal cirri. Furthermore, in Pleurotricha the cirri of all right marginal
rows are nearly equally spaced (for example, Fig. 18ge, h, i) due to a different morpho-
genesis (all marginal rows are new, that is, no parental right marginal rows are retained
after division).
The oxytrichid lineage of Coniculostomum was fully recognised only recently by de-
tailed morphogenetic studies (KAMRA & SAPRA 1989, 1990, KAMRA et al. 1994) showing
several conspicuous morphological and morphogenetical synapomorphies with the Styla-
nychia mytilus complex (Fig. 25a; BERGER & FOISSNER 1997). Conversely, a classification
within the amphisiellids (TUFFRAU & FLEURY 1994, P 141), the kahliellids (EIGNER 1995,
P 363), or the parakahliellids (EIGNER 1997) is very likely artificial. According to EIGNER
(1995), C. monilata is closely related to Neogeneia EIGNER, 1995, Kahliella CORLISS,
1960, Engelmanniella FOISSNER, 1982, and Parakahliella BERGER, FOISSNER & ADAM,
1985 because in all of these parental (old) cirral rows are retained after division. How-
ever, parallel evolution of this rather simple character (parental ciliature is not reabsorbed
after division) is a much more parsimonious explanation than parallel evolution of the
rather complicated 18 frontal-ventral-transverse cirral pattern (including identical mor-
phogenesis!) together (!) with the fragmentation of dorsal kinety 3. Furthermore, am-
phisiellids and kahliellids are flexible, whereas Coniculostomum has a firm body. This
character separates Coniculostomum also from the oxytrichids Allotricha, Parurosoma,
and Onychodromopsis, which also have an increased number of marginal rows. Further-
more, Coniculostomum should not be confused with Pleurotricha (Fig. 189), which lacks
caudal cirri and has two macronuclear nodules, or with Laurentiella (Fig. 196) and Ony-
chodromus (Fig. 193-195), which do not have, among other things, the typical 18-cirri
pattern.
Two species have been assigned to Coniculostomum, namely the type, C. monilata
and C. bimarginata. The latter is very likely the junior synonym of Pleurotricha indica.
Identification needs morphometric data, that is, protargol impregnated specimens.
 608 SYSTEMATIC SECTION

Key to species

1 Usually more than 2 right marginal rows; dorsal kinety 4 not distinctly curved anteri-
orly, 5-7 dorsomarginal kineties (Fig. 168a, e, f, i) ........... C. monilata (p 608)
Usually 2, rarely 3 right marginal rows; dorsal kinety 4 distinctly curved anteriorly.
2-4 dorsomarginal kineties (Fig. 170a--d) ...................... C. indica (p 620)

Coniculostomum monilata (DRAGESCO & NJINE, 1971) NJINE, 1979

(Fig. 15a-t, 168a-z, 169a-h, Tables 4,37)

1971 Laurentia monilala DRAGESCO & NJINE, Annis Fac. Sci. Univ. fed. Cameroun, 7-8: 124 (Fig. 168a, c, d,
g, h).
1971 Laurentiella monilata DRAGESCO & NJINE, Annis Fac. Sci. Univ. fed. Cameroun, 7-8: 97.
1972 Laurentie/Ia monilata DRAGESCO & NJINIi - DRAGESCO, AnnIs Fac. Sci. Univ. fed. Cameroun, 9: 118.
1977 Coniculostomum monilata - NJINE, Annis Stn Hrnnol. Besse, II: 24 (see nomenclature of genus
section).
1978 Coniculostomum monilafa - NJINE, Dissertation, p 167 (unpublished, see nomenclature of genus sec-
tion).
1979 Coniculoslomum monilala - NJlNE, Protistologica, 15: 353.
1979 Laurenliella (Laurosficha) monilata (DRAGESCO et NlIl~E, 1971) - JANKOWSKI, Trudy zool. Inst.,
Leningr., 86: 57.
1986 Coniculosfomum monilafa (DRAGESCO & NJINE, 1981) - DRAGESCO & DRAGESco-KERNms, Faune tropi-
cale, 26: 456 (Fig. 168b).
1989 Coniculostomum moniiataNJINE, 1979 -KAMRA & SAPRA, Int. Congr. Protozoo!., 8: 104.
1990 Conicu/oslomum monilata (DRAGESCO & NJINE, 1971) NJINE, 1978 - KAMRA & SAPRA, Europ. J. Pro-
tistol., 25: 264 (Fig. 168e, f, i-z, 169lrh).
1991 Coniculostomum monilata - KAMRA & SAPRA, Europ. J. Protisto!., 27: 331 (Fig. 15lrt).

Nom e n c I at u rea n d t a x 0 nom y: Nomenclature, see relevant chapter in genus

section. I do not agree with EIGNER (1995), who supposed synonymy of C. monilata and
C. bimarginata (the latter is here classified as junior synonym of C. indica) because sev-
eral characters show significant differences (see key and Table 37). Furthermore, KAMRA
et al. (1994) reported reproductive isolation of C. monilata and C. bimarginata, that is,
they did not conjugate. Comparison with other taxa (Laurentiella, Onychodromus), see
Coniculostomum section.

M 0 r p hoI 0 g y and b i 0 log y: Unfortunately no live size available; according to

measurements of protargol impregnated material length of live specimens may vary be-
tween 250-350 !lm; about 2.1 times as long as broad. Beads of macronucleus about

Fig. 168a-d Coniculoslomum monilata (a, c, from DRAGESCO & NJINE 1971; b, from DRAGESCO & DRAGES- ~
CO-KERNElS 1986; d, after DRAGESCO & NJINE 1971 from DRAGESCO & DRAGESCO-KERNEIs 1986. Ire, protargo!
impregnation; d, Feulgen stain). a, b) Ventral infraciliature, a = 295 j.lm, b = 280 j.lm. c) Ventral infraciliature
of posterior part of cell. d) Different shapes of nuclear apparatus. CC = caudal cirri.
Com·culostomum 609
610 SYSTEMATIC SECTION

.'
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 Coniculostomum 611

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Fig. 168i-k Coniculostomum monilata (from KAMRA & SAPRA 1990. i, j, protargol impregnation; k, Feulgen
stain). i, j) Ventral and dorsal infraciliature. Because of the widely spaced cirri of the inner right (= parental)
marginal rows, the typical oxytrichid 18 frontal-ventral-transverse cirra1 pattern is difficult to recognise. Ar-
rowheads in CD mark caudal cirri at end of dorsal kineties 1,2, and 4. k) Nuclear apparatus in dorsal view. Ma
= bead of the moniliform macronucleus, Mi = micronucleus; p 608.

10-16 Ilm long (Fig. 168a, d, k). Posterior part of paroral distinctly curved to the left and
thus optically crossing endoral (Fig. 168a, e, i). Cirrus 11112 distinctly enlarged. Arrange-
ment of frontal, ventral, and transverse cirri rather invariably as shown in Figure 168e, al-
most identical to in species of the Stylonychia mytilus complex (cp. with Fig. 149a,
IS0a). According to KAMRA & SAPRA (1990) usually 8, rarely 10 "frontal" cirri (compris-
ing frontal, buccal, and frontoventral cirri; mean = 8.1; n = 20). Invariably S ventral (3

f- Fig. 168e-b Coniculostomum monilata (e, f, from KAMRA & SAPRA 1990; g, h, from DRAGESCO & NJINE 1971.
e-h, protargol impregnation). e) Ventral infraciliature, 287 ~m. The 18 frontal, ventral, and transverse cirri are
black. Arrow marks posterior transverse cirrus (cirrus VII). f, h) Dorsal infraciliature, f= 255 ~m, h = 340 ~m.
(f) is seen from the ventral side. Arrows denote outer dorsomarginal kinety. Kinety 4 is anteriorly more or less
straight; an important difference to C. indica, where it is distinctly curved. g) FibrilIar system around a basal
body pair. BC = buccal cirrus, CC = caudal cirri, e = endoral, FC = right frontal cirrus, p = paroral, PRVC =
posterior pretransverse ventral cirrus, PVC = posterior postoral ventral cirrus, RI-R4 = right marginal rows
1-4, 1-4 = dorsal kineties 1-4; p608.
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Fig. 168HJ Coniculostomum monilata (from KAMRA & SAPRA 1990. I~, ventral infraciliature of morphogenetic stages, protargol impregnation). I, m) Very early
stages, I = 330 /lm, m = 345 /lm. n, 0) Early stages, n = 325 /lnl, 0 = 330 /lm. The 3 small arrows in (n) mark the 3 parental cirri which participate in primordia forma-
tion. MP = primordium of right marginal row ofproter, OP = oral primordium; p 608.
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Fig. 168p-s Coniculostomum monilata (from KAMRA & SAPRA 1990. p-s, ventral infraciliature of morphogenetic stages, protargol impregnation). p, q) Middle stages, p
0'1
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w
mordia of dorsomarginal kineties of proter; p 608.
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Fig. 168t-x Coniculostomum monilata (from KAMRA & SAPRA 1990. t-x, ventral and dorsal infraciliature of morphogenetic stages, protargol impregnation). t, u) Very
late stages, t =330 !lm, U =310 !lm. v, w) Proter and opisthe, v, w = ISS Jlm. x) Early stage of dorsal morphogenesis, 32S Jlm; p 608.
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.: l
--.......--------- . :' j."
• o ci CC~. ·. 'e: ': e'
y o o
.. a
.0 • , 0
o z c
Fig. 168y, Z, 169a-c Coniculostomum monilata (from KAMRA & SAPRA 1990. y, Z, a-c, dorsal infraciliature of morphogenetic stages, protargol impregnation). y, Z, a)
0'\
Middle, late, and very late stage, Y= 340 ",m, Z = 365 ",m, a = 345 ",m. Arrowheads in (y) mark the fragmentation site of dorsal kinety 3. b, c) Proter and opisthe, b =
VI
165 ",m, c = 175 ",m. Arrowheads denote new inner dorsomarginal row, CC = caudal cirri, 1-4 = dorsal kineties 1-4; p 608.
-
 616 SYSTEMATIC SECTION

d e

9 h

Fig. 169d-b Coniculostomum monilata (from KAMRA & SAPRA 1990). Diagrams showing retention of right
marginal rows and their fate during morphogenesis of cell division. d) A non-dividing cell with 4 rows
(PI-P4). e) A single new row (Nl) arises for each daughter cell. t) At cytokinesis, P4 is reabsorbed; PI-P3 are
cleaved in the middle. g) Each of the 2 daughter cells shows 1 new (Nl) and 3 retained parental (PI-P3) right
marginal rows. b) Next generation; another new right marginal row (N2) is added and P3 will be resorbed. (h)
shows same stage as (e); p 608.

postoral and 2 pretransverse) and usually 5, rarely 6 (mean = 5.1; n = 20) transverse cirri.
Transverse cirrus VII distinctly displaced posteriad (Fig. 168e, arrow). In total about
9-12 dorsal kineties. Between dorsal kinety 4 and the two outermost (= new) dorsomargi-
nal kineties a rather variable (3--6) number of parental (= old) dorsomarginal kineties.
Anterior end of dorsal kinety 4 not distinctly curved (Fig. 168f, h, j).
 Coniculostomum 617

Cyst spherical, after protargol impregnation 76-94 Ilm across (mean = 86 Jim, SD =
5.2 Ilm, n = 10); diameter of protoplast 52-63 Jim (KAMRA & SAPRA 1991). Cyst wall
made up of 4 layers, the ectocyst thrown into several folds making hexagonal patterns
(Fig. 15a). For the morphogenetic process during excystment, see chapter 1'10-3 (general
section) and Figures 15M.

M 0 r p hog e n e sis: This part of life cycle is described in great detail by I<AMRA &
SAPRA (1990). Cell division commences with the formation of an oral primordium in the
proximity of the left transverse cirrus and occasionally even close to the next transverse
cirrus (Fig. 1681, m). Some basal bodies in the anterior portion of the oral primordium
proliferate to form the anlage I, which differentiates into the undulating membranes and
the left frontal cirrus of the opisthe (Fig. 168n-p). The anterior portion of the parental pa-
roral reorganices and forms the left frontal cirrus of the proter (Fig. 168o-q). The endoral
also undergoes reorganisation (Fig. 168q). The parental adoral zone of membranelles is
retained intact for the proter.
The origin of the frontal, ventral, and transverse primordia is shown in Table 4. Cirri
1II12, IV/3, and V/4 are involved in streak formation, identically to in the species of the
Stylonychia mytilus complex (Fig. 168n, Table 4). Cirri IIJJ2 and V/4 remain partly re-
tained until primordia differentiation begins (Fig. 168q).
Two primordia originate to the right of the anterior ends of the outer right and the left
marginal row by disaggregation of 1, and sometimes 2 parental marginal cirri. Two more
primordia arise similarly near mid-body (Fig. 168o-q). These 4 primordia form a new
right and a new left marginal row for each filial product. The parental left marginal row is
resorbed in a posteriad direction. The parental right marginal rows are retained. Their fate
is depicted in Figures 169e-h. The parental outer right marginal row (P 1 in Fig. 169d)
simply takes up position 2 in the 2 daughter cells as the new row is formed to its right
(Fig. I6ge); the cirri in PI move apart as the cell elongates and the row is equatorially
cleaved during cytokinesis. Likewise, the other parental right marginal rows (P2 and P3
in Fig. 169d-h) are also reoriented to occupy the inner positions 3 and 4 respectively.
The innermost parental right marginal row P4 may be partially or completely resorbed.
Thus, the ''new'' marginal cirri formed during morphogenesis persist for 3-5 generations.
For instance, a non-dividing cell possessing 4 right marginal rows actually has persistent
ciliary structures of the previous 3 generations. Retention of parental (old) marginal cirri
occurs also in kahliellids (BERGER & FOISSNER 1987a, EIGNER 1995), which is, however,
very likely a typical case of parallel evolution.
Morphogenesis of dorsal kineties proceeds in Coniculostomum pattern (see character
23 in chapter 2 of general section). Briefly, kineties 1-4 show no peculiarities and divide
according to the widely distributed Oxytricha pattern, that is, simple fragmentation ofki-
nety 3 (Fig. 168x-z, 169a-c). Furthermore, similarly to in other oxytrichids, for example,
Stylonychia, Sterkiella, two dorsomarginal kineties for each daughter cell arise right of
the right marginal primordium (Fig. 168r, s). However, parental dorsomarginal kineties
are retained in the post dividers, which is a peculiarity within the oxytrichids (Fig. 168z,
618 SYSTEMATIC SECTION

Table 37 Morphometric data of Coniculostomum indica (ind, from AMMERMANN et aI. 1987; bim, the syno-
nym, C. bimarginata, from KAMRA et aI. 1994) and C. monilata (mol, from DRAGESCO & DRAGESco-KERNEIS
1986; mo2, from KAMRA & SAPRA 1990). All data are based on protargol-impregnated specimens. All measure-
ments in micrometres. ? = sample size not indicated. If only one value is known it is listed in the mean column,
if2 values are available they are listed as Min and Max. CV = coefficient of variation (in %), Max = maximum
value, mean = arithmetic mean, Min = minimum value, n = sample size, SD = standard deviation

Character Species mean SD7 CV Min Max n

Body, length bim 169.6 7.8 4.6 155 183 20
mol' 210 300 9
mo2 252.2 20.3 8.0 211 284 20
Body, width bim 83.3 4.6 5.5 76 93 20
mol' 115 141 9
mo2 107.8 13.0 12.1 85 128 20
Adoral zone of membranelles, bim 89.6 3.6 4.0 84 98 20
length moP 164.0 I
mo2 136.1 9.3 6.8 123 157 20
Adoral membranelles, number bim 62.7 2.4 3.9 58 67 20
ind 73.0 5 69 79 10
mol 78 104 ?
mo2 88.2 5.3 6.0 78 98 20
Left marginal row, number bim 27.3 2.0 7.2 24 31 20
of cirri ind 28.0 9 25 32 10
mol 30 44 ?
mo2 36.2 4.0 11.2 27 42 20
Right marginal rows, number bim 2.3 0.5 20.4 2 3 20
moP 4 5 ?
mo2 4.2 0.4 8.9 3 5 20
Right marginal row I, number bim 39.9 2.8 7.1 37 47 20
of cirri ind 44.0 10 40 48 10
mol 47.0 25 5 60 ?
mo2 56.2 4.7 8.3 48 65 20
Right marginal row 2, number bim 10.3 1.6 15.1 7 13 20
ofcirri4 ind 15.0 10 13 16 10
mol 11 51 ?
mo2 28.2 10.4 37.0 17 50 20
Right marginal row 3, number bim 0.6 1.0 174.4 0 3 20
ofcirri4 mol 5 27 ?
mo2 11.8 4.8 40.6 5 24 20
Right marginal row 4, number mol 2 12 ?
ofcirri4 mo2 4.1 2.2 59.2 0 9 20
Right marginal row 5, number mol 0 5 ?
ofcirri4 mo2 0.3 0.7 288.0 0 3 20
Macronucleus, number of beads bim 8.1 0.7 8.5 7 10 20
ind 2 6 ?
mol 9 16 ?
mo2 12.9 2.0 15.6 9 18 20
Micronuclei, number ind 6.0 26 3 10 10
mol 3 6 ?
mo2 4.5 1.6 36.2 2 8 20
 Coniculostomum 619

Table 37 Continued

Character Species mean SD7 CV Min Max n

Micronuclei, diameter ind 3.0 ?
mol 4 6 ?
mo2 5.5 0.4 7.7 5 6 20
Dorsal kineties, number bim 4.0 0 0 4 4 20
ind 4.0 ?
mo2 4.0 0 0 4 4 10
Dorsomarginal kineties, number bim 3.1 0.9 28.2 2 4 10
ind 2.0 ?
mo2 6.0 0.8 13.7 5 7 10
Dorsal kinety 1, number of basal bim 72.8 4.7 6.4 63 81 10
body pairs6 ind 68.0 12.0 60 79 5
mo2 102.4 19.1 8.9 88 115 10
Dorsal kinety 2, number of basal bim 59.6 4.0 6.7 54 65 10
body pairs6 ind 57.0 16.0 42 60 5
mo2 75.4 7.8 10.3 65 86 10
Dorsal kinety 3, number of basal bim 46.6 3.1 6.6 42 51 10
body pairs6 ind 49.0 12.0 41 58 5
mo2 58.8 4.2 7.2 51 66 10
Dorsal kinety 4, number of basal bim 39.3 5.0 12.7 32 47 10
body pairs6 ind 31.0 16.0 26 39 5
mo2 52.5 5.7 10.9 45 61 10
Dorsomarginal kinety 1, number bim 21.8 2.9 13.1 18 26 10
of basal body pairs6 ind 18.0 22.0 12 21 5
mo2 20.2 4.1 20.5 14 26 10
Dorsomarginal kinety 2, number bim 39.8 2.9 7.3 36 44 10
of basal body pairs6 ind 35.0 14.0 31 44 5
mo2 36.0 4.9 13.7 27 44 10
Dorsomarginal kinety 3, number bim 7.1 5.4 75.8 0 14 10
of basal body pairs6 mo2 16.4 8.0 48.6 7 29 10
Dorsomarginal kinety 4, number bim 2.0 2.7 133.3 0 6 10
of basal body pairs6 mo2 17.5 5.8 33.1 6 25 10
Dorsomarginal kinety 5, number mo2 6.9 5.1 73.8 2 20 10
of basal body pairs6
Dorsomarginal kinety 6, number mo2 3.6 3.0 81.9 0 9 10
of basal body pairs6
Dorsomarginal kinety 7, number mo2 1.1 1.8 162.7 0 4 10
of basal body pairs6

1 Population from Benin.

2 From Figure 168a.
) From DRAGESCO & Nmffi (1971). Row"Vs" in DRAGESCO & DRAGESCo-KERNEIS (1986) is likely a ventral cirrus.
4 For designation of right marginal rows, see Figure 168e.

S This value is from DRAGESCO & NJINE (1971). DRAGESCO & DRAGESCO-KERNEIS (1986) gave (erroneously?) 11

as minimum value.
6 For designation of dorsal kineties and dorsomarginal kineties, see Figures 168f, h and 170h. Dorsomarginal

kinety 1 in Coniculostomum monilata is the outer one. 10 Table 1 ofKAMRA et aI. (1994) the values of dorso-
marginal kineties I and 2 are confused.
7 In Coniculostomum indica the standard deviation is given in %.
620 SYSTEMATIC SECTION

169a-<:). The row(s) arranged closest to dorsal kinety 4 is (are) resorbed during morpho-
genesis.
KAMRA & SAPRA (1990) also described morphogenesis of reorganisation, which is in
many ways similar to that of division. However, the two outermost right marginal rows of
a newly reorganised cell have about the same number of cirri because there is no cellular
elongation associated with reorganisation. This explains why some non-dividing cells
have adjacent rows with equally spaced cirri.

o c cur r e n c e and e colo g y: Rather certainly confined to limnetic habitats and

possibly common in tropical Africa and Asia. Locus classicus is a wheel-track near the
La Mefou River, Cameroon (DRAGESCO & NJINE 1971). DRAGESCO (1972) recorded Coni-
culostomum monilata from Uganda and DRAGESCO & DRAGESCo-KERNEIS (1986) from Be-
nin. KAMRA & SAPRA (1990) isolated C. monilata from freshwater near Delhi, India.
Feeds also on Arcella sp. (DRAGESCO 1972). Cultures can be maintained at 23°C in
Pringsheim medium and with Chlorogonium elongatum as food (KAMRA & SAPRA 1990).
Biomass of 106 individuals about 1000 mg. NJINE (1977) found C. monilata at 23-26 °C
with a frequency of36 % in a temporary pond in Cameroon.

Coniculostomum indica CAMMERMANN', SAPRA & ScHLEGEL, 1987) comb. nov.

(Fig. 170a-j, 171a-c, Table 37)

1987 Pleurotricha indica AMMERMANN, SAPRA & SCHLEGEL, J. Protozool., 34: 31 (Fig. 170a-j; the slide of
holotype material is deposited in the Zoological Collection of the University ofTuebingen, Gennany).
1994 Coniculostomum bimarginata KAMRA, SAPRA & AMMERMANN, Europ. J. Protistol., 30: 55 (Fig. 171a-c;
new synonym; the holotype, that is a slide with protargol impregnated cells of a clone is deposited in
the British Natural History Museum, London, accession number 1993:8:10:1).

T a x 0 nom y: Pleurotricha indica has caudal cirri so that the classification in Pleura-
tricha, which lacks such cirri, is incorrect. The cirri of the inner right marginal row are
distinctly more widely spaced than those of the outer row (Fig. 170b, c, e, g), strongly in-
dicating that it belongs to Coniculostomum. KAMRA et al. (1994) emphasised the validity
of both P. indica and Coniculostomum bimarginata; however, they omitted to transfer P.
indica to Coniculostomum, which is hereby undertaken. They quoted the cell size, the
number of adoral membranelIes, and the number of beads in the macronucleus as main
differences, stressing that these are definitely not due to culture conditions or age factors
because both species were grown under identical laboratory conditions (Table 37). Con-
versely, I assume that these and other differences (for example, lack of parental dorso-
marginal kineties in Pleurotricha indica) are too indistinct for species separation. Fur-
thermore, both taxa were discovered in the same locality. The only significant difference
is the higher number of frontal-ventral-transverse cirri streaks in P. indica (7 against 6 in
other oxytrichids; Fig. 170t). However, KAMRA et al. (1994) found reorganisers of P. in-
dica in the type slides showing unambiguously only 6 frontal-ventral-transverse cirri
 Coniculostomum 621

.. ' .......
......

Fig. 170a-£ Coniculostomum indica (from AMMERMANN et al. 1987; t; schematic redrawing from Fig. 170g,
original. a, from life?; b-e, protargol impregnation). a) Ventral view, 235 J.lm. b, d) Ventral and dorsal infra-
ciliature of a representative specimen, 215 J.lm. Dorsal kinety 4 (arrowhead) is distinctly curved anteriorly; an
important difference to C. monilata. c, e) Ventral infraciliature of "aberrant" specimens, c, e = 215 J.lm. f) Pre-
sumed origin of the frontal-ventral-transverse cirri. Cirri which originate from the same streak are connected
by a broken line. The arrowhead marks that streak which is very probably lacking in the synonym, C. bimargi-
nata and in the type species, C. monila/a; p 620.
622 SYSTEMATIC SECTION
 Coniculostomum 623

.' . ·1
............. ····2
. .········3
.... 4

-----
......
.......
....... 4
~
~
DQ
~
#) ~ .,
~'8
::::.: 0
~" 00
0
Q C) 0

0
0
0
0
0
cc
0

'"'"
e;,'"'"
",,""
b
c

Fig. 171a-c Coniculostomum indica (from KAMRA et al. 1994. a, after live observation, protargo\ impregna-
tion, and Feulgen stain; b, c, protargol impregnation). a) Ventral view, 145 11m. b, c) Ventral and dorsal infia-
ciliature, 165 11m. Arrows mark marginal rows. Arrowheads mark inner (= parental) dorsomarginal kineties.
CC = caudal cirri, e = endoraI, p = paroral, 1-4 = dorsal kineties 1-4; p 620.

streaks (Fig. 32, 33 in KAMRA et al. 1994). Because of these uncertainties, I provide both
descriptions separately. Reinvestigation of further populations recommended to check
variability of these characters.

M 0 r p hoi 0 g y and b i 0 log y: Slightly starved cells of"Pleurotricha indica"

in life 220 x 119 /lm (± 15 % standard deviation; Table 37). Micronuclei about 3 /lm

~ Fig. 170g-j Coniculostomum indica (from AMMERMANN et al. 1987. g, h, scanning electron micrographs; i, j,
protargol impregnation). g, b) Ventral and dorsal view. Arrows mark inner right marginal row (= right mar-
ginal row 2 of Table 37). i, j) Variability of moniliform macronucleus. CC = caudal cirri, 1-6 = dorsal kineties
1-6 (kinety 6 is identical with dorsomarginal kinety 1 of Table 37); p 620.
624 SYSTEMATIC SECTION

across. Rarely specimens with more than 1 left marginal row (Fig. I70e). One supernu-
merary frontal-ventral-transverse streak, making ventral ciliature somewhat deviating
from typical I8-cirri pattern (Fig. I70a-c, e, f); see, however, taxonomy for further de-
tails. No parental dorsomarginal kineties? Dorsal kinety 6 (= outer dorsomarginal kinety)
distinctly shortened at both ends (Fig. 170d, h). Rarely 5 caudal cirri. See STEINBROCK
(1990) for the molecular biology of some macronuclear genes.
Average size of non-dividers of "e. bimarginatd' 167.3 x 84.1 ~m (from protargol
stained GI cells after complete growth; Table 37). Eight macronuclear beads in 80-85 %
of cells. Rarely additional ventral and/or transverse cirri and occasionally 1 or 2 supernu-
merary frontal cirri. Usually 2 right marginal rows, rarely a third right row with distinctly
reduced number of cirri.
Morphogenesis of cell division of the synonym, C. bimarginata proceeds essentially
similarly to that in C. monilata. For a detailed description and documentation, see KAMRA
et al. (1994). SAPRA et al. (1985) characterised the gene sized DNA molecules (desig-
nated as Pleurotricha sp.).

o c cur r e n c e and e colo g y: Locus classicus of both "Pleurotricha indicd'

(collected on February 26, 1984) and its synonym, "Coniculostomum bimarginatd' (col-
lected on March 1991) are marshes in Keoladeo Ghana Bird Sanctuary, Bharatpur, Ra-
jasthan, India (lat 27°15'N, long 77°30'E). Although these waters dry out during summer
AMMERMANN et al. (1987) could not observe any cyst formation; furthermore, they found
no stages of sexual reproduction in their cultures, which were maintained at 23°C in
Pringsheim solution with Chlorogonium elongatum as food. Generation time under labo-
ratory conditions about 10 h (AMMERMANN et al. 1987) and 9 h (KAMRA et al. 1994).

Steinia DIESING, 1866

1866 Steinia DIESING, Sber. Akad. Wiss. Wien, 52: 520 - Type (by subsequent designation, FOISSNER 1989,
P 235): Oxytricha platystoma EHRENBERG, 1831.
1866 Steinia DIESING - DIESING, Sber. Akad. Wiss. Wien, 53: 113.
1932 Steinia DIESING, 1866 - KAHL, Tierwe1t Dtl., 25: 611 (in part).
1986 Steinia DIESING, 1886 - DRAGESCO & DRAGESco-KERNElS, Faune tropicaJe, 26: 477, in part (incorrect
date).
1989 Steinia DIESING, 1866 - FOISSNER, Sber. Ost. Akad. Wiss., 196: 235.
1997 Steinia DIESING, 1866 - BERGER & FOISSNER, Arch. Protistenk., 148: 150.

C h a r act e r i sat ion: Undulating membranes in Steinia pattern. One right and 1 left
row of marginal cirri, separate posteriorly. Caudal cirri present. Six dorsal kineties. Pri-
mordia V and VI of the proter and opisthe originate from postoral ventral cirrus V/4.
Dorsal morphogenesis in Oxytricha pattern.

Add i t ion a I c h a r act e r s: Right body margin convex, left straight, margins usu-
ally converging posteriorly. Anterior end broadly, posterior narrowly rounded. Flattened
 Steinia 625

about 2:1 dorso-ventrally. Body firm. Two ellipsoidal macronuclear nodules. Contractile
vacuole in about mid-body or slightly in front of it. Cortical granules absent, pellicle col-
ourless. Cytoplasm colourless or yellowish, usually with numerous greasily shining glob-
ules and cytoplasmic crystals. Detailed description of large buccal area, see S. sphagni-
cola. Left and middle frontal cirrus about at same level, right frontal cirrus displaced pos-
teriad. Posterior-most 2 frontoventral cirri (cirri III12 and N/3) located at about same
level. Dorsal cilia 2-4 pm long. Resting cysts with spines (s. bivacuolata?). Probably
confmed to lentic or slow flowing waters.

Nom e n cIa t u rea n d t a x 0 nom y: KAHL (1932) classified Steinia as subgenus

of Oxytricha; the correct name in his revision is thus Oxytricha (Steinia). Incorrect subse-
quent spellings: Steina DIESING, 1866 (JANKOWSKI 1979, p 65); Steina platystoma (GRO-
LiERE 1978, P 309).
The correct date of pUblication is 1866 because "Band 52, Jahrgang 1865, Heft VI
bis X of the Sitzungsberichte" was only published in 1866. DIESING (l866a) did not in-
clude any species in Steinia. Somewhat later he made two new combinations, namely S.
platystoma and S. ferrugineabut did not fix a type (DIESING 1866b). Type fixation was
also omitted by later workers (FROMENTEL 1875, 1876, KAHL 1932) or Steinia was in-
cluded in Oxytricha (for example, BORROR 1972a, BOTsCHLI 1889, JANKOWSKI 1979, KENr
1882). Unfortunately, BORROR (l972a) proposed O. platystoma as type of Oxytricha.
FOISSNER (1989) thus clarified the confused situation definitively in that he fixed Oxytri-
cha granulifera as type of Oxytricha and O. platystoma as type of the reactivated Steinia.
Most species originally assigned to Oxytricha (Steinia) or Steinia do not have a frag-
mented endoral and a pit in the buccal area; FOISSNER (1989) thus transferred them to
Cyrtohymena.
The characterisation is according to BERGER & FOISSNER (1997). See chapter 2 in the
general section for detailed explanation of patterns. Morphogenesis of S. sphagnico/a re-
vealed that Steinia is very likely the sister group of Stylonychia mytilus and Coniculo-
stomum (Fig. 25a, Table 4). The strange fragmentation of the endoral in Steinia is one of
the most impressive autapomorphies within the oxytrichids. Furthermore, all species have
a conspicuous cavity (pit) at the anterior end of the buccal area so that Steinia species can
be separated by accurate live observation from Cyrtohymena and Notohymena species,
which usually also have large, anteriorly curved, and deep buccal areas. Furthermore,
Cyrtohymena and Notohymena have a flexible body (see, however, Cyrtohymena) and
(often) cortical granules and thus belong to the Oxytrichinae.

Key to species

Remarks: See also Cyrtohymena inquieta (Fig. lOla, b).

1 Transverse cirri arranged in 2 distinctly separate groups (Fig. 173a, c, k, 174k)

. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. S. sphagnico/a (p 632)
626 SYSTEMATIC SECTION

Transverse cirri not arranged in 2 groups (for example, Fig. I72k) ............. 2
2 5 transverse cirri (Fig. 172c-n) ......................... S. platystoma (p 626)
Usually only 4 transverse cirri (Fig. 175a, b) ............. S. bivacuolata (p 641)

Steinia platystoma (EHRENBERG, 1831) DIESING, 1866 (Fig. 172a:--r, Table 38)
1831 Oxytrichapiatystoma EHRENBERG, Abh. preuss. Akad. Wiss., year 1831: 118 (no figure).
1838 Oxytricha piatystoma - EHRENBERG, lnfusionsthierchen, p 365 (Fig. 172f-h).
1838 Oxytricha eurystoma - EHRENBERG, Infusionsthierchen, p 365, legend to Tafel XLI (junior objective
synonym).
1859 Oxytricha piatystoma. EHRBG. - STEIN, Organismus der lnfusionsthiere I, p 190; in part: Tafel XII, Fig.
17, 19,20 (Fig. 172i,j); not Tafel XII, Fig. 16, 18.
1866 Steinia piatystoma DIESING - DIESING, Sber. Akad. Wiss. Wien, 53: 114.
1906 Oxytricha piatystoma EHR. - EDMONDSON, Proc. Davenport Acad. Sci., II: 100 (Fig. 172r).
1932 Steinia (Oxytricha) piatystoma EHRB.-STEIN, 1859 - KAHL, Tierwelt Dtl., 25: 612 (Fig. 172a--d).
1953 Steinia piatysloma EHRENBERG-STEIN (1859) - JIROVEC, WENIG, FOTI, BARTOS, WEISER & SRAMEK-HuSEK,
Protozoologie, p 511, 513 (Fig. 172p).
1955 Sieinia piatysloma - BARWICK, BEVERIDGE, BRAZIER, CLOSE, HIRSCHFELD, PiLLAI, RAMSAY, ROBINSON, STE-
VENS & TODD, Tuatara, 5: 96 (Fig. 172e).
1969 Sieinia piatysloma - GROUERE, AnnIs Stn Iimnol. Besse, 4: 343 (Fig. 172k-Q).
1971 Steinia platystoma KAHL - CHARDEZ, Revue verviet. Hist. nat., 28: 38 (Fig. 172q).
1991 Sieinia platysloma (EHRENBERG, 1831) DIESING, 1866 - FOISSNER, BLATTERER, BERGER & KOHMANN, In-
formationsberichte des Bayer. Landesamtes fUr Wasserwirtschaft, 1/91: 336.

Nom e n c I at u rea n d t a x 0 nom y: EHRENBERG (1838) wrote that he discovered

this species on January 23, 1832. However, accepting the decision of CORLISS (1979,
P 371) to date the original description with 1831, I cite this year after the binomen.
EHRENBERG (I 831) described O. platystoma on p 118 and not on p 111 as erroneously
written in the list of synonyms in EHRENBERG (1838). KAHL (1932) classified Steinia only
as subgenus of Oxytricha; the correct name in his revision is thus O. (Steinia) platystoma.
Incorrect subsequent spellings: Oxytricha plastystoma EHRENBERG, 1830 (JANKOWSKI
1979, P 65); Oxytricha platysoma EHRBG. (DIXON 1937, P 450); Steinia platysoma EHREN-
BERG (BAMFORTH 1963, p 134); S. platyitoma (DRAGESCO 1970, P 121). Obviouslyerrone-
ously, FOISSNER & FOISSNER (I988, P 91) stated an incorrect subsequent spelling by KAL-
TENBACH (1960, P 172).
As already remarked by STEIN (1859b), S. platystoma is so insufficiently described
by EHRENBERG (1831, 1838) that any identification must be provisional. I thus agree with
FOISSNER (1989) that the redescriptions by STEIN (1859b), who mixed it up with Cyrtohy-
mena inquieta (see Fig. 100a, b), and KAHL (1932) are authoritative. Both STEIN and
KAHL draw transverse cirri which are not arranged in 2 distinctly separated groups. This

Fig. 172a-j Sieinia platystoma (a--d, from KAHL 1932; e, from BARWICK et al. 1955; f-h, after EHRENBERG ~
1838; i,j, after STEIN 1859b. a-j, from life). a, b) Buccal area in ventral view and in cross section. c, d) Ventral
view of a slender and a short specimen, c = 160 11m, d = 120 J.lm. e) Ventral view, 102 11m. f-j) Dorsal and
ventral views, f-h = 105 11m, i,j = about 150 J.lm. BC = buccal cirrus, e = endoral, G = cavity.
Steinia 627
 Rj
00

tll

!l)

: ~~ ~ f: ~ o~ ~ E 0
~

<>
· '\ ~.....\. <>
''" "'\P ~ ...•.• \) CI.l
-<
~ CI.l
\. . . \W ' '"' . ~
::1 '.
: .:~,~·i : I \~ .lot, .~, \ l.~e ~ :. k. ~I }I n
"""l
~ 0~"j 5
!' ., f,~.
\ Q 'tV 0/ Z
O
0 oWl!~ ~.G el . : · "0 Il> ~I)@ .,
• .. 0" o · "
.. IP \
•••
• •
.. 0
(1)
"
'" •
"_~ """____0 Q
0
• • ., 0 ()

k ( > " 0/ m n
" p

Fig. 172k-n Steiniaplatystoma (from GROUERE 1969. k-n, ventral infraciliature after protargol impregnation, sizes not indicated). k) Non-dividing specimen. Arrow
marks frontoventral cirrus UII2 which is arranged more posteriorly than all other frontoventral cirri. I-n) Early morphogenetic stages (reinvestigation necessary); p 626.
 Steinia 629

.·~~!IiIIt-G

Fig. 172o-r Steiniaplatystoma (0, from GROLlERJ!.I969; p, from JfROVEC et aI. 1953; q, from CHARDEZ 1971; r,
from EDMONDSON 1906. 0, protargol impregnation; p-r, from life). 0) Buccal area. Arrowheads mark frag-
mented endoral. J)-r) Ventral views, p = 100 ~m, q = 115 ~m, r = 140 ~m. G = cavity; p. 626.

difference to S. sphagnicola is confIrmed by GROLIERE (1969) on protargol impregnated

material. The DRAGESCO (1970) population is probably not conspecific with S. platystoma
because he draws neither a cavity and a helical structure in the anterior part of the buccal
area nor a fragmented endoral. Furthermore, the arrangement of the frontoventral cirri de-
viates. In my opinion, DRAGEsco'S (1970) population is identical with Cyrtohymena
primicirrata (Fig. 98h-j). Oxytricha platystoma sensu Roux (190 I; Fig. 10 1e) and sensu
BUITICAMP (l977b; Fig. 99j) are classifIed as Cyrtohymena aestuari and C. candens re-
spectively. Oxytricha platystoma sensu Gu & ZHANG (1992) is very likely Cyrtohymena
inquieta. The HORvArn (1933) population is rather surely identical with Histriculus his-
trio (Fig. 177m', n). The redescriptions by FROMENTEL (1875), DUMAS (1929, Fig. 236e),
and BIERNACKA (1959; Fig. 225s) are insufficient. A redescription of S. platystoma must
comprise a morphometric characterisation and an illustration of the dorsal infraciliature.
630 SYSTEMATIC SECTION

M 0 r ph 0 log y and b i 0 log y: After EHRENBERG (1831, 1838) in life about

105 !lm long, after STEIN (1859b) about 150 !lm, after KAHL (1932) 80-130 !lm. Accord-
ing to GROLIERE (1969) after protargol impregnation 110-170!lm long (mean = 150 !lm;
n = 100). About 2-2.5 times as long as broad. Macronuclear nodules after protargol im-
pregnation 30-40 x 15-20 !lm. Two micronuclei, IO!lm across on average (GROLIERE
1969). Cytoplasm yellowish. Cytopyge slightly subterminally on left body margin. Swims
about 520 !lm S-l (BULLINGTON 1925).
Adoral zone of membranelles about 40 % of body length, consists of about 42-45
membranelles (Fig. 1720). Frontoventral cirrus 11112 somewhat behind level of frontoven-
tral cirrus IV/3 (Fig. 172k, arrow). Two rightmost transverse cirri (cirri VII and VIII)
protrude about 2/3 beyond posterior end of cell; left transverse cm not (STEIN 1859b) to
distinctly (KAHL 1932) protruding. About 22 right and 17 left marginal cirri.
Very likely, the formation of the oral primordium commences near the left transverse
cirrus (Fig. 172I-n). CHAKRABORTY (1967) studied the macronuclear organisation and the
ultrastructure of "Oxytricha platystoma". The microphotographs and the locality (lettuce
leaves!), however, do not support his identification.

o c cur r e n c e and e colo g y: Rare, but if present then usually abundant and per-
sistent in stagnant (especially peat-bogs) and running waters (KAHL 1932). Probably cos-
mopolitan.
Locus classicus is a marsh near Berlin, Germany (EHRENBERG 1831, 1838). Records
documented by illustrations: freshwater in the Ardennes, Belgium (CHARDEZ 1971);
marshy meadow in Tharandt near Dresden, Germany, marshy ditch near Prague and vil-
lage of Wirschowitz (Czechoslovakia) in a brook on sites choked with mud (STEIN
1859b); running waters in Czechoslovakia (JiROVEC et al. 1953); forest pond in France
(GROLIERE 1969); freshwater in Iowa, USA (EDMONDSON 1906); New Zealand (BARWICK
et al. 1955).
Records from freshwater habitats not substantiated by illustrations: oxbow of Danube
River in Vienna, Austria (FOISSNER & FOISSNER 1988, HEw 1937); ditch, brook, rivers,
wet meadow, and fish pond in Bulgaria (DETCHEVA 1979b, 1993, DETSCHEWA 1972, Rus-
SEV et al. 1976); rain puddle near BrUnn, Czechoslovakia (SPANDL 1926a); pond and wet
mosses in Czechoslovakia (HAsSDENTEUFEwvA-MoRAvcovA 1955, TIRJAKovA & MATIS
1987a); creek in England (WHITELEGGE 1889); infusion of willow branches, Estonia (JA-
COBSON 1928); peat-bog in France (GROLIERE 1977, 1978); lentic water and Schussen
River, Germany (SCHMIDT 1916, WETZEL 1928a); Danube River and other sites in Hun-
gary (BERCZIK 1966, KREpUSKA 1917, 1930); spring in Italy (STELLA 1947); lake in Italy
during July (LaNGill 1895); with low abundance in a beta- to alphamesosaprobic brook,
up to 32 000 indo I-I in an alphameso- to polysaprobic water course of the Val Parma, and
in other running waters in northern Italy (MAroNI 1980, 1983, 1984, MADaNI & GHETTI
1977, 1980); Poland (JAWOROWSKI 1893, WRZESNIOWSKIEGO 1866); rare in mud infusions
from the Danube River in Romania and in wells (SPANDL 1926a-c); Sweden (QUEN-
NERSTEDT 1869); between Sphagnum in alpine bogs (up to 1900 m above sea level) in
 Steinia 631

Switzerland (HEINIS 1945, MESSIKOMMER 1954; further records from Switzerland: ANDRE
1912, 1916, BOURQUIN-LINDT 1919, MERMOD 1914, PERTY 1849a, 1852b, SAKOWSKY-
CAMPIONI 1906, THIEBAUD 1906, WOLFF 1948); USSR (WEISSE 1848a, b); Canada (PUYTO-
RAC et al. 1972); well in Yugoslavia (KARAMAN 1935); abundant in a three months old
culture of floating Spirogyra from Lake Mirror, Ohio, USA (SHAWHAN et at. 1947, STEHLE
1920); between detritus at the bottom of shallow pools in Trenton, New Jersey, USA
(STOKES 1884; = locus classicus of Trachelophyllum tachyblastum); South Eastern Lou-
isiana, USA (BAMFORTII 1963); up to 3400 ind.l· t in a Pennsylvanian pond (USA) from
August to September (WANG 1928); Brazil (CUNHA 1913, 1916).
Records from terrestrial habitats are unreliable; very likely it was confused with Cyr-
tohymena species which are common in soil and mosses (for example, DAWSON &
MITCHELL 1929, DECHEvA 1970, DETCHEVA 1972b, DIXON 1937, TIRJAKovA & MATIS
1987a). FOISSNER (1987a) listed the record by BUITKAMP (1977b), a population which is
likely identical with Cyrtohymena candens.
Feeds on algae (diatoms), flagellates, and small ciliates. GROLIERE (1969) cultured
Steinia platstoma in Eau d'Evian with Chlamydomonas sp. as food. Biomass of 106 indi-
viduals about 75 mg (FOISSNER et al. 1991).
DETCHEVA (1972a, 1975c) found Steinia platystoma in the Ogosta River, Bulgaria,
under the following conditions: frequency 1.8 %, pH 7.7, 12.7 mg 1.1 O2 (100 % satura-
tion), 5 mg 1.1 BODs, 4 mg 1.1 NHl, 11.7 mg 1.1 NOl'-N, 0.04 mg 1.1 N02'-N, 0.12 mg I-I
Fe2+_ GROuERE & NJINE (1973) counted about 3000 ind.l- I in a French peat-bog in May
under following conditions: 13 °C, pH 5.0, 1.5 mg 1"1 O2 (15 % saturation), 0.033 mg I-I
P04--P, 0_77 mg I-I NH/-N, 3.2 mg I-I N03--N. MADaNI & GHETII (198Ia) found it occa-
sionally in three alphamesosaprobic to polysaprobic sites of the Torrente Stirone, north-
ern Italy, at 3.8-19 °C, pH 7.4-8.5,26-1094 mg 1"1 C1", 6.0-12.1 mg 1-1 0 2,22-71 mg 1"1
COD, 0.5-3.7 mg I-I NH3, 0.03-2.2 mg 1"1 NOi-N, 0.5-2.6 mg 1"1 NOl--N, 0.22 to
lAO mg I-I POi--P. Hydrogen sulphide is not tolerated (WETZEL 1928b). MOCKE (1979)
counted 28 000 indo I-I in a Sphagnum-pond in Germany; when the pond began to dry up
in late August the abundance briefly increased to 104 x 106 indo 1"1; in moist Sphagnum
84 000 ind.l-I were present STATKEWITSCH (1904) studied the effect of electrical current
on Steinia platystoma.
Steinia platystoma indicates beta- to alphamesosaprobic conditions of water quality
(b-a; b = 6, a = 4, 1= 3, SI = 2.4; SLADECEK et al. 1981, WEGL 1983, FOIssNER 1988a).
This classification deviates somewhat from the statement by KAHL (1932) that it occurs
only in rather putrescent waters. The autecological data by GROLIERE & NJINE (1973) and
MADaNI & GHETII (1977, 1981a) also indicate that it obviously prefers lower quality sites
(15 % O2 saturation!, 22-71 mg I-I COD, alphameso- to polysaprobic conditions). Thus,
the saprobic classification has to be verified on reliably determined populations (FOISSNER
et al. 1991).
632 SYSTEMATIC SECTION

Steinia sphagnico/a FOISSNER, 1989 (Fig. 14b, 20£, 21f, 22b, 173a-y, 174~
Tables 3, 4, 38)

1986 Steinia platystoma (EHRENBERG-STEIN, 1859) - DRAGESCO & DRAGESCO-KERNEIS, Faune tropicale, 26:
479; in part: Planche 142, Fig. E; not Fig. A-D (misidentification; Fig. 173k).
1989 Steinia sphagnicola FOIssNER, Sber. Ost. Akad. Wiss., 196: 235 (Fig. 173a--i; 1 slide ofholotype speci-
mens and 3 slides of paratype specimens are deposited in the OberOsterreichische Landesmuseum in
Linz, Upper Austriar
1994 Steinia sphagnicola - FolSsNER, PETz, UNTERWEGER, HERzOG, SIMONSBERGER, KRAuroARTNER & ZANKL,
Katalog des OberOsterreichischen Landesmuseums Linz, 71: 109 (colour-microphotograph of a
protargol-impregnated specimen).
1996 Steinia sphagnico/a FolSSNER, 1989 - Voss & FolSSNER, Europ. J. Protistol., 32: 31 (Fig. 173j-y,
174a-j).

Nom e n c I at u rea n d t a x 0 nom y: AEsCIIT (1994, p 50) mentioned this species,

obviously erroneously, uilder the binomen "Cyrtohymena sphagnicola (FOISSNER, 1989)".
DRAGESCO & DRAGESCO-KERNEIS (1986) mixed up three species, namely Cyrtohymena
primicirrata, Steinia platystoma, and S. sphagnicola. The latter two can be separated
unequivocally by the different arrangement of the transverse cirri. According to EIGNER
(1997), Steinia sphagnicola belongs to the Parakahliellidae (details on this classification,
see Addendum 3).

M 0 r p hoi 0 g y and b i 0 log y: In life about 130-160 x 50-80 !lm (FOISSNER

1989), cultured population from Germany 160 x 80 !lm (Voss & FOISSNER 1996). Ellipti-
cal, ventral surface flat, dorsal side distinctly arched. Contractile vacuole with conspicu-
ous canals during diastole. Pellicle fragile, many specimens break during impregnation.
Movement moderately rapid.
Adoral zone of membranelies about 50 % of body length, distance between mem-
branelles conspicuously large (about 2 !lm). Buccal area broad and deep, that is, extend-
ing almost to dorsal surface (Fig. 173e, f, i, I), distal portion semicircularly curved to left
and spirally rolled up to form peculiar circular cavity (pit) also extending to dorsal sur-
face, so that it appears very bright at low magnification. Function of cavity unknown, nei-
ther in life nor after protargol impregnation any specific structure recognisable. Paroral
less distinctly curved than buccal area, commences left of left frontal cirrus and extends
left of narrow ridge (buccal lip) to proximal fifth of adoral zone of membranelles, made
of zigzagging basal bodies; cilia 25 !lm long, forming a conspicuous membrane (Fig.
173e, t). At distal end of this lip an elliptical orifice leading into the cavity (Fig. 173f, I).
Endoral sinusoidal and fragmented (!), made up of a single row of basal bodies, com-
mences about at same level as paroral but terminates more posteriorly, namely near
proximal end of adoral zone of membranelies; cilia of anterior-most fragment in life
about 17!lm long, beat into cavity; cilia of small fragments about 5 !lm, cilia of
posterior-most and longest fragment about 10 !lm long; from each fragment a strong bun-
dle of fibres extends rightward to centre of bow formed by paroral (Fig. 173i, I).
 ~~" :5)
3/ 4\ ' ( '6 J
. ~ ..,./" / ....
..
), )2), \'/
f _ ,~/., \ I, \
IP \1
13 ~
l - ./ (--4 '~""/ \' '\ J'f 'II
\ \ , \ J )
\'
\" ( . (" ,) '
• l .. ,'\
\... \' ')' ~\' . ,/ I,/
\'
\'
/\' \ ,( I r
.' \ r \'
,
p \, ,I (
l' \, (' ,J
I' '\ ~
(\)
r ')
r S·
~.

I
't"
~
-.,..,
-.- ,J
•...--- ,/ •I
J J
•/ •
t ,)
-- . -
, }
b

Fig. 173a-d Steinia sphagnico/a (from FOISSNER 1989. a, b, from life; c, d, protargol impregnation). a, b) Ventral and lateral view, a, b" ISS 11m. c, d) Ventral and dor-
sal infraciliature, 140 11m. Arrow marks posterior postoral ventral cirrus (cirrus VI3). Note the curious arrangement of the transverse cirri. e" fragmented endoral, G = 0\
W
W
cavity (function unknown), p = paroral, PRVC = right pretransverse ventral cirrus, TCI, TCr = left and right transverse cirrus, 1-6 =dorsal kineties 1-6; p 632.
634 SYSTEMATIC SECTION
 Steinia 635

AZM

AFC EM
BC . "
PM "
•
,.
PFC....
,
,
POC,
,

RMR-'
; TC
,
VC ..
, •
TC I.

Fig. 173j, k Steinia sphagnico/a (from Voss & FOISSNER 1996. j, scanning electron micrograph; k, protargol
impregnation). Ventral views, bar = SO 11m. The right half of the buccal field is covered by the cilia of the pa-
roral (arrow in j). Arrowhead in (k) marks postoral ventral cirrus V/3. Explanation of original labels: AFC =
right frontal cirrus, AZM = adoral zone of membranelles, BC = buccal cirrus, EM = endoral, LMR = left row
of marginal cirri, MA = macronuc1ear nodule, Ml = micronucleus, PFC = posterior frontoventral cirri (cirri
IllI2 and IV!3), PM = paroral, POC = postoral ventral cirrus Vl4, RMR = right row of marginal cirri, TC =
transverse cirri, VC = pretransverse ventral cirrus VJ!2; p 632.

~ Fig. 173e-i Steinia sphagnico/a (from FOISSNER 1989. e, f, interference contrast; g, bright field; h, i, protargol
impregnation). e) Ventral view. Arrow marks cavity. t) Buccal area in ventral view. Large arrow denotes paro-
ral, small arrows mark fragmented endoral. g) Resting cyst. h) Ventral view. i) Buccal area with fragmented
endoral (arrows). Dc = basal body pairs of dorsal kineties, G = cavity; P 632.
636 SYSTEMATIC SECTION

~ EM

,-

Fig. 1731, m Steinia sphagnicola (from Voss & FOISSNER 1996. I, scanning electron micrograph; m, protargol
impregnation). I) The curious buccal pit (cavity) is recognisable only in specimens with paroral cilia moved
upwards (bar = 20 Jlm). m) Oral area showing uninterrupted paroral and fragmented endoral (arrows; bar =
20 Jlm). Explanation of original labels: AZM = adoral zone of membranelles, EM = endoral, LMR = left row
of marginal cirri, MA = anterior macronuc1ear nodule, P = cavity (buccal pit), PM = paroral; p 632.

Invariably 18 frontal, ventral, and transverse cirri in 14 (FOISSNER 1989) and 25

(Voss & FOISSNER 1996) specimens. Frontal, ventral, and marginal cirri in life about
30 J-lm, transverse and caudal cirri about 35 J-lm long. Transverse cirri slightly enlarged,
left 3 cirri arranged in a rather steep row, posterior-most cirrus of this row located slightly
in front (!) of level of right pretransverse ventral cirrus (Fig. 173c, I); this pretransverse
ventral cirrus and the 2 right transverse cirri are arranged in an almost longitudinally ex-
tending row. Right marginal row begins at level of right frontal cirrus, terminates slightly
subterminally, left one terminates at posterior end. Caudal cirri located slightly right of
median (in ventral view), arranged at posterior end of dorsal kineties 1,2, and 4. Kineties
1-4 about of body length, kineties 5 and 6 (dorsomarginal rows) only 112 and 113 of body
length respectively (Fig. 173d).
Resting cysts yellowish, spherical to slightly ellipsoidal, in life 48-62 x 45-56 J-lm
(mean = 53.2 x 51.7 J-lm, without spines; n = 12). Ectocyst 1.4 J-lm thick, with many about
 Steinia 637

()

()

,
0'
o
(j
()

l)
Q
()
'" l) (]
o
a
()

0 0 n
Fig. 173n, 0 Steinia sphagnico/a (from Voss & FOISSNER 1996. n, 0, protargol impregnation). Very early di-
viders, n = 115 )lm, 0 = 118 )lm. Arrow in (n) marks oral primordium close to transverse cirrus WI. OP = oral
primordium; p 632.

7 ~m long, hollow spines (Fig. 14b). Older cysts with detritus and bacteria between
spines, indicating that a thin mucous layer is present.

M 0 r ph 0 g e n e sis (Fig. 173n-y, 174a-j): This process is described in great detail by

Voss & FOISSNER (1996) and includes many scanning electron micrographs. Briefly, im-

Fig. 173p-y Steinia sphagnico/a (from Voss & FOISSNER 1996. p-y, protargol impregnation). p) Early divider ~
showing anlage originating from oral primordium (arrow). Arrowhead marks disaggregating postoral ventral
cirrus V/4. q-u) Frontal-ventral-transverse cirri anlagen 5 and 6 originate from postoral ventral cirrus V/4 in
both proter and opisthe. A small field of basal bodies splits off from the dissolving postoral ventral cirrus V/4,
migrates anteriorly (s, t, arrows), and organises to proter's anlagen 5 and 6 (u, v). v-y) Anlagen 3 and 4 of the
proter originate from frontoventral cirri IIU2 and IV/3 (v, X, arrow). The opisthe's anlagen 3 and 4 originate
from the oral primordium (x, arrowhead) and, respectively, from postoral ventral cirrus lV/2 (y), which dis-
 638 SYSTEMATIC SECTION

<::><::>
~<::>
t:$
0
"tl ./
o /.t~ ~
", ,- §
\'~ '.~
,
1
00

·i .:
0
I
'~ Fe
'"
"
"
.. ~F. ot0> 00 DO :
I
1
-:t
."

'.
I I
" ~'~o
c>
I
I
1
1
"
~

, c> I I
c> I I
I I
POC3"
c> I
I
I l'l"""'- I
c> I 1
0 i POC1 : I 1

. .._'
c>
oi I
0 " 1I '9
I ~ O:~
0
0
c> 0 1
1 0 1 .f
1
1 ".
I
1
0 Q, rI SI
I u

...
()
() Q
c ll •
173p 4~

"
'"
4 3

65t..~
" 0

.
! ,
I .
\

I
.,

i. \.;. .
~

~ "i••
.:. ~'..::
.
.....:'",.
"

6, 5
'.:t ; A
~:
. . . . . ;!
J..!
': :-
v () <> POC3D ~.
~ .. i <> /) .~
....., ...
coO
<> a a
"t
<> 0 a POC1
VI
I
w ~ 00 ~ X
QQ tI <::> <::>

solves later than postoral ventral cirrus V/4. The whole process is accompanied by cortical growth as indicated
by increasing distances between postoral ventral cirri IV12 and V14 and between frontoventral cirri and pos-
toral ventral cirri. Postoral ventral cirrus V13 is not involved in primordia formation; an apomorphy of the Sty-
lonychinae. Bar = 10 J.(m. Explanation of original labels: Fe = frontoventral cirri IIII2 and 1V/3 (obviouslyer-
roneously designated anterior frontal cirri in the original paper), POCI-3 = postoral ventral cirri 1V/2, Vl4,
V/3, 3--6 = frontal-ventral-transverse cirri anlagen; p 632 (begin oflegend, see p 637).
 ~<;G ~'
()()G
<:::>0
<:>~ • 00
)
o 0~":) ., o°
. ""'-'
~ 0 ~/./ o • .' I.' o V/ ~
AD 5 ,6n
o .' • 4 I
I ~~
~ff \. ~ " f
, •':f'\
Of·.i ~
o :~
.",t .
'\
.\~( ~" ~
,\\\ ~"
( ~:f:.· \(f)
\~\' - / l ""
~
. '\'.
- \ -"
t ,
\,
·~\.'....\.~.. §i " •
...
. ·\1"''..
« "
.' ~
(\)
, ..I
:- • t:J .~. " s·
o
poe1 ~,'
o
'. ~. " ., .
•,•"
is'
,.s:" " <:l
" 0
0 0 0 0 .,
" ~ :j v
0 a 0 (\ () a"
"" ~~.
0 0 " () p
" ~
a 0
()
&1
"o
" -/
cOo",~~ a "• b

Fig. 174a-c Steinia sphagnicola (from Voss & FOISSNER 1996. &-c, protargol impregnation). a) Middle divider (138Ilm) with 7 anlagen in the opisthe because the left
frontal cirrus has separated from the anlage of the undulating membranes. The parental paroral commences to reorganise (arrow) and the buccal cirrus (open arrow) and
some right marginal cirri in the prospective division furrow (arrowhead) disaggregate to anlagen. b) Middle divider showing anlagen (arrows) for dorsal kinety 5 which
originate from parental marginal cirri. Arrowheads mark primordia in left marginal row. c) Middle divider showing segregation of new cirri and reorganisation of the
proter's undulating membranes. Note that fragmentation of endoral gradually disappears. Parental structures white, new ones black. ADS.6 = primordia of dorsomarginal 0'\
w
kineties (dorsal kineties 5, 6), POCI = postoral ventral cirrus 1Vfl; p 632. \0
640 SYSTEMATIC SECTION

o
o

Fig. 174d, e Steinia sphagnico/a (from Voss & FOISSNER 1996. d, e, protargol impregnation). Middle dividers
(d = 150 lim, e = 175 lim) showing developing dorsal kineties 5 and 6 (arrowheads), migration of cirri, and re-
organising undulating membranes in the proter. The fragmentation and sinusoidal bending of the endoral have
disappeared. Arrow in (d) marks postoral ventral cirrus V/3, which is not involved in primordia formation (an
apomorphic character of the Stylonychinae). Parental structures white, new ones black; p 632.

portant morphogenetic details are identical to in the Sty/onychia mytilus complex and in
Coniculostomum suggesting monophyly of these taxa (Fig. 25a). For details, see legends
to figures and Table 4.

o c cur r e n c e and e colo g y: Not very common in freshwater. Locus classicus is

the village ofKoppl, Salzburg, Austria, where FOISSNER (1989) discovered S. sphagnicola
in a Sphagnum pond ofa bog. FRYD-VERSAVEL (in DRAGESCO & DRAGESCO-KERNEIS 1986)
found it in Paris, France, and Voss & FOISSNER (1996) collected it in spring in a small
tributary (Windach) to the Amper River near Munich, Germany, about 100 m above the
sewage treatment plant of the village of Eching. CZAPIK & FYDA (1995) found Steinia
sphagnico/a in two Polish lakes.
 Steinia 641

""""
,, •
"
""" •••
.
til

-..•
til
til
••
",
~
til

,'
.", , ".-: :

--
", .. ~.

..-- •
",

'" t' P
0

.• ti
p
'"
,,, ~.'.~
,0
"
, • "~, ~

,, , • : ,:. .'". I§
~ ....
·4

.... Eff ,
,.
,,,
~

,,
,,
,,, ·
'. ·
.
•,
,
•
• '",
,,
I
,,, F -
'"
~, ..
...
~
0
'.,
0 I
'~" ,"
0

.
,,"
0 0
..
0

,,'. '
"" ..
()

0° -
0

0 IJ ~
t
"0 D
f
'f
~
o 0 0 0 °0 <> 9
Fig. 174f, g Steinia sphagnico/a (from Voss & FOISSNER 1996. f, g, protargol impregnation). t) Late divider
(190 11m) showing intersecting undulating membranes. The parental cirri (white) which were not involved in
the formation of anlagen are gradually reabsorbed. g) Very late divider, 165 11m. The anterior end of the pam-
raI becomes curved, the endoraI fragments, and most parental cirri have been reabsorbed; p 632.

Steinia sphagnicola feeds on other protists (Astasia sp., Dexiostoma campylum) and
probably also on bacteria. ForSSNER (1989) and Voss & ForSSNER (1996) cultured it in let-
tuce medium with some squeezed wheat grains to support growth of bacteria and small
ciliates. Biomass of 106 individuals about 150 mg.

Steinia bivacuolata (GELEI & SZABADOS, 1950) FOISSNER, 1989

(Fig. 175a-c)

1950 Oxytricha bivacuolata GELEI & SZABADOS, AnnIs bioI. Univ. szeged., 1: 263, 283 (Fig. 175a--c).
1989 Steinia bivacuolata (GELEI & SZABADOS, 1950) - FOISSNER, Sber. Ost. Akad. Wiss., 196: 235.
642 SYSTEMATIC SECTION

• ~
• • ~
•
, -.
~"
:::::: "
.."
• ,,
.
• • ,
,
-
•

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 Steinia 643

T a x 0 nom y: I agree with FOISS-

NER (1989) that this is a true
Steinia (and not a Cyrtohymena)
because GELEI & SZABADOS (1950)
draw the conspicuous cavity in the
anterior part of the buccal area It
differs from S. platystoma, with
which it was synonymised by BOR-
ROR (1972a), in that it usually has
only 4 transverse cirri. Steinia
sphagnicola has a different ar-
rangementof transverse cirri.
Steinia bivacuolata should not be
confused with Cyrtohymena tetra-
cirrata, which also has only 4
transverse cirri (Fig. 102a-i). Fig. 175a--c Steinia bivacuolata (from GELEI & SZABADOS
However, latter species is confmed 1950. a, opalblue stain after BRESSLAU; j, k, sublimate
fixation). Ventral and dorsal infraciliature, a = 150 Jim, j, k =
to terrestrial habitats, whereas S. 105 Jim. Differs from the other two species of Steinia in that
bivacuolata was discovered in it has only 4 transverse cirri; p 64l.
freshwater. Detailed redescription
necessary.

M 0 r p hoi 0 g y and b i 0 log y: Large specimens in life (?) about 150 11m long,
about 2.4 times as long as broad (Fig. 175a). Anterior portion of cell slightly turned to
left and somewhat broader than posterior part. However, outline of small specimens usu-
ally elliptical, causing GELEI & SZABADOS (1950) to assume that they are another variety
or even species. I also doubt that the specimen shown in Figure 175b is conspecific with
that of Figure 175a because the body shape is different and no cavity is shown. Each
macronuclear nodule with 2 hemispherical micronuclei, smaller specimens with only 2
micronuclei. Perhaps 2 contractile vacuoles; one about in mid-body, another somewhat in
front oflevel of cytostome. Rapid creeping or slow or fast swimming.
Adoral zone of membranelles about 40 % of body length, consists of approximately
35 membranelles. In anterior part of buccal area an almost transversely arranged row of 6

~ Fig. 174h-k Steinia sphagnicola (h-j, from Voss & FOISSNER 1996; k, from DRAGESCO & DRAGESco-KERNbs
1986. h-k, protargol impregnation). h) Early postdivider, 80 Jim. The curved parorai crosses (intersects) the
endorai, which fragments and becomes sinusoidal. i) Late postdivider (110 J.lm) showing how the paroral
moves rightward, producing the Steinia-specific arrangement of the undulating membranes. The buccal pit
(cavity) develops. j) Middle divider (145 J.lm) showing separation and development of dorsal kinety 4 from ki-
nety 3, that is, dorsal morphogenesis is in Oxytricha pattern. k) Ventral infraciliature, size not indicated. Arrow
marks paroral, arrowhead fragmented endoral. AD. = young dorsal kinety 4, P = cavity (buccal pit); P 632.
644 SYSTEMATIC SECTION

Table 38 Morphometric data of Steinia sphagnico/a (line I, from FOISSNER 1989; line 2, from Voss & FOISS-
NER 1996). All data are based on protargol-impregnated specimens. All measurements in micrometres. CV =
coefficient of variation (in %), Max = maximum value, mean = arithmetic mean, Min = minimum value, n =
sample size, SD = standard deviation

Character mean SD CV Min Max n

Body, length 140.7 11.3 8.0 122 157 14
2 143.0 17.5 12.2 Il2 176 25
Body, width I 60.4 7.6 12.6 46 72 14
2 68.5 10.3 15.0 55 99 25
Macronuclear nodule, length I 21.l 2.8 13.5 17 28 14
2 25.9 2.9 11.2 22 28 25
Macronuclear nodule, width I 11.2 2.2 19.6 8 15 14
2 10.4 1.9 18.3 9 18 25
Macronuclear nodules, distance 1 17.7 4.8 27.4 11 28 14
between 2 9.9 3.9 39.4 4 20 25
Micronuclei, number 1 1.7 1 2 14
2 2.4 0.6 25 2 4 25
Micronucleus, diameter 1 3.9 0.4 10.0 3 5 14
2 3.8 0.9 23.7 2 7 25
Adoral zone of membranelies, I 66.9 4.7 7.1 56 72 14
length 2 70.0 10.2 14.6 53 94 25
Adoral membranelles, number 36.4 2.3 6.3 31 39 14
2 36.4 2.3 6.3 32 40 25
Endoral, number of fragments 1 10.4 1.6 15.4 7 13 14
2 10.4 0.9 8.7 9 12 25
Distance between transverse 9.3 2.4 25.8 6 13 14
cirrus VII and end of cell 2
Right marginal row, number 21.9 1.3 6.1 20 24 14
of cirri 2 23.2 1.1 4.7 21 25 25
Left marginal row, number 1 19.6 2.3 ll.8 14 23 14
ofeirri 2 20.0 1.3 6.5 18 24 25

cilia, very likely a part of fragmented endoral (Fig. 175a). Transverse cirri protrude dis-
tinctly beyond posterior end of cell. Apparently only 2 caudal cirri.

o c cur r e n c e and e colo g y: Locus classicus of Steinia bivacuolata is a puddle

on the Ski. Istvan Square in Szeged, Hungary (GELEI & SZABADOS 1950). Feeds on auto-
trophic and heterotrophic flagellates. Biomass of 106 individuals about 150 mg. Not
found since.

Species indeterminata

SteiniaJenestrata LEPSI, 1957, Trav. Mus. Hist. nat. Gr. Antipa, 1: 90 (Fig. 225p, q). Re-
marks: In life 100-122 x 33-43 ~m. Oval, left margin usually straight, right convex. Not
contractile. Two spherical to ellipsoidal macronuclear nodules. Extrusomes or cortical
 Histriculus 645

granules lacking, cytoplasm densely granulated. Always swimming, does not jump. Ado-
ral zone of membranelles about 113 of body length. Buccal area large and deep, undulat-
ing membranes anteriorly curved. Frontal cirri distinctly enlarged, in "common arrange-
ment". Marginal cirri distinctly separate. About 5 transverse cirri, protrude distinctly be-
yond posterior end of cell. LEPSI (1957) found numerous specimens in a slightly mouldy
smelling, 11 month old culture of mosses (not Sphagnum) from a pond with distinctly
acid, slightly brown water near the mountain lake "St. Anrui" in Romania. In spite of the
extensive information on this taxon, I believe that it can never be identified unequivocally
because the drawings are too superficial and the description is too general. FOISSNER
(1989, P 239) transferred it to Cyrto!rymena, C.fenestrata.

Insufficient redescriptions
Oxytricha platystoma, EHR - DUMAS, 1929, Microzoaires, p 70 (Fig. 236e). Remarks:
Probably a redrawing from FROMENTEL (1876). Bog in France.

Oxytricha platystoma - FROMENfEL, 1876, Etudes Microzoaires, p 266, Planche xm, fig.
6, 6a. Remarks: The drawings do not give any indication that the determination is correct.

Steinia platystoma EHR. - BIERNACKA, 1959, Polskie Archwm Hydrobiol., 5: 57 (Fig.

225s). Remarks: The shape of the buccal area does not indicate a relationship to Steinia
or Cyrtohymena. Activated sludge plant of Danzig, Poland.

Histriculus CORLISS, 1960

1878 Histrio STERK!, Z. wiss. ZOO!., 31: 56 (junior homonym) - Type (by monotypy): Histrio steinii STERK!,
1878.
1882 Histrio, STERK!- KENT, Manual infusoria II, p 789.
1932 Histrio STERK!, 1878 - KAHL, TierweIt DtI., 25: 614 (in part; see nomenclature and taxonomy).
1960 Histriculus CORLISS, 1. Protozool., 7: 275 (in part) - Type (original designation): Histrio steinii STERK!,
1878.
1972 Histriculus CoRLISS, 1960 - BORROR, J. Protozool., 19: 14 (in part).
1974 Histriculus CoRLISS - STILLER, Fauna Hung., 115: 149.
1975 Parahistriculus GROLIERE, Protistologica, II: 493 - Type (by monotypy): Parahistriculus minimus
GROLIERE, 1975.
1997 Histriculus CORLISS, 1960 - BERGER & FOISSNER, Arch. Protistenk., 148: 149.

C h a r act e r i sat ion: Undulating membranes in Oxytricha pattern. One right and
one left row of marginal cirri, posteriorly confluent. Six dorsal kineties. Caudal cirri ab-
sent. Dorsal morphogenesis in Oxytricha pattern.
646 SYSTEMATIC SECTION

Add i t ion a 1 c h a r act e r s: Two macronuclear nodules. Contractile vacuole about

in mid-body. Pellicle and cytoplasm colourless. Dorsal cilia in life 2-4 Jlm long. Proba-
bly confined to freshwater and marine habitats.

Nom e n c I a t u rea n d t a x 0 nom y: Histriculus CORLISS is a new name for the

preoccupied name Histrio STERKI.Parahistricus minimus in GROLffiRE (1975, P 481) is an
incorrect original spelling. KAHL (1932) classified Histrio as subgenus of Oxytricha; the
correct name in his revision is thus Oxytricha (Histrio). Incorrect subsequent spellings:
Historio STERKI and Historia sp. (WANG 1925, P 52 and explanation of figures of plate
XI); Histriculux vorax (DIVE 1973, p 348); Histrichulus vorax (BANINA 1983, P 81).
STERKI (1878) established Histrio to separate Stylonychia histrio (basionym Para-
maecium histrio; Histrio steinii is probably only a new name for that species to honour
STEIN, who provided the first detailed description of Histriculus histrio) from other Stylo-
nychia species because the marginal cirral rows are posteriorly confluent and caudal cirri
are lacking. However, later KAHL (1932) and BORROR (1972a) added species with incon-
spicuous caudal cirri, obviously assuming that STEIN (1859b) and STERKI (1878) over-
looked them. Morphogenetic studies proved that such species indeed have inconspicuous
caudal cirri, especially "Histriculus muscorum" (now Sterkiella histriomuscorum), one of
the most widespread oxytrichids. A detailed redescription of the type species of Histricu-
Ius, H. histrio (GSCHWIND 1991; see also BERGER & ForSSNER 1997) exactly matched the
original diagnosis by STERKI (1878), demanding that species with caudal cirri assigned to
Histriculus by KAHL (1932) and BORROR (1972a) be referred to a new genus, Sterkiella
(ForSSNER et al. 1991).
The characterisation is according to BERGER & ForSSNER (1997). For a detailed de-
scription of patterns, see chapter 2 in the general section. The characterisation is based
mainly on the morphology of the type species, the only morphologically completely de-
fmed species. The validity of the other species listed in the key has to be checked by de-
tailed redescriptions.
Histrio inquietus STOKES belongs to Cyrtohymena. Histrio lemani DRAGESCO is a
synonym of Gastrostyla steinii, Histrio macrostoma GELLERT & TAMAS a synonym of
Sterkiella histriomuscorum. "Histrio muscorum KAHL", Histrio quadrinucleatus SICK,
Histriculus admirabilis FOISSNER, H. similis tricirratus BUITKAMP, and H. trici"atus
DRAGESCO & DRAGESCo-KERNEIS have been, or have to be, transferred to Sterkiella be-
cause caudal cirri are present. Histrio acuminatus MAsKELL (Fig. 222f) and Histriculus
polycirratus AGAMALIEV (Fig. 229d) are considered as species indeterminata.
Parahistriculus differs from Histriculus in the number of dorsal kineties (4 against 6)
and in that the left marginal row terminates on the right margin, that is, the marginal rows
distinctly overlap (Fig. 178b); the proposed synonymy is thus somewhat uncertain.
Pleurotricha also lacks caudal cirri and is thus possibly the sister-group of Histricu-
Ius. However, detailed morphogenetic studies on both taxa are needed to test this hy-
pothesis.
 Histriculus 647

Key to species
1 Marine .................................................... H. similis (p 665)
Linmetic ........................................................... 2
2 In life only about 30-40 Ilm long (Fig. 178b, c) ............. H. minimus (p 658)
In life >80 Ilm long .................................................. 3
3 Posterior end rather broadly truncated, sometimes even slightly indented (Fig. 178i,
j) ..................................................... H. vorax (p 659)
Posterior end rounded or slightly tapered (for example, Fig. 177e, 178<1, g) ..... 4
4 Body margins in middle area almost parallel, in posterior quarter converging and nar-
rowly rounded; transverse cirri very prominent, displaced distinctly anteriad so that
only the 2 posterior-most cirri reach posterior end of cell (Fig. 176a-e, 177a--p) ....
. " ................................................... H. histrio (p 647)
Outline elliptical or with posteriorly converging margins; transverse cirri not very
prominent, at least the 2 posterior-most cirri protrude beyond posterior end of cell
(Fig. 178d-g) ....................................................... 5,
5 Outline elliptical; in life about 80 Ilm long (Fig. 178g) ..... H. complanatus (p 657)
Body margins converging posteriorly, left margin sometimes indented; in life about
110 Ilm long (Fig. 178d-e) ................................. H. sphagni (p 655)

Histriculus histrio (MOLLER, 1773) CORLISS, 1960 (Fig. 22d, 176a-j, 177a-q,
Tables 3, 39, 41)

1773 Paramaecium histrio MOLLER, Vennium Terrestrium et Fluviatilium, p 55 (no figure).

1786 Kerona histrio - MOLLER, Animalcula Infusoria, p 235 (Fig. 177a, b).
1803 Trichoda histrio - SCHRANK, Fauna Boica, p 98.
1830 Sty/onychia his/rio - EHRENBERG, Abh. preuss. Akad. Wiss., year 1830: 43.
1838 Sty/onychia histrio - EHRENBERG, lnfusionsthierchen, p 373 (Fig. 177c).
1852 Cerona histrio M. - PERTY, Kenntniss k1einster Lebensfonnen, piSS.
1859 Sty/onychia his/rio. EHRBG. - STEIN, Organismus der lnfusionsthiere I, p 166 (Fig. 177d).
1861 Sty/onychia histrio (EHR. STEIN) - WRZESNIOWSKl, AnnIs Sci. nat., Zoologie, 16: 336 (Fig. 1771).
1862 Sty/onychia histrio EHREN. - ENGELMANN, Z. wiss. Zool., II: 357 (Fig. 177j).
1865 Sty/onychia histrio - QuENNERSTEDT, Acta Univ. lund., 2: 57 (Fig. 177g).
1878 Histrio steinii STERK!, Z. wiss. Zool., 31: 56 (objective? synonym).
1882 Histrio stein ii, MOLL. sp. - KENT, Manual infusoria II, p 789.
1887 Histrio erethesticus STOKES, Proc. Am. phil. Soc., 24: 252 (Fig. 177h).
1888 Histrio erethisticus, STOKES - STOKES, 1. Trenton nat. Hist. Soc., I: 297 (incorrect subsequent spelling).
1901 Histrio steinii STERK! - Roux, Mem. Inst. natn. genev., 19: 105 (Fig. 177i).
1906 Histrio steinii MOLL. - EDMONDSON, Proc. Davenport Acad. Sci., II: 101 (Fig. 177k).
1932 His/rio (Kerona) histrio (0. F. MOLLER, 1786) STEIN, 1859 - KAHL, Tierwelt Dtl., 25: 615 (incorrect
dating).
1932 Histrio erethisticus STOKES, 1887 - KAHL., Tierwelt Dtl., 25: 615 (incorrect subsequent spelling; Fig.
177e).
1933 His/rio histrio (0. F. MOLLER) STEIN 1859 - WANG & NIE, Contr. bioI. Lab. Sci. Soc. China, 10: 62
(Fig. 177t).
 648 SYSTEMATIC SECTION

f •

•
t • •'

"

Fig. 176a-e Histriculus histrio (a, from BERGER & FOISSNER 1997; b--e, from GsCHWIND 1991; a, d, e, from
life; b, c, protargol impregnation). a) Ventral view, 110 Ilm. b, c) Ventral and dorsal infraciliature,
80-120 Ilm. Arrow marks confluent marginal rows. Note the very prominent, distinctly anteriorly displaced
transverse cirri. d) Right lateral view. e) Dorsal view showing oral apparatus and very prominent transverse
cirri; p 647.

Fig. 176f-j Histriculus histrio (t: g, from BERGER & FOISSNER 1997; b-j, from GsCHWIND 1991; f-j, protargol ~
impregnation). I) Very early divider in ventral view, 115 Ilm. g) Middle divider in ventral view, 120 Ilm. Post-
oral ventral cirrus V/3 is not involved in primordia formation. Parental structures white. b-j) Early, middle,
and late stage of dorsal morphogenesis, which proceeds in Oxytricha pattern, h = 125 Ilm, i = 130 Ilm, j =
140 Ilm. Note that no caudal cirri are formed at the posterior ends of the new dorsal kineties (arrows inj). Ar-
rowheads in (i) mark fragmentation of dorsal kinety 3, arrowhead in (j) denotes new dorsomarginal kineties
(kineties 5 and 6) of the proter. OP = oral primordium, Vl3 = postoral ventral cirrus V/3, 1-6 = dorsal kineties
1-6; p 647.
 Histriculus 649

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650 SYSTEMATIC SECTION

1950 Histrio hyalinus VOROSVAAY, Annis bioI. Univ. szeged., I: 377, 384 (Fig. 1770, p; new synonym).
1960 Histriculus his/rio - CORLISS, 1. Protozool., 7: 275.
1960 Histriculus erethisticus (STOKES, 1887) - CORLISS, 1. Protozool., 7: 275 (incorrect subsequent spelling).
1960 Histriculus hyalinus (VOROSVARY, 1950) - CORLISS, 1. Protozool., 7: 275.
1968 Oxytricha (Histriculus) histrio (STEIN, 1859) CORLISS, 1960 - CHORlK, Free-living ciliates, p 137 (Fig.
177m).
1976 Oxytrichafallax STEIN, 1859 - SCHMALL, Diplomarbeit, p 38 (Fig. 177q; misidentification).
1991 Histriculus histrio (MOlLER, 1773) CoRLISS, 1960 - GsCHWIND, Dipiomarbeit, p 52 (Fig. 176b-e, b-j;
collection, determination, and live observations were done by W. FOlSSNER).
1997 Histriculus histrio - BERGER & FOISSNER, Arch. Protistenk., 148: 136 (Fig. 22<1, 1763, 1; g).

Nom e n c I at u rea n d t a x 0 nom y: CALKINS (1926, p 392) erroneously desig-

nated the present species as "Histrio pel/ionella". KAHL (1932) classified Histrio as sub-
genus of Oxytricha; the correct names in his revision are thus Oxytricha (Histrio) histrio
and O. (Histrio) erethisticus. Incorrect subsequent spellings: Histrio erecthisticus STOKES
(MOcKE 1979, P 273); Histrio erethysticus STOKES (EDMONDSON 1920, P 186); Histrio
steini STERK! (ANDRE 1912, p 138); 0. (Histrio) erethisicus STOKES (MARGALEF LOPEZ
1945, p 377); s. histrie (WRZESNlOWSKI 1861, p 334); Stylonichia histrio (MAUPAS 1883,
p 619); Stylonichia histrix (SERNOV 1929, P 557); Stylonichia hystrio (ENRIQUES 1913,
p 105); Stylonychnia histria (BALDENSPERGER 1927, P 200); Stylonychnia histrio (BAL-
DENSPERGER 1927, P 229).
The identity of Histriculus histrio and H erethesticus was already supposed by HEM-
BERGER (1982, P 237) and ForSSNER in GSCHWIND (1991) although STOKES (1887a) illus-
trated and mentioned 9 frontoventral cirri (Fig. 177h), similar to in H sphagni. However,
all other characters of Histriculus histrio and H erethesticus agree almost perfectly so
that synonymy is very likely. The arrangement of the postoral and pretransverse ventral
cirri in Histriculus hyalinus is rather probably misobserved (Fig. 1770, p), and "Oxytri-
cha platistoma" sensu HORvArn (1933) is presumably also H histrio (Fig. 177m', n). I
have never observed specimens with 9 frontoventral cirri. The redescriptions by BORGER
(1905; Fig. 232f), CONN (1905; Fig. 232b), and FROMENTEL (1876) are insufficient.
Histriculus histrio should not be confused with Sterkiella or Stylonychia species; it is
easily identified by its trembling movement, the tapered posterior end, and the prominent
transverse cirri which do not protrude beyond posterior end of cell.

M 0 r ph 0 log y and b i 0 log y: In life 90-140 x 40-70 Ilm (GSCHWIND 1991),

86-115 Ilm long (EHRENBERG 1838), larger specimens 130-150 Ilm long (STEIN 1859b),
175-215 x 70-80 Ilm (WANG & NlE 1933), 150-170 x 70-80 Ilm (original measure-
ments), 100 x 40 Ilm (VOROSVAAY 1950). Almost parallel-sided or slightly elliptical,
somewhat tapered posteriorly, anteriorly moderately broadly rounded. Right margin
slightly convex, left straight or slightly concave. Flattened about 2: 1 dorso-ventrally, an-
terior and posterior part rather thin and thus very translucent. Two folds parallel to poste-
rior margin, recognisable in life, possibly due to outer dorsal kineties (Fig. 176a). Macro-
nuclear nodules in median of cell, ellipsoid, in life about 30 x 15 Ilm, fmely granulated.
Each macronuclear nodule invariably (n = 20; GsCHWIND 1991) with an ellipsoid, in life
 Histriculus 651

Fig. 177a-m Histriculus histrio (a, b, after MOLLER 1786; c, after EHRENBERG 1838; d, after STEIN 1859b; e,
from KAHL 1932; t: from WANG & NlE 1933; g, after QUENNERSTEDT 1865; h, from STOKES 1887a; i, after Roux
190 I; j, from ENGELMANN 1862; k, from EDMONDSON 1906; 1, after WRZESNIOWSKI 1861; m, from CHORIK 1968).
Ventral views in life, a-d, g, h = size not indicated, e = 150 11m, f = 200 11m, i, j = 170 11m, k = 160 11m, 1=
210 11m, m = 155 11m. The specimen in (I) is defecating; p 647.
652 SYSTEMATIC SECTION

o p

Fig. 177m'-p Histriculus histrio (m', n, from HORvArn 1933; 0, p, from VOROsvARy 1950. m', sublimate-
toluidinblue stain after HORVArn; n-p, opal-blue stain). m', 0) Ventral and lateral view, m' = 110 J.1m, n =
115 11m. 0, p) Ventral and dorsal infraciliature, 155 11m. The arrangement of some cirri is unreliable and very
likely dorsal kinety 6 is too long, possibly due to inappropriate stain method; p 647.

about 5-6 x 4-5 Ilm sized micronucleus. According to HAsHIMOTO (1953; identification
not documented by illustrations) ectoplasm with so-called refractile bodies, 4-6 !lm in di-
ameter, which have an affinity to lead. Contractile vacuole with inconspicuous canals.
Cytoplasm usually with many greasily shining, 3-5 !lm sized globules, except in very thin
anterior and posterior part of cell. Food vacuoles 5-15 Ilm or more in diameter. Move-
ment very characteristic: swims continuously forward, moves back like a flash in a wide
curve, then repeatedly swims back and forth very rapidly, the distances becoming shorter
and shorter, that is, appears to be trembling.
Adoral zone of membranelles about 50 % of body length. Buccal area flat and mod-
erately broad. Undulating membranes only moderately curved and almost not intersecting
(that is, nearly parallel), endoral shorter than paroral (Fig. 176b). Invariably 18 frontal,
ventral, and transverse cirri (n = 20; GSCHWIND 1991), arranged as shown in Figure 176b.
Frontal cirri and buccal cirrus distinctly enlarged. Buccal cirrus at anterior end of paroral.
Transverse cirri in life 25-30 Ilm long, very prominent and thus easily recognisable even
at low magnification, displaced distinctly anteriad so that only cirri IVII and VII reach
posterior end (rarely protrude slightly), sometimes fringed. Transverse cirrus VIII some-
 Histriculus 653

what separate from cirrus VII, usually slightly

larger than other transverse cirri (WANG & NIE
1933, own observations). Usually all transverse
cirri point toward posterior tip of cell. Marginal
cirri in life about 12-20 Ilm long, anteriorly
thicker than posteriorly. Marginal rows conflu-
ent posteriorly in median of cell. Right row be. .
gins at level of distal end of adoral zone of
membranelles. Dorsal cilia in life about 3-4 Ilm
long. Invariably (n = 20; GSCHWIND 1991) 6
dorsal kineties (including 2 dorsomarginal rows
= kineties 5, 6). Kineties 1-5 about of body
-
,
0:-
length, kinety 6 terminates in about mid-body.
Distance between kineties 3 and 4 distinctly
wider than between other kineties (Fig. 176c).
VOROSVA.R.y (1950) illustrated 6 kineties of body
length, which is probably a misobservation due
to inappropriate stain method (Fig. 177p).
The formation of the oral primordium com-
mences near the left transverse cirrus (Fig.
176f). The posterior postoral ventral cirrus V/3
is not involved in primordia formation, as in all Fig. 177q Histriculus histrio (from SOIMALL
other Stylonychinae (Fig. 176g). Dorsal mor- 1976). Ventral infraciliature after protargol
phogenesis is in Oxytricha pattern (Fig. impregnation, 160 JUll; p 647.
176h-j). As already stated by STEIN (1859b), no
caudal cirri are formed. There are some older
papers concerning adoral zone of membranelles (MAmR 1903), morphogenesis (HAsm-
MOTO 1957, NUSSBAUM 1886, STERKI 1878), conjugation (BALBIANI 1882, P 17; ENGEL-
MANN 1862, 1876, STEIN 1859b), refractile bodies at the cystic stage, and encystment
(HAsmMOTO 1956, 1960).

o c cur r e n c e and e colo g y: Common in eutrophic lentic freshwater such as

ponds, lakes, and in slow running waters rich in macrophytes and filamentous algae; usu-
ally absent in alpine, fast flowing brooks and rivers.
Locus classicus is a freshwater habitat in Denmark, where MOLLER (1773, 1786) dis-
covered Histriculus histrio between filamentous algae. STOKES (1887a) discovered the
synonym H. erethesticus in shallow pools with Lemna and algae, probably near Trenton,
New Jersey, USA, where he lived. Locus classicus of the synonym H. hyalinus is a brook
in Hungary, where VOROSVA.R.y (1950) discovered it on the water surface of lentic areas
during summer.
Records documented by illustrations: in old samples with living filamentous algae in
Copenhagen, Denmark, and in Berlin and Landshut, Germany (EHRENBERG 1838); on tufts
654 SYSTEMATIC SECTION

Table 39 Morphometric data of Histriculus histrio (from GsCHWIND 1991). All data are based on protargol-
impregnated specimens. All measurements in micrometres. CV = coefficient of variation (in %), Max = maxi-
mum value, mean = arithmetic mean, Min = minimum value, n = sample size, SD = standard deviation

Character mean SD CV Min Max n

Body, length 102.2 10.3 10.0 81 122 21
Body, width 49.2 5.0 10.1 40 60 21
Macronuc1ear nodule, length 22.5 2.4 10.7 17 27 20
Macronuclear nodule, width 8.4 1.2 14.4 6 10 20
Adoral membranelles, number 47.1 2.0 4.2 41 50 18
Right marginal row, number of cirri 42.2 1.9 4.6 39 46 20
Left marginal row, number of cirri 29.0 1.7 5.9 26 33 20

of Beggiatoa and in the upper layer of small basins filled with water from a eutrophic
pond (Poppelsdorfer Weiher) in Bonn, Germany (SCHMALL 1976); running and stagnant
waters near Leipzig, Germany (ENGELMANN 1862); common among Lemna and algae in
Germany (KAHL 1932); in clear running waters among Callitrichacea, Lemna, and fila-
mentous algae (often very abundant in several day old samples from such sites) in
Prague, Czechoslovakia, and near Tharand, Germany, abundant in a trout pond supplied
with clear spring water (STEIN 1859b; see also SRAMEK-HuSEK 1953); moderately frequent
in the periphyton ofa lake (Teufelsee) near Berlin, Germany, in August (GSCHWIND 1991;
collector: W. ForSSNER); pool containing sodium hydroxide in Hungary, cultivated in
straw infusion (HORvATH 1933, see taxonomy); Poland (WRZESNIOWSKI 1861, WRZESN-
IOWSKIEGO 1866); Sweden (QUENNERSTEDT 1865, 1869); various clear waters in Switzer-
land (Roux 1901, STERK! 1878); frequent in the water of an aquarium, collected from
various ponds in Nanking, China (WANG & NIE 1933). I found it in small, eutrophic
ponds with many macrophytes in the city of Salzburg, Austria.
Records not documented by illustrations: Austria (ForSSNER & ForSSNER 1988); in
September in a pond near a lake (Grabensee) in Austria (GSCHWIND 1991; collector: W.
ForSSNER); rare in running waters in Upper Austria (BLATTERER 1994); surface layer
(Kahmhaut) of an infusion from the hypertrophic pond of the Salzburg University, Aus-
tria (BLATTERER 1989); in mud and on putrefying leaves in a spring near Vienna, Austria,
in April (KUHN 1940); Belgium (BERvOETS 1940, CHARDEZ 1987); Bulgarian running wa-
ters (DETCHEVA 1981, 1986, RUSSEV et al. 1976, 1984; see also Table 41); saprobic brook
near Prague, Czechoslovakia (BUCIfAR 1957); at 29-32 °C and pH 5.7-7.2 in a brook in
Czechoslovakia (MATIS & STRAKOVA-STRIESKOvA 1991); ponds in England (CRAIGIE 1921,
DIXON 1920); meadow pond in Estonia (JACOBSON 1928); mouth of the Schussen River,
Germany (WETZEL 1928a); drainage canal with a saprobic index of 2.4 in Germany
(HEuss 1975); among detritus and in the surface film of ditches regularly contaminated
with pesticide sprays and in unsprayed ditches in the Elbe flood plain, Germany (CASPERS
& HECKMAN 1982); in beta- to alphamesosaprobic brooks and in Amper River in Ger-
many (FOISSNER et al. 1992a, GSCHWIND 1991, HEuss 1976); with low abundance in basins
with largely mineralised water and in clear pond water near Bonn, Germany (GREISER
 Histriculus 655

1974, SCHMIDT 1913, 1916); mesotrophic lake and ponds in Germany (LEVANDER 1892,
MOcKE 1979); caves and mines in Germany (GRIEPENBURG 1933, 1934, 1935, WETZEL
1929); aquariums in Germany (GONKEL 1989a, b, 1990, 1991); ponds (oxbows) of the
Rhine River, Germany (BAIER 1934); periphyton of unpolluted foothill stream (Breiten-
bach) near the village of Schlitz, Germany (PACKROFF & ZWICK 1996); Tisza River, proba-
bly in still areas of flooded parts, Hungary (STILLER 1942; further record from Hungary:
DUDICH 1947); Italy (CuNEo 1891); betamesosaprobic area of a lowland river in Poland
(WIACKOWSKI 1981); Russia (EICHWALD 1844, 1847, WEISSE 1845, 1848a, c); Danube
River, Slovakia (SZENTIVANY & TIRJAKovA 1994); waters rich in organic matter, Spain
(MARGALEF LoPEZ 1945); in September abundant in the botanical garden of Basle, Swit-
zerland (RIGGENBACH 1922); old infusion, Switzerland (PERTY 1852b; further records from
Switzerland: ANDRE 1912, MERMOD 1914, THIEBAUD & FAVRE 1906); pond in Kiev and
reservoirs of the Seversky Donets Basin, Ukraine (KovALCHUK & SVINTSOVA 1994, KRAv-
CHENKO 1969); at 4-19 °C among algae (Ulothrix, Tetraspora) in lentic water of the litto-
ral of Lake Baikal (GAJEVSKAJA 1927, GAJEWSKAJA 1933, ROSSOLIMO 1923); Iowa, USA
(SHAWHAN et al. 1947); among Ruppia in Devil's Lake complex, North Dakota, USA (ED-
MONDSON 1920); in April, June, July, and November in small numbers in a fresh water
pond in Pennsylvania, USA (WANG 1928); Argentina (SECKT 1924); swamps in Australia
(WHITELEGGE 1889). Records from terrestrial habitats (MARTIN & SHARP 1983) are cer-
tainly misidentifications.
Feeds on bacteria, diatoms (Synedra acus), Chlorella-like green algae, and flagel-
lates (ANDRE 1912, GSCHWIND 1991, V6R6svARy 1950, WANG & NIE 1933, own observa-
tions). Biomass of 106 individuals about 120 mg (NESTERENKO & KOVALCHUK 1991).
Some autecological data are shown in Table 41. MATIS (1975) found H. histrio occasion-
ally in temporary pools and brooks near Bratislava, Czechoslovakia, under the following
conditions: 7-23.5°C, pH 6.0-6.9, 1.5-2.7 mg 1. 1 O2• GROLrERE et al. (1990) counted
13 000-15 000 ind.I-1 in beta- to alphamesosaprobic and polysaprobic areas ofa running
water at following conditions: 10-90 mg I-I COD, 0-4.6 mg I-I NH/-N, 0.1-0.26 mg I-I
POl--P. According to WETZEL (1928b), hydrogen sulphide is not tolerated.
Histriculus histrio is not classified in recent lists of indicator species of water quality
(ForSSNER 1988a, ForssNER et al. 1991, SLAoECEK et al. 1981, WEGL 1983). Previously, H.
histrio was classified as oligosaprobic (KOLKWITZ & MARSSON 1909, KOLKWITZ 1950,
MAUCH 1976, MEz 1898). According to the faunistic and ecological data H. histrio can be
found in oligosaprobic to beta- to alphamesosaprobic waters; I thus suggest the following
classification: b-o; 0 = 4, b = 4, a = 2, 1= 2, SI = 1.8.

Histriculus sphagni (STOKES, 1891) CORLISS, 1960 (Fig. 178d-f)

1891 Histrio sphagni STOKES, JI K microsc. Soc., year 1891: 702 (Fig. 178d, f).
1932 Histrio sphagni STOKES, 1891- KAHL, Tierwelt Dtl., 25: 617.
1960 Histriculus sphagni (STOKES, 1891) - CORLISS, J. Protozool., 7: 275.
1975 Histriculus sphagni (STOKES, 1891) CORLISS, 1961- GROLIERE, Protistologica, II: 492 (Fig. 178e).
 656 SYSTEMATIC SECTION

~/ ..
;j~1 •. .

Fig. 178a, g Histricu/us comp/anatus (a, from VUXANOVICI 1961; g, after STOKES 1887b). Ventral views in life,
a = 110 Ilm, g = 80 Ilm; p 657. b, c Histricu/us minimus (from GROLlERE 1975. b, c, protargol impregnation).
Ventral and dorsal infraciliature, 35 Ilm; p 658. d-f Histricu/us sphagni (d, t; after STOKES 1891; e, from GRO-
LIERE 1975. d, t; from life; e, protargol impregnation). Ventral views and left margin, d, e = 110 Ilm. Arrow-
heads mark an additional cirrus behind the middle frontal cirrus; p 655. b-j Histricu/us vorax (h, j, from
CURDS 1966a; i, after STOKES 1891. h-j, from life). b) A small cell in ventral view, 115 Ilm. i, j) Ventral views
of typical specimens, i = 180 11m, j = 205 11m. eb = endogenous bud, Ma = macronuclear nodule; p 659.
 Histriculus 657

Nom e n c I a t u rea n d t a x 0 nom y: KARL (1932) classified Histrio as subgenus

of Oxytricha; the correct name in his revision is thus Oxytricha (Histrio) sphagni. This is
a senior secondary homonym of O. (Opisthotricha) sphagni in KARL (1932). The GRO-
LIERE (1975; Fig, 178e) population agrees rather well with the original description (Fig.
178d), especially in that both STOKES and GROLffiRE found an additional cirrus in the fron-
tal area, which should be checked once more. The undulating membranes are almost in
Stylonychia pattern; however, GROLIERE (1975) did not observe caudal cirri; indicating
that the classification in Histriculus is appropriate. Identity of H. sphagni and H. compla-
natus cannot be excluded, whereas BORROR (1972a) synonymised it with the type species,
H. histrio. Histriculus sphagni sensu GROLIERE (1969; Figure 10 on Planche III) is proba-
bly a different species because the transverse cirri are separated into two groups. Rede-
scription necessary.

M 0 r p h ci log y and b i 0 log y: In life about 110 J.lm long (11225 inch; STOKES
1891 wrote, obviously erroneously, 111225 inch). According to GROLffiRE (1975) in life
(1) 85-115 x 40-50 J.lm. Macronuclear nodules after protargol impregnation 10-12 times
5-6 J.lm. Two micronuclei. Adoral zone ofmembranelles about 45-50 % of body length,
35-42 membranelles. Undulating membranes rather straight, that is, similar to in Stylony-
chia (see taxonomy). Frontal and transverse cirri distinctly enlarged. One additional cir-
rus behind frontal cirri (Fig. 178d, e, arrowhead). 50-60 left and right marginal cirri, ac-
cording to STOKES (1891) slightly elongated posteriorly, after GROLIERE (1975) all of same
length. Five dorsal kineties (GROLIERE 1975). ALONSO (1975) and ALoNSO & PEREZ SILVA
(1966a) described the polytene chromosomes in the macronuclear anlage; however, their
identifications are not documented by illustrations.

o c cur r e n c e and e col 0 g y: Rare freshwater species. Locus classicus is proba-

bly Trenton in New Jersey, USA, where STOKES (1891) discovered Histriculus sphagni in
standing water with decaying Sphagnum, collected from shallow pools and ponds. GRO-
LIERE (1975, 1977, 1978) and GROLIERE & NJINE (1973) found it in ponds and peat-bog
moss in France. Records not substantiated by illustrations: 16 indo cm-2 in the Parma
River, Italy (MAnoNI & GHETTI 1977, as Oxytricha sphagni STOKES); brook and pond in
Bulgaria (DETCHEVA 1976a); lakes in Canada (PUYTORAC et al. 1972); Amazon River,
Peru (CAIRNS 1966a). Feeds on small algae such as Chlamydomonas sp. (GROLIERE 1975).

Histriculus complanatus (STOKES, 1887) CORLISS, 1960 (Fig. 178a, g)

1887 Histrio complanatus STOKES, Ann. Mag. nat Hist., 20: 113 (Fig. 178g).
1888 Histrio complanatus, STOKES - STOKES, J. Trenton nat. Hist. Soc., I: 296.
1932 Histrio complanatus STOKES, 1887 - KAHL, Tierwelt Dtl., 25: 615.
1960 Histriculus complanatus (STOKES, 1887) - CORLISS, 1. Protowol., 7: 275.
1961 Histrio complanatus STOKES, 1887 - VUXANOYICI, Studii Cere. BioI., 13: 437 (Fig. 178a).
658 SYSTEMATIC SECTION

Nom e n c I at u rea n d t a x 0 nom y: KAHL (1932) classified Histrio as subgenus

of Oxytricha; the correct name in his revision is thus Oxytricha (Histrio) comp/anatus.
Incorrect subsequent spelling: Histrio companatus STOKES, 1887 (BORROR 1972a, P 14).
This species shows no peculiarities so that defmite identification is rather difficult.
Identity with H. sphagni or Sterkiella histriomuscorum cannot be excluded. The rede-
scriptions by CHORIK (1968; Fig. 232g), CONN (1905; Fig. 232a), and LEPSI (1965; Fig.
222g, 224f) are insufficient. Detailed redescription necessary.

M 0 r p hoi 0 g y and b i 0 log y: In life about 80 x 40 11m, according to VUXANO-

VICI(1961) 80-120 11m long. Twice as long as broad. Outline elliptical, anterior end
evenly rounded, posterior inconspicuously pointed. Typical18 frontal, ventral, transverse
cirral pattern. Three rightmost transverse cirri protruding beyond posterior end of cell.
Marginal cirri "uninterrupted, longest and largest posteriorly" (STOKES 1887b), indicating
that inconspicuous caudal cirri are present.

o c cur r e n c e and e colo g y: Rare freshwater species. Locus classicus is Tren-

ton, New Jersey, USA, where STOKES (1887b) discovered Histricu/us comp/ana/us in a
shallow pool in early spring. VUXANOVICI (1961) found some specimens in a culture with
marsh-plants in a lake in Bucharest, Bulgaria, in December.
Records not documented by illustrations: pelagial of a clear lake in Styria, Austria
(KRAmER 1988); sandy soil from the flood plains of the Tisza River in Szeged, Hungary,
an area inundated at most once a year (HORvATH 1950); 1000 indo I-I in activated sludge
ofa Polish plant with a Sheffield aeration tank (KLIMOWICZ 1973); low salinity lagoons of
the Caspian Sea (AGAMALIEV 1973, 1986); at 22 °C in a metal tank with much blue-green
algae and sulphur bacteria on a hillside farm in the Mountain Lake region, Virginia, USA
(BOVEE 1960).
Feeds on bacteria and algae (HORVATH 1950). DAWSON (1926, 1928) and RICHARDS &
DAWSON (1927) found an average daily division rate of 1.24 (maximum value about 1.8)
in a culture medium of dried timothy hay and wheat; usually a grain of wheat was added.
See also ZAIKA (1970) for rate of reproduction. DETCHEVA (1979c) found H. comp/anatus
once in the Strouma River, Bulgaria, under the following conditions: 17 °C, pH 7.7,
9 mg 1-1 O2 (79 % saturation), 9.2 mg I-I BOD5, 88 mg 1-1 Ca+, 25 mg 1-1 Mg+, 72 mg 1-1
SO/-, 23 mg}"1 Cl-, 0.04 mg I-I NH/-N, 1.7 mg I-I N03--N, 0.17 mg I-I N02--N,
0.26 mg 1"1 Fe2+.

Histriculus minimus (GROLrERE, 1975) BERGER & FOISSNER, 1997

(Fig. 178b, c)

1975 Parahistriculus minimus GROLIERE, Protistologica, 11: 493.

1997 Parahistriculus minimus GROLIERE, 1975 - BERGER & FOISSNER, Arch. Protistenk., 148: 149 (see nomen-
clature).
 Histriculus 659

Nom e n c I at u rea n d t a x 0 nom y: BERGER & ForSSNER (1997) synonymised

Parahistriculus with Histriculus; they are thus considered as combining authors of the
only included species. Incorrect subsequent spelling: Parahistriculus minimum (GROLIERE
1977, P 347). Histriculus minimus is very similar to Oxytricha setigera and O. balladyna
in terms of the small size and the single micronucleus (Fig. 178b). However, both Oxytri-
cha species have caudal cirri and 10-15 /lm long dorsal cilia. Redescription, especially
live observation necessary.

M 0 r ph 0 log y: Very small, after protargol impregnation 30-40 x 15-20 /lm (mean =
36 x 17 /lm). Oval. Not contractile. Often 1 micronucleus between macronuclear nodules
(Fig. 178b); however, sometimes 1-6 micronuclei per nodule (Fig. 178c). This extreme
variability is very uncommon and indicates that possibly two species were mixed up.
Adoral zone ofmembranelles about 50 % of body length, 22-30 membranelles. 18 fron-
tal, ventral, and transverse cirri. Frontal and transverse cirri slightly to distinctly enlarged,
latter protrude beyond posterior end of cell. Buccal cirrus near anterior end of rather
straight undulating membranes. Pretransverse ventral cirri displaced distinctly anteriad.
18-21 right marginal cirri; left marginal row with 20-26 cirri, terminates on right side;
marginal rows thus overlapping. Four dorsal kineties of about body length. Dorsal cilia
2-3 /lm long.

o c cur r e n c e: Locus classicus is a peat-pit at Landie, France (GROLIERE 1975, 1977).

Not found since.

Histriculus vorax (STOKES, 1891) CORLISS, 1960 (Fig. 178h-s, 179a-n, 180)

1891 Histrio vorax STOKES, JI R. microsc. Soc., year 1891: 703 (Fig. 178i).
1932 Histrio vorax STOKES, 1891 -KAHL, Tierwelt Otl., 25: 617.
1960 Histriculus vorax (STOKES, 1891) - CORLISS, J. Protozool., 7: 275.
1966 Histriculus vorax (STOKES) CORLISS - CURDS, J. Protozool., l3: 155 (Fig. 178h,j-.'l, 179a--n).
1969 Histriculus vorax CORLISS, 1960 - CURDS, Wat. Pollut. Res., No. 12: 68 (incorrect author).
1991 Histriculus vorax (STOKES, 1891) CoRLISS, 1960 - FOISSNER, BLATTERER, BERGER & KOHMANN, Informa-
tionsberichte des Bayer. Landesamtes filr Wasserwirtschaft, 1/91: 309.

Nom e n c I at u rea n d t a x 0 nom y: KAHL (1932) classified Histrio as subgenus

of Oxytricha; the correct name in his revision is thus Oxytricha (Histrio) vorax, which is
a secondary homonym of 0. (Stylonychia) vorax in KAHL (1932, P 619).
The CURDS (1966a) population is described after wet silver impregnation. A detailed
redescription of the live aspect and after protargol impregnation is thus necessary. CURDS
(l965b, c, 1966a) described endogenous bud formation (see below), a very unusual
(doubtful?) form of reproduction in hypotrichs. The illustration in KUTIKOWA (1984) is a
redrawing from CURDS (1966a).
 660 SYSTEMATIC SECTION

,.:-,-
rp

••
•
•
. ,-... •.-.... ..
••
\

"
· k q

eb

Fig. 178k-s Histriculus vorax (from CURDS 1966a). k, I) Ventral and dorsal infraciliature after wet silver ni-
trate impregnation, sizes not indicated. m) Lateral view of a large specimen containing a mature endogenous
bud. n-s) Reorganisation and division of the nucleus apparatus during endogenous bud formation. Replication
bands form at extreme outer ends of the 2 macronuclear nodules, move centrally, and disappear. Micronuclei
in anaphase (n-p). One macronuclear nodule bends toward budding area (q, arrow). Constriction ofmacronuc-
lear nodules continues, the nodules divide and migrate to budding area (r, arrows). Large specimen carrying
bud containing 2 macronuclear nodules and 4 micronuclei (s); p 659. t Histriculus similis (after QUENNER-
STEDT 1867). Ventral view in life, 135 /lm; p 665. u-w Histriculus simi/is (insufficient redescription; from
CHARDEZ 1986). u) Ventral infraciliature after nickel sulphate fixation, size not indicated. v, w) Lateral view of
a specimen without and with endogenous bud. eb = endogenous bud, rp = replication band; p 667.
 Histriculus 661

&l
lj,. I-
(0\
'J
I TOTAL NUt.4BER.AND STANDARD DEVIATION. OF
CILIATES PER MILLILITRE
~
0:
!zUJ Fig. 179a-f Histrieulus vorax
o :i 250 + ~ t:l (from CURDS 1966a). a) Growth
ffi 3 200
NUt.4BERS OF LARGE ADULTS AS PERCENTAGES OF
TOTAL CILIATE POPULATION 50 lI.. 0: showing occurrence of large
~- O~
adult cells in a clonal culture. b)
~ ~ 150 UJ ~
Size distribution in the same
ZUJ ~~
;i ~ 100 25 ~ tl clonal culture at various ages.
OUJ50
~~
~!:i
UJ~
c-f) Cyst development of His-
:i L-..J.....Ji:::~-""'.J,.....I---'-..J.....J---L--'-"''-'--'-L....J.--L-.l-L-' 0 ~ Q trieu/us vorax. c) Newly encysted
iJ 0 2 <4 6 8 10 12 14 16 18 20
a bud; period of rest d) Develop-
TIME (dayS) ing cyst 3--4 h after end of resting
period. e) Developing cyst 6 h af-
ter end of resting period. Rotation
(arrow) begins and contractile
vacuole becomes apparent f) De-
veloping cyst 12 h after end of
rest period and 12 h before
excystment Cirri noted for flfSt
time (arrow). CV = contractile
vacuole; p 659.
b
100 150 200 250
LENGTH (microns)

M 0 r p hoi 0 g y and b i 0 log y: After STOKES (1891) in life about 170 Jlm long.
According to CURDS (1966a) size and shape rather variable (Fig. 178b, j, 179a, b). Small
adult cells 70-140 x 50-60 Jlm, large adult cells 190-250 x 100-120 Jlm. KAHL (1932),
obviously erroneously, stated 80 Jlm in the legend to the figure, which is a redrawing
from STOKES (1891). Large cells broader and blunter than small cells (Fig. 179g, h).
Broadly elliptical, often almost rectangular, then left margin concave and right one con-
vex (Fig. 178i, j, 179h). Posterior end in large specimens often indented. Flattened about
2:1 dorso-ventrally. Large cells usually with dark cytoplasm rich in small, dark granules
(no cortical granules; Fig. 179h). Each macronuclear nodule usually with 2 micronuclei.
Contractile vacuole in about mid-body or slightly in front.
Adoral zone of membranelIes about 50 % of body length. Frontal cirri, buccal cirrus,
and 1 or 2 left frontoventral cirri distinctly enlarged. Transverse cirri enlarged, displaced
somewhat anteriad so that only the 3 rightmost cirri protrude beyond posterior end of
cell. Comparatively few marginal cirri, widely spaced, longer and more prominently pro-
jecting beyond posterior end of cell (possibly redescription will show that caudal cirri are
 662 SYSTEMA TIC SECTION

Fig. 179g-1 Histricu/us vorax (from CURDS 1966a). g) Small adult. b) Large adult The outline is very similar
to that in Figure 178i, indicating that the identification is correct. i) Large adult containing developing endoge-
nous buds. j) Large adult containing a large, single endogenous bud just prior to deposition. k) Non-trophic
embryo approximately 30 min after excystment. I) Trophic embryo approximately 4 h after excystment e.b. =
endogenous bud; p 659.

present, demanding that it be transferred to Sterkiella). Six dorsal kineties, kineties 5 and
6 (= dorsomarginal rows) only 112 and 113 of body length (Fig. 1781).
CURDS (1965b, c, 1966a) described endogenous bud fonnation (Fig. 178m-s, 179c-f,
i-n). PANG & ZHANG (1981) assumed that CURDS observed cannibalism induced by over-
feeding. Conversely, it cannot be excluded that this type of reproduction, which is, for ex-
ample, common in some suctorians (MATIHES 1988), also evolved in hypotrichs. Another
example of parallel evolution of a curious reproductive strategy is the division in cysts, a
 Histriculus 663

Fig. 179m, n Histricu/us vorax (from CURDS 1966a m, n, basic fuchsin stain). m) Division of the macronu-
cleus during bud fonnation. Reorganised macronucleus nodule nearest oral region constricts first and bends to-
ward budding area. n) Large adult bearing 2 buds, each with its own macronuclear nodules; p 659.

common strategy, for example, in some colpodids (FOISSNER 1993), but only known from
Keronopsis and Paraholosticha (DIECKMANN 1989) within the hypotrichs. I have thus pro-
vided some of CURDS' figures and micrographs (see legends for some details). CHARDEZ
(1986) reported a similar phenomenon in "Histriculus similis" (Fig. 178u-w).

o c cur r e n c e and e colo g y: Rare; probably confined to freshwater. Locus clas-

sicus is a pond with decaying vegetation in the USA, probably near Trenton in New Jer-
sey, USA, where STOKES lived. CURDS (1965b, c, 1966a) found Histriculus vorax in an
activated-sludge sample collected from the Northern Outfall Sewage Works of the Lon-
don County Council, England. CURDS et al. (1968) reported on 2 x 106 to 3 x 106 indo 1"1
activated sludge in a small scale plant. CURDS & COCKBURN (1970a) found it with a fre-
quency of 21 % in the eftluent of 52 percolating filters. Not recorded from Imhoff-tanks
(CURDS 1975). Records not documented by illustrations: pond in California, USA (STOUT
1980); activated sludge (BANINA 1983, REID 1969).
CURDS (l965a) and CURDS & VANDYKE (1966) studied the feeding habits and growth
rates. They tested 19 strains (15 species) of bacteria and found that Klebsiella aerogenes,
Azotobacter chroococcum, Bacillus cereus, B. subtilis, B. subtilis var. niger, Escherichia
664 SYSTEMATIC SECTION

9
~---------------------.. ([]) -.J
m Z
1/1

1 c(
0:
::>
0
~
W \I) 0 0
z
..Jz ~
u; 0
ID 0 U
c( z
f 0: ....
::>0
::>
Oz
~O
U.U ~
"::-::;: h

f 1
c d @ .........

GROWTH
1
fJA 15k j :':".=.::.:

a~ b~
l,---~ ~
GROWTH
~~
I k
Fig. 180 Complete asexual life cycle of Histriculus vorax (from CURDS 1966a). a) Small adult. b) Division of
small adult. c) Large adult. d) Division of large adult. e) Development of endogenous bud within large adult. f)
Large adult after deposition of bud. g) Newly deposited bud. b) Encysted bud. i) Developing cyst. j) Newly
hatched embryo. k) Embryo becomes trophic. I) Trophic embryo grows to become small adult; p 659.

coli, Proteus vulgaris, Pseudomonas ovalis, P. putrijaciens, P. jluorescens, Streptococ-

cus faecalis, and Vibrio cuneatus are favourable. Enterobacter cloacae is unfavourable
and Chromobacterium violaceum killed the ciliates within 3 h. Number of divisions per
 Histriculus 665

day varied between 2 and 6 when feeding on Pseudomonas ovalis and cultured at 25°C.
CURDS (1966a) cultured a clone in an infusion of 0.l5 % dried lettuce in tap water with
Bacillus cerus as the sole food organism. Upper temperature limit 40°C (NISBET 1984).
Biomass of 106 specimens about 350 mg (FOISSNER et al. 1991).
Histriculus vorax is classified as strict alphamesosaprobic indicator of water quality:
a; a = 10, I = 5, SI = 3.0 (SLAoECEK et al. 1981, WEGL 1983, FOISSNER 1988a, FOissNER et
al. 1991). This classification has to be verified by detailed autecological and saprobio-
logical investigations on reliably determined populations.

Marine species

Histriculus similis (QUENNERSTEDT, 1867) CORLISS, 1960 (Fig. 178t)

1867 Sty/onychia simi/is QuENNERsTEDT, Acta Univ. lund., 4: 38 (Fig. 178t).

1932 Histrio simi/is QUENNERSTEDT, 1867 - KAHt., Tierwelt Otl., 25: 615, in part Fig. 121,6; not Fig. 116,2.
1960 Histricu/us similis (QUENNERSTEDT, 1867) - CoRLISS, J. Protozool., 7: 275.

Nom e n c I at u rea n d t a x 0 nom y: KAHL (1932) classified Histrio as subgenus

of Oxytricha; the correct name in his revision is thus o. (Histrio) similis, which is a sec-
ondary homonym of o. (Opisthotricha) similis in KAHL (1932, P 611). I agree with FOISS-
NER (1988a) that KAHL's (1932) identification with a marine, quadrinucleate form (Fig.
186b) cannot be accepted; the fact that SICK (1933) established a new species for KAHL's
taxon, namely Histrio quadrinucleatus, was always overlooked.
According to WENZEL (1953), Sterkiella histriomuscorum is the junior synonym of H.
similis. However, this has to be proved on a detailed redescription of a marine
population. At this stage, I propose classifying H. similis (marine, 2 macronuclear nod-
ules!) and Sterkiella histriomuscorum as separate species. Since it cannot be excluded
that H. similis in fact lacks caudal cirri, I do not transfer it to Sterkiella. There is some
faunistic and ecological literature available on "H. simi/is". When it concerns freshwater
or terrestrial populations it is listed in Sterkiella histriomuscorum. When it concerns ma-
rine populations and if the determination was done expressly according to KAHL (1932) it
is listed in Sterkiella quadrinucleatus (illustrations from BORROR 1972b, MARGALEF 1948,
REuTER 1961). The populations in NIETO et al. (1984; Fig. 2321) and CHARDEZ (1986; Fig.
178u-w) are insufficiently redescnoed.

M 0 r p hoi 0 g y: In life 110-130 Ilm long, about 3 times as long as broad. Long ellipti-
cal to parallel sided. Two (and not four!) macronuclear nodules. Contractile vacuole
about at level of cytostome. Adoral zone of membranelles about 38 % of body length.
Undulating membranes slightly curved. According to Figure 178t three frontal cirri, 1
buccal cirrus, 4 frontoventral cirri, 6 postoral and pretransverse ventral cirri, and 5 some-
666 SYSTEMATIC SECTION

what enlarged transverse cirri. Transverse cirri protrude slightly beyond posterior end of
cell. Caudal cirri lacking (?; see taxonomy).

o c cur r e n c e: Locus classicus is the Kattegat (connection between Northern and

Eastern Sea) near Varberg, Sweden (QUENNERSTEDT 1867, 1969). No other reliable re-
cords known (see taxonomy).

Species indeterminata

Histricu/us po/ycirratus AGAMALIEv, 1978, Zool. Zh., 57: 606 (Fig. 229d). Remarks: See
also AGAMALIEV (1983, p 111). Histrio polycirratus in AGAMALIEV (1976a, P 91) is a no-
men nudum. I doubt that this taxon is correctly described. About 80-90 x 50 urn. 40-42
adoral membranelles. According to illustration about 30 right and 36 left marginal cirri.
Number offrontoventral cirri increased (6 against 4 in typical oxytrichids). Caspian Sea
(locus classicus) and Bay of Biscay, Atlantic Ocean (FERNANDEZ-LEBORANs & NoVILLO-
VILLAJOS 1993).

Histrio acuminatus MAsKELL, 1886, Trans. Proc. N. Z. Inst., 19: 59 (Fig. 222f). Remarks:
See also MAsKELL (1887a). In life about 140 flm long. Margins almost in parallel, poste-
rior end somewhat pointed. Buccal area large. Frontal cirri distinctly enlarged. Five trans-
verse cirri, protrude beyond posterior end of cell. Only 1 macronuclear nodule! (possible
an exconjugant). Freshwater in Wellington, New Zealand. KAHL (1932, p 617) assumed
that it is a valid species because of the pointed posterior end. Since KAHL classified His-
trio only as subgenus of Oxytricha, the correct name in his revision is Oxytricha (Histrio)
acuminatus. CoRLISS (1960, p 275) transferred it to Histricu/us, H acuminatus. BORROR
(1972a) stated that BRANDON is the author of this species. However, since it is not clear
from the contents of the publication that BRANDON alone is responsible both for the name
and for satisfying the criteria of availability other than publication, MAsKELL and not
BRANDON is the author of the name (lem 1985, Article 50 a). BRANDON is probably the
collector of the sample.

Insufficient redescriptions

Historio sp. - WANG, 1925, Contr. bioI. Lab. Sci. Soc. China, 1: 53 (Fig. 242g). Remarks:
Perhaps a Sterkiella histriomuscorum. Small pond in Nanking, China.

Histriculus simi/is - NIETO, CALVO, MARTIN & TORRES, 1984, Acta Protozool., 23: 188
(Fig. 2321). Remarks: NIETO et a!. (1984) found this quadrinucleate population in a small
stream at Puebla del Rio, Sevilla, Spain. Thus I do not believe that it is identical with the
marine species of QUENNERSTEDT (1867) which has only 2 macronuclear nodules or with
the marine form of KAHL (1932) now classified as Sterkiella quadrinucleatus. The
straight undulating membranes and the body shape in Figure 2321 strongly resemble Sty-
 Histriculus 667

lonychia sp. Conversely, the micrographs in NIETO et al. (1984) and CALVO et al. (1986a)
indicate that it is possibly identical with Sterkiella cavicola, which, however, prefers ter-
restrial habitats. In life 140-180 x 60-85 ~m, rigid; 4 macronuc1ear nodules; 35-44 ado-
ral membranelles; rather invariably 18 frontal, ventral, transverse cirri; 52-58 left and
right marginal cirri. Further literature on this taxon: CALVO et al. (1983, 1984, 1986b,
1988).

Histriculus similis (QUENNERSTEDT) CORLISS 1960 - CHARDEZ, 1986, Acta Protozool., 25:
33 (Fig. 178u-w). Remarks: CHARDEZ (1986) found his population in a running water in
Belgium. Thus I do not believe that it is identical with the marine species of QUEN-
NERSTEDT (1867). The body shape and the resting cyst in some micrographs in CHARDEZ
(1986, Planche I, Fig. 12, 15, 16) remind strongly of Stylonychia pustulata. He also de-
scribed endogenous bud formation (Fig. 178w), which is, however, possibly a misobser-
vation. It cannot be excluded that he observed specimens feeding on their own cysts. The
deposition of the endogenous bud is possibly only the defecation of an undigested cyst
(see also H. vorax for some details, where CURDS 1966a reported a similar curiosity).
Small specimens 85-95 Jlm, large individuals 100-130 Jlm long. Two macronuclear nod-
ules, 16-20 ~m long. 35-40 adoral membranelles. 29-32 right and 22-27 left marginal
cirri. Number and arrangement of frontal, ventral, and transverse cirri, see Figure 178u.
3-4 somewhat elongated caudal cirri, a further indication that it is possibly Stylonychia
pustulata.

Histriculus (sp. nov.?) - WICKLOW, 1991, J. Protozool., 38: 2A, Abstract 11. Remarks:
WICKLOW (1991) observed predator-induced polymorphism in a (possibly new) species of
Histriculus. Freshwater, USA?

Histrio comp/anatus STOKES - CONN, 1905, Bull. Conn. St. geol. nat. Hist. Surv., 2: 60
(Fig. 232a). Remarks: Freshwater in Connecticut, USA.

Histrio complanatus STOKES - LEPSI, 1965, Protozoologie, p 35, 977 (Fig. 222g, 224f).
Remarks: Romania.

Histrio erethisticus STOKES - CONN, 1905, Bull. Conn. St. geol. nat. Hist. Surv., 2: 60
(Fig. 232b). Remarks: Incorrect subsequent spelling. The body shape and the arrange-
ment of the cirri is not at all as in H. histrio, the senior synonym. Freshwater in Connecti-
cut, USA.

Histrio sp. - CONN, 1905, Bull. Conn. St. geol. nat. Hist. Surv., 2: 60 (Fig. 232c). Re-
marks: The arrangement of the cirri is surely incorrect. Freshwater in Connecticut, USA.

Kerona histrio - FRoMENTEL, 1876, Etudes microzoaires, p 272, Planche XIV, fig. 7, 11.
668 SYSTEMATIC SECTION

Oxytricha (Histriculus) complanatus STOKES, 1887 - CHORIK, 1968, Free-living ciliates,

p 137 (Fig. 232g). Remarks: The drawing is very general so that the identification is un-
certain. Freshwater in Moldova.

Stylonychia histrio (0. F. MOLL.) - BORGER, 1905, An. Univ. Chile, 117: 436 (Fig. 232t).
Remarks: Because of the large size (in life about 280 x 140 JIm) and the body shape I
doubt that BORGER'S identification is correct, although he mentions that caudal cirri are
lacking. Chile (see also BORGER 1908).

Sterkiella FOISSNER, BLATIERER, BERGER & KOHMANN, 1991

1991 Sterkie//a FolSsNER, BLATIERER, BERGER & KOllMANN, Infonnationsberichte Bayer. Landesamtestllr
Wasserwirtschaft, 1/91: 311- Type (original designation): Oxytricha cavico/a KAm., 1935.
1997 Sterkie//a FOISSNER, BLAITERER, BERGER & KOHMANN, 1991 - BERGER & FOISSNER, Arch. Protistenk.,
148: 151.

C h a r act e r·i sat i on: Undulating membranes in Oxytricha pattern. One right and 1
left row of marginal cirri, distinctly separate posteriorly. Six dorsal kineties. Caudal cirri
present. Primordia V and VI of the opisthe originate de novo. Dorsal morphogenesis in
Oxytricha pattern.

Add i t ion a I c h a r act e r s: Both ends rounded, flattened about 2:1 dorso-
ventrally. Body firm to slightly flexible. Pellicle and cytoplasm colourless, cortical gran-
ules lacking. Movement usually rapid. Posterior postoral ventral cirrus distinctly dis-
placed posteriorly. Transverse cirri protrude distinctly beyond posterior end of cell. Usu-
ally 3 inconspicuous, that is, not distinctly elongated caudal cirri, in life and sometimes
even after protargol impregnation difficult to distinguish from marginal cirri. Dorsal cilia
in life 2-4 JIm long.

T a x 0 nom y: The characterisation is according to BERGER & FOIsSNER (1997). For a

detailed description of patterns, see chapter 2 in the general section. The type species of
Histriculus, H. histrio has no caudal cirri. Thus, we established Sterkiella to include those
Histriculus species which have (usually inconspicuous) caudal cirri. Sterkiella is well de-
fmed by a unique, but sophisticated combination of morphological characters, that is,
body rigid or only slightly flexible, undulating membranes curved and intersecting, mar-
ginal rows separate posteriorly, and caudal cirri present. However, the only autapomor-
phy so far known is the new fonnation of primordia V and VI of the opisthe (character 18
in the cladogram, Fig. 25a). Sterkiella differs from Stylonychia in that it has distinctly
bent undulating membranes and (less important) in that the caudal cirri are rather short.
Seven taxa are assigned to Sterkiella; however, only 3 of them are well defined,
namely S. cavico/a, S. histriomuscorum, S. thompsoni. Sterkiella histriomuscorum is a
sibling species complex (ForSSNER & BERGER 1998) comprising possibly also S. tricirrata
 Sterkiella 669

(5 dorsal kineties) and S. terricola (4 macronuclear nodules). In ecological studies we

suggest designating them as Sterkiella histriomuscorum complex.

Key to species
1 2 macronuclear nodules ............................................... 2
3 or 4 macronuc1ear nodules ........................................... 3
2 3 transverse cirri; 5 dorsal kineties ....................... S. tricirrata (p 693)
4-5 transverse cirri; 6 dorsal kineties (Fig. 183c, d, g) . S. histriomuscorum (p 683)
3 (l) 3 macronuc1ear nodules ............................ S. thompsoni (p 679)
4 macronuclear nodules ................................................ 4
4 In life about 400 Ilm long (Fig. 182a) ..................... S. admirabilis (p 677)
In life 100-250 Ilmlong .............................................• 5
5 Saline waters (Fig. 186b) ......................... S. quadrinucleatils (p 694)
Freshwater or terrestrial habitats ........................................ 6
6 In life 140-250llm long; 5 transverse cirri (Fig. 18Ia-l) ...... S. cavicola (p 669)
In life about 120 Ilm long; only 3 transverse cirri (Fig. 184n) .. S. terricola (p 682)

Sterkiella cavicola (KARL, 1935) FOISSNER, BLATIERER, BERGER & KOH-

MANN, 1991 (Fig. 6a, 7f, g, 21b, 181a-q, Table 40)

1888 Onychodromus grandis - MAUPAS, Archs Zoo\. expo gen., 6: 217 (Fig. 181o--q; misidentification, see
nomenclature and taxonomy).
1932 Stylonychia (Onychodromus) grandis MAUPAS - KAlIL, Tierwelt Dtl., 25: 620 (see nomenclature and
taxonomy).
1935 Oxytricha cavicola KARL, Tierwelt Otl., 30: 841 (Fig. 181n).
1950 Oxytricha magna GELEI & SZABADOSZ, AnnIs bioI. Univ. szeged., 1: 269, 288 (Fig. 181m).
1969 Opisthotricha monspessulana CH. et S., 1940 - GROLIERE, Annis Stn limno\. Besse, 4: 335 (Fig. 181j-l;
misidentification).
1970 Opisthotricha monspessulana CH. et S. 1940 - GROuERE, C. r. hebd. Senac. Acad. Sci., Paris, 270: 366
(misidentification).
1972 Oxytricha cavicola KARL, 1935 - BORROR, J. Protozoo\., 19: 13.
1984 Histriculus muscorum - MATSUSAKA, NAKAMURA & NAGATA, J. Electron Microsc., 33: 218 (misidentifi-
cation; Fig. 7f, g).
1987 Histriculus cavicola (KARL, 1935) - BERGER & FOISSNER, Zool. Jb. Syst., 114: 213 (Fig. 181a-g;
authoritative redescription; 1 slide of protargo!-impregnated specimens is deposited in the OberOsterrei-
chische Landesmuseum in Linz, Upper Austria).
1991 Sterkiella cavicola (KARL, 1935) - FOISSNER, BLATTERER, BERGER & KOllMANN, lnformationsberichte des
Bayer. Landesamtes fUr Wasserwirtschaft, 1/91: 311.
1992 Histriculus cavicola - LEHLE, Mikrokosmos, 81: 21, Bild 7 (microphotograph ofa protargol-impregna-
ted specimen).
1994 Histriculus cavicola (KARL, 1935)- SHIN & KIM, Korean J. Zoo\., 37: 114 (Fig. 181h, i).
1994 Histriculus cavicola (KARL, 1935) - SHIN, Dissertation, p 124 (Fig. 181h, i).
670 SYSTEMA TIC SECTION

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 Sterkiella 671

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Fig. 181h, i Sterkiella cavico/a (h, i, from SInN & KIM 1994. h, from life; i, protargol impregnation). h) Ven-
tral view, 160 flm. i) Ventral (left) and dorsal (right) infraciliature, 150 flm; p 669.

T a x 0 nom y: There is no doubt that the identification by MAUPAS (1888) is incorrect

because his population lacks dorsal horns. KAHL (1932) intended making a new species
for Onychodromus grandis sensu MAUPAS (1888). Unfortunately, he only transferred the
species name "grandis" to Stylonychia and attributed it to MAUPAS (1888) instead of es-
tablishing a new species. Furthennore, he classified Stylonychia only as subgenus of Oxy-
tricha so that the correct name in his revision is Oxytricha (Stylonychia) grandis. I am
not entirely sure that the synonymisation with Sterkiella cavicola is correct because "Sty-
lonychia grandis" has straight undulating membranes. Conversely, the size, the nuclear
apparatus, and the cyst are almost identical. Because of the uncertainty, I provide a sepa-
rate description of "Onychodromus grandis" sensu MAUPAS (1888).
The synonymy of S. cavicola and Oxytricha magna is evident and was already sug-
gested by BORROR (1972a). Sterkiella admirabilis also has 4 macronuclear nodules but is

-f- Fig. 181a-g Sterkiella cavico/a (from BERGER & FOISSNER 1987a a-d, from life; e-g, protargol impregnation.
a-d, f, g, Austrian population; e, Japanese population). a-c) Ventral, lateral, and dorsal view, a = 195 flm, b =
145 f1ffi, C = 165 flm. d) Cyst, diameter without protrusions about 53 flm. e-g) Ventral and dorsal infracilia-
ture, e= 105 flm, f= 150 flm, g = 135 flm; p 669.
 672 SYSTEMATIC SECTION

Fig. 181j, k Sterkie/fa cavico/a (from GROLIERE 1969. j, k, protargol impregnation). j) Ventral infraciliature.
Arrowhead marks posterior postoral ventral cirrus. k) Very early morphogenetic stage in ventral view. OP =
oral primordium, TC = left transverse cirrus; p 669.

distinctly longer (390-450 Jim) and has more adoral membranelles (50--60) and marginal
cirri. However, synonymy cannot be excluded.

M r ph 0 log y and b i 0 log y: In life 140-220 x 70-100 Jim (BERGER & FOISS-
0
NER 1987a), according to KAHL (1935) 180-250 Jim long, after GELEI & SZABADOSZ
(l950) 200 x 90 Jim, after GROLIERE (1969) 220-250 x 100-120 Jim, after SHIN & KIM
(l994) 150-220 x 70-100 Jim. Ovoid or margins almost in parallel. Flattened about 2.5:1
 Sterkiella 673

•
•

• ••
f •

Fig. 1811-q Sterkiella cavicola (I, from GROLIERE 1969; m, from <JELE! & SZABA-
DOS 1950; n, from KAlIL 1935; o-q, from MAUPAS 1888. I, protargol impregnation;
m, sublimate fixation; n-q, from life). \) Ventral infraciliature, size not indicated.
One of the 3 undulating membranes is likely a misobservation (cp. Fig. 18Ij). m,
n) Ventral views, m = 140 ~m, n = 200 ~m. 0) Ventral view of a "normal" speci-
men, 150 ~m. p) Large specimen with supernumerary frontoventral cirri, 235 ~m.
q) Cyst, 55 ~; P 669.

dorso-ventrally, anterior and posterior part very thin. Dorsal furrow pronounced, even in
overfed specimens. Macronuclear nodules in life about 20 x 15 J,lm, located along me-
dian or slightly left of it. Anterior macronuclear nodule usually slightly larger than poste-
rior one (Fig. 181e, g, Table 40). 2-6, usually 3-5, in life about 4 J,lm sized, spherical mi-
cronuclei. Contractile vacuole in about mid-body, during diastole with 2 large, dilated
collecting canals.
Adoral zone of membranelles 40-45 % of body length, bases of largest mem-
branelles in life about 11 J,lm wide. Buccal area rather broad. According to GROLIERE
(1969, 1970) and FERNANDEZ-LEBORANs (1985) a third undulating membrane is present
(Fig. 1811); however, in the micrographs by GROLIERE (1969) it is not discernible (Fig.
181j), indicating a misobservation. Invariably 3 frontal and 4 frontoventral cirri and 1
buccal cirrus, genus-specifically arranged; frontal and ventral cirri in life about 25 J,lm
long. Usually 5, rarely up to 7 postoral and pretransverse ventral cirri. Usually 5, rarely 6,
674 SYSTEMATIC SECTION

Table 40 Morphometric data of Sterkiella cavicola (cal = Austrian population, ca2 = Japanese population,
both from BERGER & FOISSNER 1987a; ca3, from SHIN & KIM 1994), S. histriomuscorum (hi! = population from
Guttal, Glockner area, Austria, hi2 = population from the SchloBalm area, Bad Hofgastein, Austria, from
BERGER et a1. 1985; hi3, from AUGUSTIN & FOISSNER 1992; hi4, from SHIN & KIM 1994), S. thompsoni (tho,
from FOISSNER 1996b), and S. terricola and S. tricirrata (ter, tri, from BUITKAMP 1977b). All data are based on
protargol-impregnated specimens. All measurements in micrometres. ? = sample size not indicated. CV = coef-
ficient of variation (in %), Max = maximum value, mean = arithmetic mean, Min = minimum value, n = sam-
ple size, SD = standard deviation

Character Species mean SD CV Min Max n

Body, length cal 135.3 25.2 18.6 88 186 21
ca2 102.9 8.3 8.1 87 121 16
ca3 155.6 13.9 8.9 123 180 16
hi! 81.7 4.5 5.5 72 88 15
hi2 64.9 5.4 8.3 57 75 15
hi3 99.0 13.2 13.3 85 129 20
hi4 105.6 4.9 4.6 100 114 10
tho 97.0 10.3 10.7 82 115 15
Body, width cal 73.9 18.4 24.9 42 112 21
ca2 53.7 6.8 12.7 43 67 16
ca3 74.4 5.7 7.6 61 82 16
hil 42.8 5.8 13.4 33 50 15
hi2 32.7 4.0 12.1 28 42 15
hi3 47.7 4.9 10.3 41 62 20
hi4 53.0 3.6 6.8 47 59 10
tho 40.1 7.1 17.7 31 56 15
Adoral zone of membranelles, cal 60.5 15.2 25.2 39 96 21
length ca2 41.6 2.1 4.9 39 46 16
ca3 66.6 5.3 7.9 60 80 16
hi! 31.1 2.6 8.2 27 37 15
hi2 27.5 1.9 6.9 25 32 15
hi3 45.2 3.8 8.5 39 53 20
hi4 43.4 2.0 4.6 41 47 10
tho 40.3 3.0 7.4 35 48 15
Adoral membranelles, number cal 48.1 9.3 19.3 32 70 21
ca2 33.3 1.5 4.5 30 36 16
ca3 37.8 1.9 5.1 35 43 16
hil 31.0 2.3 7.4 26 34 9
hi2 29.2 1.7 5.9 27 32 9
hi3 39.3 22.9 7.3 34 44 20
hi4 28.4 1.3 4.5 27 31 10
ter 22 23 ?
tho 34.4 2.1 6.0 32 39 15
tri 23 25 ?
Right marginal row, number cal 24.8 1.9 7.7 22 29 21
of cirri ca2 23.3 1.2 5.1 22 26 16
ca3 24.2 2.3 9.5 17 26 16
hi! 20.7 2.1 10.2 17 24 12
hi2 19.0 1.0 5.5 17 21 12
hi3 27.2 2.0 7.4 24 32 20
hi4 21.6 1.2 5.4 20 23 10
 Sterkiella 675

Table 40 Continued

Character Species mean SD CV Min Max n

Right marginal row, number ter 20.0 ?
of cirri tho 22.3 1.8 8.2 19 25 15
tri II 13 ?
Left marginal row, number cal 21.6 2.6 12.1 18 28 21
of cirri ca2 21.3 1.2 5.6 19 24 16
ca3 21.4 2.5 11.6 IS 24 16
hil 19.0 1.1 5.8 17 21 II
hi2 15.9 1.6 10.3 12 18 II
hi3 22.7 1.9 8.4 18 25 20
hi4 18.2 1.6 8.5 IS 20 10
ter 17.0 ?
tho 17.8 2.2 12.4 15 24 15
tri 10 13 ?
Transverse cirri, number hil 4.0 0.4 10.2 3 5 13
hi2 4.2 0.4 9.0 4 5 13
hi3 4.7 0.6 12.6 4 6 20
hi4 4.0 0 0 4 4 10
1st macronuclear nodule, lengthl cal 22.3 5.4 24.4 12 29 IS
ca2 14.5 1.1 7.5 13 17 16
ca3 17.9 2.5 14.0 13 23 16
1st macronuclear nodule, width cal 16.0 4.2 26.1 9 23 15
ca2 10.7 1.1 9.8 9 13 16
ca3 10.1 1.6 15.7 7 12 16
2nd macronuclear nodule, length cal 17.4 5.0 28.7 10 25 IS
ca2 12.1 1.5 12.2 10 IS 16
2nd macronuclear nodule, width cal 14.3 3.6 25.6 9 20 IS
ca2 9.2 0.7 7.1 8 10 16
3rd macronuclear nodule, length cal 18.1 5.6 30.8 10 28 IS
ca2 12.9 2.0 15.3 10 16 16
3rd macronuclear nodule, width cal 13.6 2.8 20.5 9 19 15
ca2 8.9 0.6 7.3 8 10 16
4th macronuclear nodule, length cal 20.4 5.4 26.6 12 28 IS
ca2 12.1 1.5 12.0 10 14 16
ca3 15.8 2.4 15.4 12 20 16
4th macronuclear nodule, width cal 13.2 2.7 20.7 9 18 15
ca2 9.2 1.0 10.8 7 II 16
ca3 9.2 1.3 14.5 7 12 16

I This is the anterior macronuclear nodule.

in life about 35 Ilm long, slightly enlarged transverse cirri. Marginal cirri in life 20 Ilm
long; right marginal row begins almost at level of right frontal cirrus, anterior-most cirrus
of this row usually slightly detached and not involved in primordia formation of right
marginal row of proter (Fig. 181e, f). Dorsal kineties 3 and 4 slightly shortened
anteriorly, kineties 5 and 6 distinctly shortened posteriorly (Fig. 18Ig). Caudal cirri in
life about 20 Ilm long, arranged at end of dorsal kineties 1,2, and 4 (Fig. 18Ig).
676 SYSTEMATIC SECTION

Cysts spherical, in life about 48-62 !lm (mean = 55 !lm; n = 25) across, with an ir-
regularly serrated surface, teeth about 2.5 !lm high. Inner wall about 1.4 !lm thick. Cyto-
plasm densely filled with small globules (Fig. 181 d). In most strains several days are
needed for the cysts to respond to excystment stimuli (NAKAMURA 1989). The results by
NAKAMURA & MArsusAKA (1991) suggested that excystment is influenced, among other
things, by the age of the cysts, strain differences, and the medium in which the cysts were
formed or aged. Further details on the cyst and on en- and excystment, see CALVO & DE
MIGUEL (1995/96), CALVO et al. (1992), LOPEZ et al. (1995), MArsusAKA (1977, 1979),
MArsusAKA & HONGO (1984), MArsusAKA & KIMuRA (1981), MArsusAKA et al. (1984),
NAKAMURA & MArsusAKA (1985, 1992a, b), and VELASCO et al. (1992,1995).
Southern hybridisation analysis using an a-tubulin probe from Stylonychia lemnae
have been performed in order to estimate the number of a-tubulin genes present in the ge-
nome of Sterkiella cavicola (DELGADO et al. 1994, 1995). The analysis revealed a close re-
lation between Euplotes and Sterkiella and suggested that both taxa are really closer than
generally suggested by rRNA analysis. Further literature: PEREZ-ROMERO et al. (1998).
Morphogenesis of cell division is described by GROLIERE (1969, 1970); however, I
doubt that all illustrations are correct. Figure SA in GROLIERE (1969), for instance, shows
a late stage with 8 frontal-ventral-transverse cirral streaks. The oral primordium origi-
nates near the left transverse cirrus (Fig. 181k).
Morphology and biology of "Onychodromus grandis" sensu MAUPAS (1888; Fig.
18lo-q): Well-fed specimens (giants) in life about 300 x l50!lm (Fig. l81p), small
(probably normal) specimens about 150 !lm long and 2.3 times as long as broad (Fig.
1810). Large specimens are bulging, small ones with parallel margins. Both ends
rounded. Contractile vacuole in about mid-body. Cytoplasm cloudily granulated. Large
specimens with about 12 frontal, buccal, and frontoventral cirri (Fig. 181 p), small speci-
mens with the typical number of 8 cirri (Fig. 1810). Invariably 5 postoral and pretrans-
verse ventral cirri and 5 transverse cirri. "Stylonychia grandis" sensu HEMBERGER (1982,
P 233), however, usually has 4 transverse cirri. According to the illustration by MAUPAS
(1888) only the posterior-most transverse cirrus protrudes distinctly beyond posterior end
of cell. Caudal cirri arranged slightly right of median. Cyst in life about 50 !lm across,
with prominent, blunt processes (Fig. l8lq).

o c cur r e n c e and e colo g y: Widely distributed in terrestrial habitats and some-

times in the aufwuchs and detritus of freshwater. In litter of beech-forests often associated
with Rigidocortex octonucleatus and Territricha stramenticola. Probably cosmopolitan.
Locus classicus is a tree hole with water in Germany, presumably near Hamburg,
where KARL lived. GELEI & SZABADOS (1950) discovered the synonym, Oxytricha magna,
in rain puddles in Szeged, Hungary. Further records, largely substantiated by illustrations:
upper soil layer of an arable field near the town of Salzburg, Austria (BERGER & ForssNER
1987a, "Versuchsflache D" in ForssNER et al. 1987a); litter of beech-forests in Germany
and Austria (BONKOWSKI 1996, LEHLE 1989, 1992, own observations); pond in a forest
near Clermont-Ferrand, France (GROLIERE 1969, 1970, GROLrERE & NJINE 1973); soil on
 Sterkiella 677

the campus of the Faculty of Science, Kumamoto University, Japan (MATSUSAKA 1977,
MATSUSAKA et al. 1984); terrestrial habitats near Seoul and on Ullung Island, Korea (SlDN
& KIM 1994, 1995); soil in Amazonian rain forest near Iquitos, Peru (FOisSNER 1997b);
soil (0-3 cm, pH 4.7) from cloud rain forest near summit of Monteverde Preserve, Costa
Rica (FOISSNER 1997b); soil south of Sydney, Australia (BLATIERER & FOISSNER 1988).
Records by ROHNERT (1951; Oxitricha cavicola par lapsus) from tree holes of Austria and
Germany are not substantiated by illustrations.
Feeds on bacteria (Enterobacter aerogenes, E. cloacae, Chromobacter violaceus),
autotrophic and heterotrophic flagellates (Chlamydomonas sp., Chilomonas sp., Pan-
dorina morum, Gonium sp.), fungal spores, cysts of naked amoebas, ciliates (Nassula sp.,
Hastatella sp., Tetrahymena sp.); also cannibalistic {BERGER & FOISSNER 1987a, GEi.EI &
SZABADOSZ 1950, GROLIERE 1969, KARL 1935). MATSUSAKA et al. (1984) cultured Sterk-
iella cavicoia at 25°C in 0.1 % (w/v) lettuce extract in Petri dishes with Tetrahymena sp.
as food. SENDO & MATSUSAKA (1982) used following culture method to obtain mass cul-
ture: a 200 ml aliquot of 0.1 % lettuce extract in a 500-ml Erlenmeyer flask was inocu-
lated with 2.8 x 106 Tetrahymena sp. and incubated for 30-40 h at 25°C. The flask was
then maintained at 25 °C while positioned on a reciprocal shaker set for 70 strokes min'\
with an amplitude of 3 cm. After a lag phase of 10 h, the cultures entered an exponential
growth phase. Subsequent to a short period during which the growth rate decelerated, the
cultures reached a stationary phase about 60 h after inoculation. The cell density at the
stationary phase was about 1000 cells ml'\ and generation time during exponential phase
was about 6 h. Biomass of 106 individuals about 540 mg (FOISSNER 1987a).
Records of "Onychodromus grandis" sensu MAUPAS (1888): MAUPAS found it in an
infusion of cress collected in Algeria. Records not substantiated by illustrations: once
with high abundance at a water temperature of 10°C in the Tiber River, Italy (STElLA &
CAMPEA 1948); mud ofa fishpond in Poland (CZAPIK 1959); in sand, gravel, and silt sedi-
ments of small rivers in Latvia (LIEPA 1990, VEYLANDE & LlYEPA 1985); freshwater habi-
tats in Japan (MATSUOKA et al. 1983); Copper Creek, Colorado, USA (CAIRNS & YONGUE
1973a). Feeds on small and large ciliates.

Sterkiella admirabilis (FOISSNER, 1980) comb. nov. (Fig. 182a-e)

1979 Histriculus admirabilis- FOISSNER & ADAM, Jb. Univ. Salzburg, 1977-1979: 153 (nomen nudum).
1980 Histriculus admirabilis FOISSNER, Ber. Nat.-Med. Ver. Salzburg, 5: 108 (Fig. 182a--c).
1988 Stylonychia quadrinucleata ALEKPEROV & MUSAYEV, Zoo!. Zh., 67: 1907 (Fig. 182d, e; new synonym).

Nom e n c I at u rea n d t a x 0 nom y: FOISSNER & ADAM (1979) published His-

trieulus admirabilis in a species list without description. The large size, the limnetic habi-
tat, the shape of the undulating membranes, and many morphometric data separate it from
other quadrinucleate species of the Oxytrichidae, such as Sterkiella cavicoia, S. quadri-
nucleatus, or Cyrtohymena quadrinucleata. Redescription necessary.
678 SYSTEMATIC SECTION

Fig. 182a~ Sterkiella admirabilis (a-c, from FOISSNER 1980a; d, e, from ALEKPEROV & MUSAYEV 1988. a-c,
from life; d, wet silver impregnation; e, Feulgen stain). a, b) Ventral and lateral view, a =415 JIm, b = 430 JIm.
c, d) Ventral infi"aciliature, c =? JIm, d = 350 JIm (according to bar 490 JIm long!). Note that in (d) only 2 pos-
toral ventral cirri are shown. e) Nuclear apparatus. CV = contractile vacuoles, Ma = posterior macronuclear
nodule, Mi = micronucleus; p 677.

The superficially described Stylonychia quadrinucleata is very likely not a Stylony-

chia because the undulating membranes are distinctly curved. (Fig. 182d). The size, the
nuclear apparatus (two pairs of nodules), and the number of adoral membrane lies (55-60)
are rather similar to in S. admirabilis, indicating synonymy.

M 0 r ph 0 log y and b i 0 log y: In life 390-450 !lm long, about 2.5-3.1 times as
long as broad (FOISSNER 1980a), according to ALEKPEROV & MUSAYEV (1988) about
350 !lm long (see, however Fig. 182d). Body margins more or less parallel. Anterior and
posterior strongly flattened dorso-ventrally (Fig. 182b). Slightly flexible (but never as
much as in Oxytricha species). Macronuclear nodules usually arranged in 2 distinct pairs
slightly left of median. Usually 4, sometimes only 2 micronuclei. Two contractile vacu-
oles, namely a small one at distal end of undulating membranes, and a large one at level
of cytostome. Cytoplasm with few, large, orange-yellow inclusions and opaque by many
irregularly shaped granules.
Adoral zone of membranelies almost 50 % of body length, about 50 membranelles
(FOISSNER 1980a), according to ALEKPEROV & MUSAYEV (1988) 55-60 membranelles.
Rather invariably 3 frontal and 4 frontoventral cirri and 1 buccal cirrus. Usually 3 pos-
toral and 2 pretransverse ventral cirri; however, occasionally up to 10 ventral cirri. Usu-
 Sterkiella 679

ally 5, sometimes 6 transverse cirri. All frontal, ventral, and especially transverse cirri
distinctly enlarged. About 30 left and 42 right marginal cirri (ALEKPEROV & MUSAYEV
1988). Three inconspicuous caudal cirri in median of cell (FOISSNER 1980a).

o c cur r e n c e and e colo g y: Locus classicus is a melt water pool (pond 23 in

FOISSNER 1980c) north of the FuschertOrl, GroBglockner HochalpenstraBe, Austrian Alps,
about 2400 m above sea-level. FOISSNER (1980a, 1980c) found S. admirabilis also in the
benthic area, the periphyton, and the pelagial of some other small water bodies near the
GroBglockner. Locus classicus of the synonym, Stylonychia quadrinucleata, is a garden
soil on the Apsheron peninsula, Azerbaijan (ALEKPEROV & MUSAYEV 1988).
Feeds on bacteria and coccale green algae (FOISSNER 1980a, c). FOISSNER & ADAM
(1979) and FOISSNER et al. (1982a) provided following autecological data (6 analyses):
b-a, 0.2 x 106 to 5 X 106 bacteria mI- 1 (direct counting), 4.6-14.4 °C, 8.1-9.9 mg 1-1 O2
(85~124 % saturation), pH 4.8-5.0, 1.1-2.0odH total hardness, 36-54 mg 1-1 KMn04-
consumption, ~.31 mg 1-1 NH/-N.

Sterkiella thompsoniFOISSNER, 1996 (Fig. 182f-k, Table 40)

1964 Opisthotricha sp. - SUDZUKI, Sci. Rep. Jap. Antarctic Res. Expedition, 19: 5, 11 (Fig. 224g; see taxon-
omy).
1972 Oxytricha sp. - THOMPSON, Antarctic Terrestrial Biology Antarctic Res. Ser., 20: 283 (Fig. 182i,222a;
see taxonomy).
1996 Sterkiella thompsoni FOISSNER, Acta Protozool., 35: 112 (type slides are deposited in the OberOsterrei-
chische Landesmuseum in Linz, Upper Austria).

T a x 0 nom y: THOMPSON (1972) provided a description of this species after silver im-
pregnation, but did not name it because he considered his data insufficient. However,
they match the original description very well. Conversely, SUDZUKI'S (1964a) population
is rather superficially described; only the number of macronuclear nodules and the loca-
tion (Antarctica) indicate that it is S. thompsoni. Three macronuclear nodules are very
rare in hypotrichs and ciliates in general. However, the detailed morphometric characteri-
sation (see below) and the records by SUDZUKI and THOMPSON prove that S. thompsoni is
not a teratological population of the very common S. histriomuscorum (2 nodules) or S.
cavicola (4 nodules). SESHACHAR & KAsTIJRI BAI (1963) very briefly described a ''new
species of Oxytrichd' from India also having 3 macronuclear nodules; however, this still
unnamed population is much larger, namely 200-450 x 100-150 Jlm (Fig. 222c).
The classification of S. thompsoni is somewhat uncertain because the dorsal ciliature
originates in the Urosomoida pattern (fragmentation ofkinety 3 lacking; Fig. 24b), which
is common in the Oxytrichinae, especially in Urosomoida and Urosoma. All other char-
acters (body inflexible, lack of cortical granules, long adoral zone of membranelles, dis-
tinct fibres around dorsal cilia) strongly indicate that it is a Stylonychinae. The present
classification implies that the Urosomoida pattern (loss of fragmentation of kinety 3)
680 SYSTEMATIC SECTION

.-
'\
-. .

---.. -- --..,

-----, /'
./'

,,

.
•
. ..• ... •• . t
 Sterkiella 681

evolved twice independently. Perhaps detailed ontogenetic data will reveal more insight
into the true position of this species.

M 0 r p hoi 0 g y and b i 0 log y: In life about 90-130 x 40-60 ~m. Usually

broadly parallel-sided, rarely slightly bursiform, anterior end broadly rounded, posterior
one narrowly rounded and often inconspicuously pointed. Rather rigid like, for example,
S. histriomuscorum and Stylonychia pustulata, dorsoventrally flattened up to 2: 1. Usually
3, very rarely 2 (in 6 out of 120 specimens; possibly postdividers) or 4 slightly to dis-
tinctly ellipsoidal macronuclear nodules in median of middle third of cell; middle nodule
often slightly smaller than anterior and posterior ones; nodules after protargol impregna-
tion 11-18 ~m long (mean = 13.1 J.1m, SO = 3.5 J.1m, CV = 26.5 %, n = 15) and 7-12 J.1m
broad (mean = 8.9 J.1ffi, SO = 1.5 ~m, CV = 17.0 %, n = 15). Micronuclei almost spheri-
cal, after protargol impregnation 2-3 x 1.5-2.1 J.1m, number highly variable (1--6, mean =
2.0, SD = 1.7, CV = 84.2 %, n = 13), however, most specimens have only one. Contrac-
tile vacuole in mid-body with 2 long collecting canals extending anteriorly and posteri-
orly, anterior canal often with small dilations at level of buccal cavity. Cortex colourless,
rigid, without special granules. Cytoplasm colourless, contains many small, yellowish
crystals, some 2--6 J.1m sized, colourless fat globules, and food vacuoles up to 10 J.1m in
diameter. Moves moderately fast, often resting for some time and thus easy to observe.
Adoral zone of membranelles about 42 % of body length. Buccal cavity rather large
and deep, right margin thickened, possibly by fibres or backward directed endoral cilia,
right half of cavity covered by hyaline lip. Paroral and endoral slightly curved, incon-
spicuously to distinctly intersecting optically, both very likely composed of dikinetids.
Pharyngeal fibres inconspicuous. Frontal cirri and transverse cirri in life about 30 J.1m
long. Frontal and transverse cirri slightly enlarged. Arrangement of frontal-ventral-
transverse cirri as in other Sterkiella's (Fig. 182j). Invariably 3 frontal,4 frontoventral,
and 2 pretransverse ventral cirri (n = 15), 1 buccal cirrus (n = 15), usually 3 (mean = 3.1,
SD = 0.6, CV = 19.3 %, Min = 2, Max = 5, n = 15) postoral ventral cirri and usually 5
(mean = 5.1, Min = 5, Max = 6, n = 15) transverse cirri. Marginal rows distinctly separate
posteriorly, gap occupied by caudal cirri right of cell median. Invariably 4 (n = 15) dorsal
kineties. Kinety 1 slightly shortened anteriorly, kineties 2 and 3 ± as long as body, kinety
4 slightly shortened posteriorly. Dorsal cilia in life about 3 ~m long, associated with dis-
tinct, oblique fibrillar structures. Invariably 2 (n = 15), in life about 30 ~m long caudal
cirri, one each on kineties 1 and 3 (Fig. 182k). Dorsal morphogenesis probably in Uroso-
moida pattern (Fig. 24b), that is, fragmentation ofkinety 3 lacking.

~ Fig. 182f-k Sterkiella thompsoni (f-h,j, k, from FOISSNER 1996b; i, from THOMPSON 1972. f, g, h, from life; i,
wet silver impregnation; j, Ie, protargol impregnation). f) Ventral view of representative specimen, 105 11m. b,
g) Ventral and lateral view of oviform specimen showing oral apparatus and contractile vacuole (g). i, j) Ven-
tral infraciliature, i = about 100 11m, j = 110 11m. k) Dorsal infraciliature, 110 11m. Large arrow marks posterior
end of dorsal kinety 4 (which is possibly a dorsomarginal row), tiny arrows denote fusiform fibre bundles sur-
rounding dorsal cilia; p 679.
682 SYSTEMATIC SECTION

Opisthotricha sp. sensu SUDZUKI (1964a; Fig. 224g) in life 96-120 x 40-80 JIm. 2-3
macronuclear nodules. More than 23 micronuclei (misobservation?). 20-26 right mar-
ginal cirri, marginal rows separate posteriorly. Three caudal cirri.
Oxytricha sp. sensu THOMPSON (1972; Fig. 182i) after wet silver nitrate impregnation
91-120 x 39-50 JIm (mean = 108 x 43 JIm; n = 25). Both ends rounded. 3-4 oval to
elongate macronuclear nodules in posterior 2/3 of cell. Adoral zone of membranelIes
51 JIm long on average. Four dorsal kineties.

o c cur r e n c e and e colo g y: Possibly endemic to Antarctica. Locus classicus is

a Drepanocladus uncinatus moss from Signy Island, South Orkney Islands, Antarctica
(ForSSNER 1996b). THOMPSON (1972) found S. thompsoni in a rock pool at the highest ele·
vation behind Old Palmer Station, Antarctic Peninsula. SUDZUKJ (1964a) isolated it from a
moss sample from Langhovde, Antarctica. Further records: moss near Casey Station and
fellfield south-west of Casey Station, Wilkes Land, continental Antarctica (PETZ & Forss-
NER 1997a). Feeds on bacteria, heterotrophic flagellates, green algae, diatoms and wheat
starch (ForSSNER 1996b).

Sterkiella terricola nom. nov. (Fig. 184n, 0, Table 40)

1977 Histriculus similis (QUENN.) CORUSS, fonna tricirratus BUITKAMP, Acta Protozool., 16: 273 (Fig. 184n,
0).
1986 Histriculus tricirratus BUITKAMP, 1977 - DRAGESCO & DRAGESCO-KERNEIS, Faune tropicale, 26: 485 (see
nomenclature).

Nom e n cIa t u rea n d t a x 0 nom y: BUITKAMP (1977b) published the name "tri-
cirratus" expressly only for a "forma" of Histriculus similis. Thus, according to Articles
10c, 23j, 45g, and 50c (i) of the leZN (1985) DRAGESCO & DRAGESCo-KEloo:IS (1986) are
the authors of Histriculus tricirratus because they first used this name for the species
rank. However, both H. tricirratus DRAGESCO & DRAGESCO-KERNEIS, 1986 and Oxytricha
tricirrata BUITKAMP, 1977 have to be transferred to Sterkiella so that they become secon-
dary homonyms and the junior is invalid. Thus, the DRAGESCO & DRAGESCo-KERNEIS
(1986) species requires a replacement name in Sterkiella: S. terricola nom. nov. (basio-
nym: Histriculus tricirratus). Histriculus triccirratus BUITKAMP, 1977 in DRAGESCO &
DRAGESCO-KERNEIS (1986, P 485) is an incorrect spelling. Differs from Sterkiella his-
triomuscorum in the number of macronuclear nodules and transverse cirri; however, syn-
onymy cannot be excluded. Detailed redescription necessary.

M 0 r p hoI 0 g y and b i 0 log y: In life (?) about 120 JIm long, elliptical. Macronuclear
nodules arranged in a chain left of median. 4 micronuclei. Contractile vacuole in about
mid-body. Arrangement of frontal, ventral, and transverse cirri Sterkiella-typical, but with
small cirrus between middle and right frontal cirrus (Fig. 184n, arrow). Cirri in life about
15 JIm long. 6 dorsal kineties. Oral primordium originates at left transverse cirrus.
 Sterkiella 683

o c cur r e n c e and e colo g y: Locus classicus is the soil of a unburned savannah

in the Ivory Coast. Not found since. Feeds on bacteria and cyanobacteria. Biomass of 106
individuals about 234 mg (FOlSSNER 1987a; value to high), according to BUITKAMP (1979,
as Histriculus similis) only 84 mg. BUITKAMP (1979) found 29 and 54 indo g-I dry soil in
cultures incubated at 20°C and 35 °C, respectively.

Sterkiella histriomuscorum (FOISSNER, BLATTERER, BERGER & KOHMANN,

1991) FOISSNER, BLATTERER, BERGER & KOHMANN, 1991 (Fig. 6b, lOa-v,
20a, 183a-l, 184a-m, 186e, f, Tables 3, 4, 40,41,52, Addendum 2)

1932 Histrio muscorum KAlIL, Tierwelt Dtl., 25: 617 (Fig. 184k).
1938 Stylonychia curvata - GIESE & ALDEN, J. expo Zool., 78: 117 (Fig. 186f; misidentification).
1950 Opisthotricha terrestris HORVA1ll, AmIls Inst. BioI. hung., 19: 157, 158 (nomen nudum; see nomencla-
ture).
1953 Histrio simi/is (QuENNERSTEDT 1867) - WENZEL, Arch. Protistenk., 99: 113 (Fig. 184h; misidentifica-
tion).
1954 Oxytricha minor- MOTE, Proc. Iowa Acad. Sci., 61: 578, 588 (Fig. 1841; misidentification).
1954 Ciliate sp. No. 1- MOTE, Proc.lowaAcad. Sci., 61: 578, 588 (Fig. 184m).
1956 Opistotricha te"estris HORVA1ll, Arch. Protistenk., 101: 275 (Fig. 183i,j; new synonym; incorrect sub-
sequent spelling of Opisthotricha).
1957 Oxytricha histrioides GELLERT, AnnIs Inst. bioI. Tihany, 24: 20 (Fig. 1831; new synonym).
1958 Histrio macrostoma GELLERT & TAMAs, AnnIs Inst. bioI. Tihany, 25: 229, 240 (Fig. 183k; new
synonym).
1960 Histriculus muscorum (KAlIL, 1932) - CORUSS, J. Protozool., 7: 275.
1970 Opistotricha terrestris HORVA1ll - DEUIEZ & CHARDEZ, Annis Speltol., 25: 135 (Fig. 186e; incorrect
subsequent spelling of Opisthotricha).
1970 Histriculus muscorum KARL, 1932 - DRAGESCO, AmIls Fac. Sci. Univ. fed. Cameroun (Numero hors-
serie): 116 (Fig. 184g).
1972 Histriculus muscorum (KARL, 1932) CORUSS, 1960- BORROR, J. Protozool., 19: 14.
1972 Oxytricha bimembranata SIllBUYA, 1929 - MAns & DANISKovA, Acta Fac. Rerum nat. Univ. comen.,
Bratisl., 17: 49 (Fig. 184i,j; misidentification).
1982 Histriculus muscorum KARL, 1932 - FOISSNER, Arch. Protistenk., 126: 80 (Fig. 184a-f; authoritative re-
description; 1 slide of protargol-impregnated specimens is deposited in the Oberosterreichische Landes-
museum in Linz, Upper Austria).
1985 Histriculus muscorum (KARL, 1932) - BERGER, FOISSNER & ADAM, Protistologica, 21: 303 (Fig. lOa-v).
1986 Histriculus muscorum KARL, 1932 - DRAGESCO & DRAGESCo-KERNEIS, Faune tropicale, 26: 483.
1986 Oxytricha terrestris (HORWA1ll, 1958) - DRAGESCO & DRAGESCO-KERNEIS, Faune tropicale, 26: 471.
1989 Oxytricha trifaUax GRESLIN, PRESCOTT, OKA, LoUKIN & CHAPPELL, Proc. natn. Acad. Sci. U. S. A, 86:
6264 (see nomenclature).
1991 Oxytricha histriomuscorum - FOISSNER, BLATTERER, BERGER & KOHMANN, Informationsberichte des
Bayer. Landesamtes filr Wasserwirtschaft, l/91: 311 (see nomenclature).
1991 Sterkiella histriomuscorum (FOISSNER, BLATTERER, BERGER & KOHMANN, 1991) - FOISSNER, BLATTERER,
BERGER & KOHMANN, Informationsberichte des Bayer. Landesamtes filr Wasserwirtschatl, l/91: 311 (see
nomenclature).
1992 Histriculus muscorum - LEHLE, Mikrokosmos, 81: 21, Bild 6 (microphotograph of a protargol-
impregnated specimen).
684 SYSTEMATIC SECTION

1992 Sterkiella histriomuscorum (FOisSNER, BLATIERER, BERGER & KOHMANN, 1991) FOISSNER, BLATIERER,
BERGER & KOHMANN, 1991 - AUGUSTIN & FOISSNER, Arch. Protistenk., 141: 279 (Fig. 183a-e).
1994 Histriculus muscorum KAHL, 1932 - SHIN & KIM, Korean J. Zoo!., 37: 115 (Fig. 183f-h).
1994 Histriculus muscorum KAHL, 1932 - SHIN, Dissertation, p 131 (Fig. 183f-h).

Nomenclature and taxonomy: KAHL (1932) classified Histrio as subgenus of Oxy-

tricha; the correct name in the original description is thus Oxytricha (Histrio) muscorum.
Consequently, it is a primary homonym of 3 other O. muscorum species described by
KAHL (1932; namely O. (Steinia) muscorum, O. (Opisthotricha) muscorum, and 0. (Sty-
/onychia) muscorum) because the presence of a subgeneric name does not affect ho-
monymy between species group names (leZN 1985, Article 57d). This was always over-
looked and replacement names were only provided by ForSSNER et al. (1991). At that time
I did not consider "Opistotricha terrestris", Oxytricha histrioides, Histrio macrostoma,
or Oxytricha trifallax as synonyms of "Histrio muscorum" and thus proposed the re-
placement name Oxytricha histriomuscorum (lCZN 1985, Article 60a). After the addition
of these synonyms it would seem appropriate to use the name of the oldest synonym,
namely "Opistotricha terrestris" HORvArn (lCZN 1985). Nevertheless, I continue to use
the replacement name in order to stabilise the nomenclature of this common species
(ForSSNER & BERGER 1999). HEMBERGER (1982, p 239) proposed "Sty/onychia para/lela"
as replacement name for "Histrio muscorum KAHL". However, since his thesis is unpub-
lished, this name is not available. "Sty/onychia parallel/a" in DRAGESCO & DRAGESCo-
KERNElS (1986, P 485) is an incorrect subsequent spelling of HEMBERGER'S name.
The name Opisthotricha terrestris, published by HORvArn (1950), was not accompa-
nied by a description allowing a differentiation from other taxa so that it must be consid-
ered as nomen nudum. The body size, the outline of the cell, and the ventral (especially
the distinctly posteriorly displaced postoral ventral cirrus V/3 and the 4 transverse cirri)
and dorsal infraciliature are reminiscent of Sterkiella histriomuscorum. However, accord-
ing to Table B in HORvArn (1950) O. terrestris has a flexible body, whereas it is almost
frrm in S. histriomuscorum. Nevertheless, I consider them as synonyms as both species
were discovered in terrestrial habitats.
The synonymy of S. histriomuscorum and Histrio macrostoma is evident and was al-
ready proposed by HEMBERGER (1982, P 239). The little-known, terrestrial Oxytricha his-
trioides invariably has 4 transverse cirri and is very likely a further synonym of S. his-
triomuscorum, although GEllERT (1957) interpreted the body as metabolic and the pos-
toral ventral cirrus V/3 is not very distinctly posteriorly displaced. It could thus also be a
synonym of Oxytricha auripunctata or O. quadricirrata, which also have only 4 trans-
verse cirri but are smaller and have cortical granules. BORROR (1972a) synonymised O.
histrioides with 0. tal/ax, which is, however, confined to limnetic habitats and has 5
transverse cirri.
WENZEL (1953) synonymised S. histriomuscorum with Histriculus similis, which is in
fact rather similar (Fig. 178t). However, I avoid a synonymisation because no detailed re-
description of a marine population exists. Since it cannot be excluded that H. simi/is in
fact lacks caudal cirri, it is preliminarily not transferred to Sterkiella.
 Sterkiella 685

BC

FVC

PVC

RM f
••
f

Fig.183a-c Sterkiella histriomuscorum (from AUGUSTIN & FOISSNER 1992. a, b, from life; c, protargol impreg-
nation). a, b) Ventral and left lateral view, a = 105 ~m, b = 120 ~m. c) Ventral infraciliature, 105 ~m. BC =
buccal cirrus, FVC = frontoventral cirri, LMR = left marginal row, PVC = postoral ventral cirri, RMR = right
marginal row, TC = transverse cirri; p 683.

Histriculus muscorum sensu DRAGESCO (1970; Fig. 184g) and Zou & ZHANG (1992)
have, among other things, undulating membranes in Stylonychia pattern, indicating mis-
identification. Morphogenetic features (formation of oral primordium left of postoral
ventral cirri; origin of primordia from frontoventral cirrus IV/3) in Zou & ZHANG (1992)
suggest that the Chinese population is possibly Stylonychiapustulata. Histriculus mus-
corum sensu REUTER (1961) also has straight undulating membranes and the transverse
cirri are arranged in two distinct groups; it is thus classified as Stylonychia hi/aria (Fig.
159a).
Recently, we (ForSSNER & BERGER 1998) studied an original strain of "Oxytricha tri-
fallax", a nomen nudum species (see p 254) because no morphological data are pub-
lished. We found no significant differences to Sterkiella histriomuscorum. GRESLIN et al.
(1989) got it from Robert lIAMMERSMITII, Ball State University. Further literature and in-
formation on "0. tri/allax", see p 254. Oxytricha nova also belongs to the Sterkiella his-
686 SYSTEMATIC SECTION

.,
'. '

W;o:::"::::-.
;~:::::'':!::

Mi+-=--t

'-.... I I / J J

.
II \ I / 1 J
f ~ \ J I
I
j
J

~ \ / I I
#
, ~
§: \
\
\ J
I
J
J
j
I
; ~,
,~ , \\ . \ ( )
,
.'

\r:~

f~ t}
/
\
.
, )
\ 0'· :.•. / )
, I
, \ \ I ) )
,

oCI.
)
\ J
, \ )
\ I
~
, )
, \. ,.'.•' I J
.
". \. ~. I /
; , .. I /

••••
\. ! /
'. I /
\.
I
"" /
• •
\
)
\
9
'- h
"
 Sterkiella 687

Fig. 183i-1 Sterkiella histriomuscorum (i, j, from HORVAlH 1956; k, from GELLERT & TAMAs 1958; L from
GELLERT 1957. i-l, opal-blue stain after BRESSLAU). Ventral and dorsal infraciliature, i,j = 80 Jun, k = 110 11m,
1= 140-150 11m; p 683.

triomuscorum complex (FOISSNER & BERGER 1999); further details and literature, see spe-
cies indeterminata in Oxytricha (p 250).

M 0 r p hoI 0 g y and b i 0 log y: In life 80-160 x 40-70 !lID, after KAm.. (1932)
100-150 !lm, according to WENZEL (1953) up to 180!lm long, after HORvAru (1956)
80 !lm long and 2 times as long as broad, after GELLERT (1957) about 140-150 !lm long
and 2.5 times as long as broad, according to FOISSNER (1982) 100-150 x 40-60 !lm. Body
usually slightly flexible, that is, not as rigid as, for example, in Stylonychia mytilus but
not as flexible as, for example, in Oxytricha granulifera; very probably because of this
intermediate state of flexibility it was often classified in Oxytricha. Body margins usually
parallel, both ends rounded. Macronuclear nodules in life about 20-30 x 12-20 !lID, after
protargol impregnation 9-21 x 5-12!lm (mean = 14-17 x 7-10 !lm; n = 60), arranged
slightly left of median. Distance between macronuclear nodules after protargol impregna-
tion 2-15 !lm (mean = 6-9 !lm; n = 60). 1-3, usually 2 micronuclei, after protargol im-
pregnation 2-3 !lm across. Contractile vacuole slightly in front of mid-body.

~ Fig. 183d-b Sterkiella histriomuscorum (d, e, from AUGUSTIN & FOISSNER 1992; f-h, from SHIN & KIM 1994.
d, g, h, protargol impregnation; e, silver carbonate impregnation; f, from life). d, b) Dorsal infraciliature, d =
105 11m, h = 110 11m. e) Adoral zone of membranelies and undulating membranes. C, g) Ventral view and ven-
tral infraciliature, f= 100 11m, g = 110 11m. AM = adoral membranelles, CC = caudal cirri, e = endoral, Ma =
macronuclear nodule, Mi = micronucleus, p = paroral; p 683.
688 SYSTEMATIC SECTION

•
.•
•
•
"
III

"... • .• b c

, ~
"
, "
, ,
~

,, II

• "\PRV

... .
I

-.
. , III
• • J C
CC-" .~~---. - TC
~~~
f
- .
 Sterkiella 689

Adoral zone of membranelles about 35-40 % of body length. Buccal area medium
sized (Fig. 183a, 184a, k). Undulating membranes according to AUGUSTIN et al. (1986)
and AUGUSTIN & FOISSNER (1992) composed of a double row of basal bodies (Fig. 183e),
after FERNANDEZ-LEBORANs (1985) and FERNANDEz-LEBORANS & MARTIN-GONZALEZ (1982)
consist of a single row with a parallel fibre. Frontal and transverse cirri in life about
30 J.1m, marginal cirri about 20 J.1m long. Usually 5, rarely 4 or 6 postoral and pretrans-
verse ventral cirri. Some populations predominantly with 4, some predominantly with 5
transverse cirri. Rather invariably 6 (Fig. lOb, 183b), according to AUGUSTIN & FOISSNER
(1992) rarely 7 dorsal kineties. Kinety 4 slightly shortened anteriorly, kineties 5 and 6
distinctly shortened posteriorly. Kinety 1 consists of 16-28 (n = 40) basal body pairs, ki-
nety 2 of 17-26 (n = 32), kinety 3 of 11-19 (n = 32), kinety 4 of 7-19 (n = 36), kinety 5
of6-14 (n = 40), and kinety 6 of 2-7 (n = 52). Invanably 3 caudal cirri on dorsal kine-
ties 1,2, and 4.
M 0 r ph 0 g e n e sis of cell division is described by NIETO et al. (1984; without
figures), BERGER et al. (1985), and PETZ & FOISSNER (1997a; see Addendum 2). It pro-
ceeds very similarly to in related species, so that only some details are mentioned (Fig.
lOa-v, Table 4). Stomatogenesis commences just left of the left transverse cirrus and the
proliferation of basal bodies yields a long and narrow oral primordium. About 88 % of
the specimens investigated by BERGER et al. (1985) possess 4 transverse cirri, 6 % have 5,
and 6 % have only 3 transverse cirri. The highly dominant set of 17 frontal, ventral, and
transverse cirri arises within the streaks I-VI as follows: 1,2, 3, 3,4,4. The right mar-
ginal primordium of the proter originates from the second (from anterior) cirrus (Fig.
10i). Morphogenesis of dorsal ciliature is in Oxytricha pattern (Fig. 100).

o c cur r e n c e and e colo g y: Very common in terrestrial habitats (soil, moss)

and in the benthal of running waters and lakes, rarely in waste water treatment plants.
Probably cosmopolitan.
KAHL (1932) gave no exact locality; he discovered "Histrio muscorum" in alpine
mosses from Germany but found it also in mosses from California, USA. I hereby fix a
locality from FOISSNER'S (1982; Fig. 184b) authoritative redescription as locus classicus,
namely "S03", an Alpine pasture at the Hochmaisalm near the GroBglockner Hochalpen-
strafie, Austrian Alps (about 1850 m above sea-level). This site is heavily eutrophified by
cattle manure and thus densely stocked with Rumex sp. and Urtica sp. Locus classicus of
the synonym "Opistotricha terrestris" is Szeged, Hungary, where HORVATH (1956) dis-
covered it in the soil of the garden of the educational college. Locus classicus of the
synonym Oxytricha histrioides is the humus layer of a Hungarian deciduous forest

+- Fig. 184a-g Sterkiella histriomuscorum (a-f, from FOISSNER 1982; g, after DRAGESCO 1970 from DRAGESCO &
DRAGESCo-KERNEIS 1986. a, d, e, from life; b, c, 1; 'g, protargol impregnation). a) Ventral view, 130 11m. b, c)
Ventral and dorsal infraciliature, b, c = 65 11m. d, e) Dorsal and left lateral view, d, e = 140 11m. f) Ventral in-
fraciliature of posterior part, width = 25 11m. g) Ventral infraciliature, 65 11m. The identification is somewhat
uncertain because the undulating membranes are straight and arranged in parallel; reminiscent of Sty/onychia.
CC = caudal cirri, PRVC = pretransverse ventral cirri, TC = transverse cirri; p 683.
 690 SYSTEMATIC SECTION

-<f.:"
~ ~
.~ "1-40-

CC n

Fig.l84h-m Sterkiella histriomuscorum (h, from WENZEL 1953; i,j, from MATIS & DANISKovA 1972; I., from
KAHL 1932; I, m, after MOTE 1954. h, I, m, various staining methods; i-I., from life). Ventral views, h =
140 11m, i, j, I, m = sizes not indicated, k = 100 11m; p 683. n, 0 Sterkiella terricola (from BVITKAMP 1977b).
Ventral infraciliature and nuclear apparatus after protargol impregnation, 120 11m. Tiny arrow marks a very
small cirrus between middle and right frontal cirrus; p 682.
 Sterkiella 691

Table 41 Autecological data of Sterkiella histriomuscorum (columns 1-3), Histriculus histrio (columns 4,5),
and Urosoma caudata (columns 6-8). References: column 1, from DETCHEVA (1972a, 1975a, 1979a, 1983c;
many analyses from Bulgarian running waters); column 2, from FERNANDEZ-LEBORANS (1982; 36 analyses from
the Manzanares River in Madrid, Spain); column 3, from FOISSNER et al. (1991; 10-15 analyses from various
running waters in Austria); column 4, from DETCHEVA (1978, 1983c; many analyses from Bulgarian running
waters); column 5, from PATRICK et a1. (1967; 2 analyses from the Savannah River, USA); column 6, from
RUTHVEN (1972) and CAIRNS & DICKSON (1972; 1 analysis from an American running water); column 7, from
DETCHEVA (1983c; Bulgarian running waters; number of analyses not indicated); Column 8, from REcK (1987;
total range from a eutrophic lake in Germany; number of analyses not indicated)

Parameter' Reference
2 3 4 5 6 7 8
Saprobity a-b a-p b-p b-a a
Frequency (%) 0.3-2.5 66 11 0.6-1 0.6
Temperature eC) 11-21 -0.5-28 0.3-13 13-16 17-20 23 18-22 4-9
pH 7.2-8.3 6.8-9.5 6.6-8.3 6.8-7.4 7.6 7.2-7.4 7.3-7.8
O2 (mgl-') 0.8-10.4 0.8-10.2 3.4-13.3 7.4-8 7-9 3.6-4.5 4.6-7.4 3-7.8
O2 (% saturation) 9-95 31-115 70-80 53-78 23-65
BODs (mg 1-') 3.6-27 1.7-8 2.8-12 0.3-<3 2_3 8.5
NH/-N (mg 1-') 0.03-0.3 0-32 0.01-9 0.06-0.5 <0.0092 0 0.3-0.6 0.06-0.32
NOi-N (mg 1-') o.ot-O.1 0-35 0-0.06 0.02-0.25 0.04 0.04-0.36
NOi-N (mg 1-') 0-1.3 0-4.5 0.4-10.6 0-0.02 0_14-<0.5 0.06 0
pol--p (mg 1-') 0.02-2.8 0.016-0.03 1.9
, The figure "0" denotes only that the parameter in question could not be detected with the analytical method
used.
2 NH,-N.

(GELLERT 1957; no further records are available of this synonym!). Locus classicus of the
synonym Histrio macrostoma is the eastern shore of the Tihany peninsula in Lake Bala-
ton, Hungary, where GELLERT & TAMAs (1958) discovered it in detritus drifts (see also
TAMAS & GELLERT 1959).
Further records substantiated by illustrations: various terrestrial and limnetic habitats
and activated sludge samples from Austria (AUGUSTIN & FOISSNER 1992, AUGUSTIN et al.
1986, 1987, FOISSNER 1987b, C, FOISSNER & FOISSNER 1988, FOISSNER et al. 1990); limnetic
habitat in a cave in Belgium (DELHEZ & CHARDEZ 1970); Bavaria, Germany (WENZEL
1953); beech forest and limed spruce forest in Germany (LEHLE 1989, 1992, 1993); shal-
low rain water pool in Czechoslovakia (MArrs & DANISKOVA 1972); Turiec River in Slo-
vakia (TIRJAKovA 1993); soils in Himalayan region (FOISSNER 1986a); moss-covered soil
on a wall near Seoul and in mosses from Ullung Island, Korea (SHIN & KIM 1994, 1995);
soil of a virgin prairie near Crocker, Iowa, USA (MOTE 1954); soils in Australia (BLAT-
TERER & FOISSNER 1988).
Records not substantiated by illustrations: garden soil in Upper Austria (PETZ et al.
1988); in activated sludge plant (Asten near Linz) in Upper Austria 3-30 d after sam-
pling (OBERSCHMIDLEITNER & AEsCHT 1996); soil in Belgium (CHARDEZ 1967, 1987); soil
692 SYSTEMATIC SECTION

and lotic waters in Bulgaria (DECHEVA 1968, 1970, DETCHEVA 1972b, 1979b, 1981,
TOMEscu 1987); oligosaprobie to betamesosaprobic brooks and beta- to alphamesosapro-
bie rivers in Upper Austria and Bavaria (ForSSNER 1997a, FOISSNER & MOOG 1992, Forss-
NER et al. 1992b); backwash sludge from oxidators and filters of a water works in Ger-
many (FOISSNER 1996a); activated sludge from the sewage treatment plant Marburg-
Cappel, Germany (ETIL 1996); soils in Hungary (GELLERT 1957); mesosaprobic rivers in
Spain (FERNANDEZ-LEBORANS etal. 1990, NIETO et al. 1984); freshwater in Azerbaijan
(ALEKPEROV 1988); various sites of a Donor water reservoir, USSR (ALEKPEROV 1989);
temporary pool (32 % frequency, 23-27 0c) in Cameroon (NJINE 1977); limnetic habitats
in Hunan Province, China (SHEN & GONG 1989); soil in Japan (TAKAHASID & SUHAMA
1991); soil from Venezuela, Brazil, Costa Rica, Australia, and Tasmania (FOISSNER
1997b); soil from Marion Island, South Atlantic Ocean (FOISSNER 1996c); soil from Ant-
arctica (FOISSNER 1996b, SUDZUKlI979).
Feeds on bacteria (Pseudomonas ovalis), algae (green algae, diatoms, Chlorogonium
sp.), fungi, flagellates, amoebas, and ciliates (Colpidium colpoda, Pseudocyrtolophosis
alpestris, Pseudochilodonopsis mutabilis, Tetrahymena sp., Vorticella astyliformis;
ForSSNER 1981, 1982, GIESE 1951, HORvArn 1950, 1956, NIETO et al. 1982, TAMAs &
GELLERT 1958, WENZEL 1953); also cannibalistic (GIESE & ALDEN 1938). Up to 4.47 divi-
sions per day at 26 °C when fed on Colpidium colpoda (GIESE & ALDEN 1938); according
to NIETO et al. (1982), length oflife cycle 6 ± 1 h (± SD?) at 20 °C. Biomass of 106 indi-
viduals about 70 mg (ForSSNER 1987a).
Some autecologieal data of freshwater populations are summarised in Table 41, for
autecological data of terrestrial popUlations, see ForSSNER (1981) and ForSSNER & PEER
(1985). FERNANDEZ-LEBORANS (1982) counted up to 16000 indo I-I in the Manzanares
River in Spain; he found the following optimum values of occurrence: 12.5 °C, pH 7.5,
3.5 mg I-I O2, 0 mg I-I NH/-N, 0.14 mg I-I NOi-N, 0.13 mg I-I NOl--N. ForSSNER et al.
(1991) found S. histriomuscorum once with high abundance in a polysaprobic brook
(saprobie index of total biocoenosis = 3.4) under the following conditions: 8.7°C,
pH 8.1, 540 /lS cm-1 (20°C), 13.8°dH total hardness, 20 mg I-I Cl-, 20 mg I-I SOlo,
9 mg I-I NHt-N, 0.06 mg I-I NOi-N, 0.065 mg I-I NOl--N, 0.16 mg I-I POi--p, 2.2 mg I-I
P.,..., 3.4 mg I-I O2 (31 % saturation), >3.4 mg I-I BOD2, 30 mg I-I KMn04-consumption,
55 mg I-I COD, 184000 bacteria mI-1 (plate method, 22°C), 330000 endotypic coli per
100 ml (plate method, 44 0C). MATIS & STRAKOVA-STRIESKOvA (1991) recorded it from a
brook and two thermal lakes in Czechoslovakia at 9-42°C and pH 5.7-7.2. 100 /lg I-I
lead acetate and 20 /lg I-I zinc chloride were lethal (FERNANDEZ-LEBORANS & ANTONIO-
GARCIA 1986). Further data on the effect of heavy metals such as lead, zinc, and mercury,
see FERNANDEZ-LEBORANS et aI. (1987). Much less sensitive to ultraviolet radiation than
Euplotes patella (GIESE 1938).
FERNANDEZ-LEBORANS (1982) classified S. histriomuscorum as alphamesosaprobic: a;
a = 7, P = 3, 1=4, SI = 3.3. This agrees largely with the experience of ForSSNER et al.
(1991) and the autecological data (Table 41), that is, S. histriomuscorum usually occurs
in organically polluted waters. It is, however, also found in old soil suspensions contain-
 Sterkiella 693

ing many bacteria as well as in betamesosaprobic waters. Thus, FOISSNER et al. (1991)
proposed the following classification: a; b = 2, a = 6, P = 2, I = 3, SI = 3.0. This has to be
checked on reliably determined populations.
In this paragraph some faunistic records and ecological data of limnetic and terres-
trial populations of "Histriculus similis" are listed because these are very likely misiden-
tified Sterkiella histriomuscorum. Faunistic records: inundated meadow in Belgium (BER-
VOETS 1940); submerged, wet, moist, and dry mosses in Czechoslovakia (TIRJAKOVA &
MATIS 1987a); Manzanares River, Spain (FERNANDEZ-LEBORANS & ANTONIO-GARCIA 1988,
FERNANDEZ-LEBORANS & NOVILLO-Vn..LAJOS 1993); mountain lake in Romania (LEPSI
1957); percolating filters and activated sludge in England (CURDS 1975, CURDS & COCK-
BURN 1970a); freshwater in China (YANG 1989); water samples with detritus from the
Mountain Lake Region, Virginia (BOVEE 1960); grassland soils in Canada (BICK & Burr-
KAMP 1976). Ecology: according to CURDs & COCKBURN (1970b), "Histriculus simi/is" in-
dicated an eflluent BODs range of 0-10 mg 1-1 in activated sludge plants. NOLAND (1925)
found it in various types of freshwater in Wisconsin, USA, at 8.5-22.5 °C, 1.5-8.3 mg 1"1
O2 (18-132 % saturation), pH 7.0-8.7. FERNANDEZ-LEBORANS et al. (1985) found 4-8,
about 1.2-2.4 J.1m sized lead particles in specimens treated with 500 J.1g 1-1 lead acetate.

Sterkiella tricirrata (BUITKAMP, 1977) comb. nov. (Fig. 185, Table 40)

1977 Oxytricha tricirrata BUITKAMP, Acta Protozool.,

16: 273.

T a x 0 nom y: The only slightly flexible

body, the arrangement of the frontoventral
cirri, and the distinctly posteriorly displaced

o·
postoral ventral cirrus V/3 indicate that the . ~~il;;::"" ~
species belongs to Sterkiella. It differs from S. ~
terricola, which also has only three transverse
cirri, in that it has only two macronuclear nod-
ules. Sterkiella histriomuscorum has more
adoral membranelles, more marginal cirri (Ta-
ble 40), and more (6) dorsal kineties; never-
theless, synonymy cannot be excluded. Rede-
scription necessary, especially live observa-
tions and morphometric characterisation. ".,',

~:~
-$- ....::. 4;.:;' is;
M 0 r p hoi 0 g y: In life (?) about 80 J.1m
long. Elliptical. Two micronuclei. Contractile
vacuole in about mid-body. Undulating mem- Fig. 185 Sterkiella tricirrata (from BUITKAMP
branes each consist of a single row of basal 1977b). Ventral infraciliature and nuclear appa-
ratus after protargo\ impregnation, 80 J.Im.
694 SYSTEMATIC SECTION

bodies. Arrangement of frontal, ventral, and transverse cirri as in some other Sterkiella
species. Invariably (n = ?) only 3 transverse cirri. Cirri in life about 10 J.1m long. Dorsal
cilia 2 J.1m long, arranged in 5 kineties, however, it cannot be excluded that BUITKAMP
(1977b) overlooked the short kinety 6.

o c cur r e n c e and e colo g y: Locus classicus is the soil of a burnt savannah in

the Ivory Coast; not found since. Feeds on small amoebas and ciliates. Biomass of 106 in-
dividuals about 55 mg (FOIsSNER 1987a), according to BUITKAMP (1979) only 24 mg.
BUITKAMP (1979) found 115 and 29 indo g-' dry soil in cultures incubated at 20°C and
25 °C respectively.

Sterkiella quadrinucleatus (SICK, 1933) comb. nov. (Fig. 186a~, g)

1932 His/rio similis QUENNERSTEDT, 1867 - KAHL, Tierwelt Dtl., 25.: 615; in part, Fig_ 116,2 (Fig. 186b) ; not
Fig. 121,6.
1933 Histrio similis QUENNERSTEDT 1867 - KAHL, Tierwelt N.-u. Ostsee., 23 (Teil IT.C3): 114.
1933 Histrio quadrinuc/eatus KAHL n. sp. - SICK, Z. wiss. Zool., 2: 66 (see taxonomy).
1948 Oxytricha (His/rio) similis. QUENN. - MARGALEF, Publnes Inst BioI. apI., Barcelona, 5: 70 (Fig. 186g).
1961 Histrio simi/is QUENNERSTEDT, 1867 - REUTER, Acta zool. fenn., 99: 20 (Fig. 186a).
1972 Histriculus similis (QUENNERSTEDT, 1867) - BORROR, Acta Protozool., 10: 64 (Fig. 186c).

Nom e n cia t u rea n d t a x 0 nom y: KAHL (1932) classified Histrio as subgenus

of Oxytricha; the correct name in his revision is thus Oxytricha (Histrio) simi/is, which is
a secondary homonym of Oxytricha (Opisthotricha) simi/is in KAHL (1932).
Histrieu/us similis was originally described with two macronUclear nodules (Fig.
178t). KAHL (1932, Fig. 186b) designated a very similar, but quadrinucleate species as H.
similis, assuming a misobservation by QUENNERSTEDT (1867). Recognising this shortcom-
ing, SICK (1933) established a new species for the KAHL (1932) population, namely His-
trio quadrinucieatus. It is probably confined to saline habitats. Thus, all records of "His-
trieu/us similis" or "Histrio simi/is" from marine or inland saltwater which were identi-
fied according to KAHL (1932) are adjudged Sterkiella quadrinucleatus.
The BORROR (1972b; Fig. 186c) population agrees rather well with REuTER'S (1961;
Fig. 186a) population, especially in body shape and in that they have an increased num-
ber of postoral ventral cirri and 6 transverse cirri. However, since KAHL (1932) men-
tioned that his populations sometimes also had 6-8 transverse cirri, I preliminarily avoid
a further splitting. The REUTER (1961) and BORROR (1972b) populations are reminiscent
of Gastrosty/a stein;; (rather invariably 4 transverse cirri) and Territricha stramentico/a
(2 macronuclear nodules, lives in beech litter). Detailed redescription necessary.

M 0 r ph 0 log y and b i 0 log y: According to KAHL (1932) in life 150-250 J.1m,

after MARGALEF (1948) 125-150 J.1m, after REUTER (1961) 110-130 J.1m long, after BOR-
ROR (1972b) 83-150 x 40-59 J.1m. Body margins parallel or slightly converging posteri-
 Sterkiella 695

Fig. 186a-c, g Sterkiella quadrinuc!eatus (a, after RElJfER 1961; b, from KAm. 1932; c, from BORROR 1972b;
g, from MARGALEF 1948. a, b, g, from life; c, from life and after nigrosin-mercuric- fonnalin method). a, b, g)
Ventral views, a = 110-130 11m, b = 150 11m, g = 125-150 11m. The small figure shows a lateral view. c) Ven-
tral infraciliature of a specimen with supernumerary cirri, 90 11m; p 694. d Oxytricha durhamiensis (from
BORROR 1972b). Ventral view from life and after nigrosin-mercuric-fonnalin method, 90 11m; p 228. e, r Sterk-
iella histriomuscorum (e, from DELHEZ & CHARDEZ 1970; f, from GIESE & AwEN 1938. e, f, from life). e, f)
Ventral views, e = 80 11m, f= 105 11m. Nuclear apparatus in (1) stained with FEULGEN reaction; p 683.
696 SYSTEMATIC SECTION

orly. Contractile vacuole in about mid-body (Fig. 186b), according to BORROR (1972b)
nrnnerous water expulsion vesicles aborally, in a series slightly left of median, indicating
that this population is possibly a species of its own. BORROR (1972b) also mentioned
small cortical granules.
Adoral zone of membranelles about 40 % of body length, about 40 membrane lies
(BORROR 1972b). Frontal and transverse cirri distinctly enlarged. Nrnnber and arrange-
ment of cirri according to Figure 186b more or less Sterkiella specific; according to
REUTER (1961) and BORROR (1972b) 2 or 3 supernrnnerary postoral ventral cirri and an
additional transverse cirrus present. However, KAlIL (1932) also mentioned that the nrnn-
ber of transverse cirri is occasionally 6-8. At least 4 dorsal kineties (BORROR 1972b).

o c cur r e n c e and e colo g y: KAlIL (1932, 1933) found Sterkiella quadrinuclea-

IUs in the Northern Sea near Sylt, Germany (locus classicus) and also in the Bay ofKiel,
Eastern Sea, and in saline sites in Oldesloe near Hamburg, Germany. MARGALEF (1948)
observed it in the littoral of a brackish pond near Barcelona, Spain. REUTER (1961, 1963a)
recorded it from Scandinavian rock pools with a salinity of 0.43-1.09 %. BORROR
(l972b) collected S. quadrinucleatus from New Hampshire tidal marshes, USA.
Records not substantiated by illustrations: brackish water in the Bay of KieI, Ger-
many (SICK 1933); sub-littoral sediments in the Eastern Sea (BOCK 1952, HARTWIG 1974);
rarely in the aufwuchs community of a heavily polluted site in a harbour (Klinigshafen)
near List, Sylt, Germany (KOSTERS 1974); regularly in ground water (0.54--0.79 %
salinity) up to the 8 m isotele of the coast of Hiddensee Island, Eastern Sea (MUNCH
1956); saline lake in Romania (TUCULESCO 1962b). STELLA (1947) found "Histrio similis"
in a spring in Italy (misidentification?).
Feeds on green algae and diatoms (BORROR 1972b). MrrwLOWITSCH (1989) recorded
it from small, saline running waters in Germany under the following conditions (n = 7):
12.3-17.8 °C, pH 7.4-7.7, 12-35 mg I-I CO2 , 2.5-9.8 mg 1-1 0 2,0.05-0.39 mg I-I NH/-N
(n = 6),0.04--0.1 mg I-I N02--N (n = 5), 1.7-6.9 mg I-I NOJ--N (n = 6),36 to 410 mg I-I
Cl-, and 418-1943 IlS cm-I specific conductivity. KAlIL (1932) classified Sterkiella quad-
rinucleatus as meso- to polysaprobic; however, this has to be verified by further aute-
coiogical studies on reliably determined populations.

Parastylonychia DRAGESCO, 1963

1963 Parasty/onychia DRAGESCO, Cah. BioI. mar., 4: 263 - Type (original designation): Parastylonychia
plouneouri DRAGESCO, 1963.
1972 Parastylonychia DRAGESCO, 1963 - BORROR, J. Protozool., 19: 14.
1985 Parastylonychia - SMALL & LYNN, Phylum Ciliophora, p. 460.
1997 Parastylonychia DRAGESCO, 1963 - BERGER & FOISSNER, Arch. Protistenk., 148: 149.
 Parastylonychia 697

C h a r act e r i sat ion: Undulating mem-

branes in Stylonychia pattern, that is, more or
less straight and parallel. One right and 1 left row
of marginal cirri, distinctly separate posteriorly.
Caudal cirri absent.

T a x 0 nom y: The characterisation is accord-

ing to BERGER & FOISSNER (1997). Description of
Stylonychia pattern, see chapter 2 in the general
section. Monotypic; for details, see Parastylony-
chia plouneouri.

Single species

Parastylonychia plouneouri DRAGESCO,

1963 (Fig. 187)

1963 Parastylonychia plouneouri DRAGESCO, Cah. BioI.

mar., 4: 263.
1972 Parastylonychia p/euneouri DRAGESCO, 1963 - BOR-
ROR, J. Protozool., 19: 14 (incorrect subsequent spell-
ing).

T a x 0 nom y: The fum body indicates that it is

a Stylonychinae. However, a detailed redescrip-
tion and morphogenetic data are needed for a
more exact classification of this marine species. Fig. 187 Parastylonychia plouneouri (from
Reminiscent of "Stylonychia grandis", which, DRAGESCO 1963). Ventral view in life and
after protargo\ impregnation ("aspect gene-
however, has caudal cirri (Fig. 1810).
ral schematique"), 200 Ilm.

M 0 r p hoi 0 g y: In life about 200 !lm long,

approximately 2.5 times as long as broad. Ellip-
tical to parallel-sided, both ends broadly rounded. Body frrm. Four macronuclear nod-
ules, 1 or 2 micronuclei. Contractile vacuole not mentioned. Adoral zone of mem-
branelles about 37 % of body length. Three enlarged frontal cirri, 5-6 frontoventral cirri
(including buccal cirrus), 4-5 postoral ventral cirri, 2-3 pretransverse ventral cirri, and
5 distinctly enlarged transverse cirri, which protrude beyond posterior end of cell. 25-38
right and 16-20 left marginal cirri (according to the illustration, however, 31 left mar-
ginal cirri!). Four dorsal kineties, dorsal cilia very likely short, because neither men-
tioned nor drawn.
698 SYSTEMATIC SECTION

o c cur r e n c e: Locus classicus is the Atlantic Ocean near the town of Roscoff, France,
where DRAGESCO (1963) discovered Parastylonychia plouneouri in the saprobic psammal
at "de Plouneour-Trez". Not found since.

Pleurotricha STEIN, 1859

1859 Pleurotricha STEIN, Lotos, 9: 4 - Type (original designation): Stylonychia Ianceolala EHRENBERG, 1835.
1859 Pleurotricha. STEIN - STEIN, Organismus der Infusionsthiere I, p 168.
1932 Pleurotricha STEIN, 1859 - KAHL, TierweltDtl., 25: 593.
1972 Pleurotricha STEIN, 1859 - BORROR, J. Protozool., 19: 14 (see nomenclature and taxonomy).
1974 Pleurotricha lanceolala STEIN - STILLER, Fauna Hung., ll5: 105 (in part).
1979 Pleurotricha s. str., subgen. n. - JANKOWSKI, Trudy zool. Inst., Leningr., 86: 62 - Type (original desig-
nation): Pleurotricha grandis STEIN, 1859 (see nomenclature and taxonomy).
1997 Pleurotricha STEIN, 1859 - BERGER & FOISSNER, Arch. Protistenk., 148: 149.

C h a r act e r i sat ion: Undulating membranes in Oxytricha pattern. Two or more

right and one or more left rows of marginal cirri. Caudal cirri absent. No parental mar-
ginal cirri retained after division. Dorsal morphogenesis in Oxylricha pattern.

Add i t ion a I c h a r act e r s: Large to vel)' large. Body fInn. Two macronuclear
nodules. Contractile vacuole at about level of cytostome. Cortical granules lacking. Cyto-
plasm colourless, however, after STEIN (1859b) the cytoplasm of P. lanceolala is usually
pale and bluish-grey. Adoral zone of membranelies 40-45 % of body length, buccal area
large. 18 frontal-ventral-transverse cirral pattern. Left and middle frontal cirrus arranged
at about same level, right one displaced distinctly posteriad Postoral ventral cirrus V/3
near anterior pretransverse ventral cirrus (V/2). Transverse cirri arranged in 2 distinctly
separated groups with 3 left (cirri 1111, lIllI, IV/l) and 2 right (cirri VII, VIII) cirri re-
spectively; only cirri of right group protrude distinctly beyond posterior end of cell. Fron-
tal, pretransverse ventral, and transverse cirri and buccal cirrus distinctly enlarged. Outer
marginal rows almost confluent posteriorly. Dorsal cilia 2-4 Ilm long. Freshwater, lentic
waters obviously preferred.

Nom e n c I at u rea n d t a x 0 nom y: STEIN (1859a) established Pleurotricha une-

quivocally for Stylonychia lanceolata and not for Pleurotricha grandis, as erroneously
assumed by BORROR (1972a) and JANKOWSKI (1979). The latter author established the sub-
genera Pleurotricha (Pleurotricha) (see above) and Pleurotricha (Allotricha) (JANKOWSKI
1979, P 50,62). However, the type species were wrongly fIxed (Icm 1985, Article 44a).
Furthennore, Pleurolricha (Allotricha) was already classifIed as subgenus, albeit with
some doubt, by BOTsCHLI (1889). Incorrect subsequent spellings: Pleurosticha (HYMAN
1931, P 56); Pleurtricha (BUCHHOLZ & BUCHHOLZ 1974, P 251).
The characterisation is according to BERGER & FOISSNER (1997). For a detailed de-
scription of patterns mentioned, see chapter 2 in the general section. Pleurotricha has the
typical 18 frontal-ventral-transverse cirral pattern of the oxytrichids, but additional mar-
 Pleurotricha 699

ginal rows and no caudal cirri (the latter character, however, is not substantiated by mor-
phogenetic data). The lack of caudal cirri (and the almost confluent outer marginal rows)
indicate that it is possibly the sister-group of Histriculus. However, detailed morphoge-
netic studies are needed for clarifying the cladistic relationships. Laurentiel/a, Onycho-
dromus, and Coniculostomum, all of which usually also have an increased number of cir-
ral rows, differ (among other things) in that they have prominent caudal cirri. Coniculo-
stomum has, furthermore, a different mode of marginal row formation, namely parental
cirri are retained after division. Allotricha STERKI, 1878 was often considered to be ajun-
ior synonym of Pleurotricha (for example, BERGER & FOISSNER 1997, BORROR 1972a,
KAHL 1932). However, it has a flexible body and cortical granules and thus belongs to the
Oxytrichinae.
Pleurotricha includes two valid species, namely P. lanceolata and P. grandis. Pleu-
rotricha planensis FERNANDEZ-LEBORANS, 1984 is a species indeterminata. Pleurotricha
indica AMMERMANN, SAPRA and SCHLEGEL, 1987 is classified as ConiculostorilUm because
it has caudal cirri. Pleurotricha variabilis REurnR, 1961 is flexible and thus classified in
Allotricha. Pleurotricha setifera ENGELMANN, 1862 belongs to Gastrostyla. Pleurotricha
macrostoma DRAGESCO, 1970, was transferred to Holosticha by BORROR (1972a).

Key to species
1. 1 left marginal row (for example, Fig. 188t) .............. P. lanceolata (p 699)
More than 1, usually 3 left marginal rows (Fig. 190a, b) ........ P. grandis (p 712)

Pleurotricha lanceolata (EHRENBERG, 1835) STEIN, 1859 (Fig. 188a-z,

189a-x, Tables 27, 42)
1835 Stylonychia lanceolala EHRENBERG, Abh. preuss. Akad. Wiss., year 1835: 164 (no illustration).
1838 Stylonychia lanceolata - EHRENBERG, lnfusionsthierchen, p 373 (Fig. 188a-d).
1841 Kerona lanceolata - DUJARDIN, Zoophytes, p 427.
1852 Cerona lanceolata - PERlY, Zur Kenntniss Ideinster Lebensforrnen, piSS.
1859 Pleurotricha lanceolata - STEIN, Lotos, 9: 4 (combining author).
1859 Pleurotricha lanceolata. STEIN - STEIN, Organismus der lnfusionsthiere I, p 170 (Fig. 188e-g).
1862 Pleurotricha lanceolata STEIN - ENGELMANN, Z. wiss. Zool., 11: 364 (Fig. 188h, i, 0).
1882 Pleurotricha lanceolata, EHR. sp. - KENT, Manual infusoria II, p 783.
1898 Pleurotricha lanceolata (EHBG). - JOUKOWSKY, Verh. naturh.-med. Ver. Heidelb., 6: 21 (Fig. 188m).
1925 Pleurotricha grandis STEIN - GULATI, J. Bombay nat. Hist. Soc., 30: 752 (Fig. 1891; misidentification).
1928 Pleurotricha lanceolata- MANwELL, BioI. Bull. mar. bioI. Lab., 54: 417 (Fig. 1881, 189n).
1928 Pleurotricha lanceolata (STEIN) - MANwELL, BioI. Bull. mar. bioI. Lab., 55: 433 (Fig. 188k).
1932 Pleurotricha lanceolala (EHRB., 1838) - KAHL, Tierwelt Dtl., 25: 593 (incorrect dating).
1932 Pleurotricha spec. - KAHL, Tierwelt Dtl., 25: 593,601 (Fig. 188n).
1933 Pleurotricha lanceolala (EHRENBERG) STEIN 1859 - WANG & NIE, Contr. bioI. Lab. Sci. Soc. China, 10:
60 (Fig. 189k).
1942 Pleurotricha lanceolala - LILLY, Physiol. Zool., 15: 147 (Fig. 188j).
1963 Pleurotricha lanceolala (EHRENBERG) - LUNDIN & WEST, Free-living protozoa, p 68 (Fig. 189m).
700 SYSTEMA TIC SECTION

Fig. 188a--d: Pleurotricha lanceolata (after EHRENBERG 1838. a-d, from life). a) Ventral view, 170-210 ~m.
b) "Dissolving" specimen. c) Lateral view. d) Dorsal view; p 699.

1968 Pleurotricha lanceolata-1EFFRlES & MELLOTI, 1. Protozool., 15: 741 (Fig. 188s-z, 189a--t).
1970 Pleurotricha Ianceolata (EHRENBERG, 1838) - DRAGESCO, AnnIs Fac. Sci. Univ. fed. Cameroun
(Numero Hors-serle): 125 (Fig. 188p-r).
1972 Pleurotricha tchadensis DRAGESCO, AnnIs Fac. Sci. Univ. fed. Cameroun, 11: 85 (Fig. 189g; new syno-
nym).
1979 Pleurotricha (Allotricha) lanceolata-1ANKwosKl, Trudy zool.lnst., Leningr., 86: 62 (erroneously fixed
as type of subgenus, see nomenclature of genus).
1980 Pleurotricha Ianceolata - GRIMES, McKENNA, GoLDSMllli-SPOEGLER & KNAUPP, Science, 209: 281, Fig.
la (scanning electron micrograph).
1984 Pleurotricha lanceolata - MARTlN-GoNzALEZ, SERRANO & FERNANoEZ-GALIANO, J. Protozool., 31: 347
(Fig. 189i, j).
1986 Pleurotricha Ianceolata (EHRENBERG, 1838) STEIN, 1859 - DRAGESCO & DRAGESco-KERNEIs, Faune
tropicale, 26: 476 (Fig. 189h).
1986 Pleurotricha tchadensis DRAGESCO, 1972 - DRAGESCO & DRAGESCO-KERNliIS, Faune tropicale, 26: 477.

Nom e n c I at u rea n d t a x 0 nom y: Incorrect subsequent spellings: Pleurotri-

cha lanccolala STEIN (GURWITSCH 1934, P 17); Pleurotricha lanceolalum (KAy 1945b,
P 272; KAy 1946, P 133); Pleurotricha lchadiensis (DRAGESCO 1972b, P 71).
Pleurotricha lanceolala is insufficiently described by EHRENBERG (1835, 1838) so
that the identification by STEIN (1859a, b) is somewhat uncertain. STEIN (1859b) even sup-
posed the insufficiently known Trichoda calvilium MOLLER, 1773 (= Kerona calvitium in
 Pleurolricha 701

Fig. 188e--g: Pleurotricha lanceolata (after STEIN 1859b). Ventral views from life, 170-297 11m. STEIN, one of
the most outstanding protozoologists, recognized almost every detail of the infraciliature, including the lack of
caudal cirri; p 699.

MOLLER 1786) to be identical with the present species. Pleurotricha spec. sensu KAHL
(1932) is vel)' likely also P. lanceolala because it has the same ventral infraciliature and
size; a third right marginal row is not as unusual as supposed by KAHL (1932; see Fig.
188e, n, z).
Pleurotricha tchadensis differs slightly from P. lanceolala in the arrangement and
number of frontoventral and postoral ventral cirri, which is vel)' likely, at least to some
extent, due to inexact observations on the African population. Furthermore, DRAGESCO
(1970) drew 7 dorsal kineties, which is probably a further misobservation; he obviously
confused the outer right marginal row with a dorsal kinety (Fig. 188r). I thus synonymise
P. tchadensis with the type, as already proposed by HEMBERGER (1982, P 222). However,
further African populations should be studied.
 702 SYSTEMATIC SECTION

Fig. 188h--o: Pleurolricha lanceolala (h, i, 0, after ENGELMANN 1862;j, from LILLY 1942; k, I, from MANwELL
1928a, b; m, after JOUKOWSKY 1898; n, from KAlIL 1932. h-j, 1tl--Q, from life; k, I, fixation with SCHAUDINN'S
fixative and haematoxylin staining). h) Conjugation, 260 11m. i) Exconjugant with macronucleus anlage (ar-
rowhead), 220 11m. j) Ventral view, 145 11m. k, I) Ventral infraciliature as seen from dorsal, 127 11m. m, 0)
Ventral views, m = size not indicated, n = 220 11m. 0) Resting cyst with its distinct spines, 100 11m; p 699.
Pleurotricha 703

...
. ' .1

..

·'.0
•t
•
•

q
11\\\"'"

f
/IIIf
\
, '. -:".
\ .......... >. .
...
: 00. '0 .....
0

.....
•

. .
...
': "0• • • • •

..
'0 "0 :

..:

.... .. "

Fig. 188p-r: Pleurotricha lanceolala (from DRAGESCO

1970. p-r, protargol impregnation). p, q) Ventral infra-
ciliature, p, q = 150 j.lm. r) Dorsal infraciliature, 150 j.lm.
The long dorsal kinety on the left margin (arrowhead) is
very likely a misinterpreted right marginal row; p 699.
 704 SYSTEMATIC SECTION

,, ,
,", ,
, I
,, ,I
.
, , I

,, ,' ,
,,'".
' I
,, , , I
'"
•• , , , ••
; I

,,
, I I ,'I

,, , I
, I I
I ,
I, "
•t .
I'
I I
I I

:
,.
I , ',', I
,I I' ,
,I I I
I,','.
I

", ,
I I

•,
~

I
I :
I
,f,.
, :
:-
'. "~I ....

Fig. 188s-z, 189a, b: Pleurotricha lanceolata (from JEFFRIES & MELLOlT 1968. s-z, a, b, protargol impregna-
tion). s-z, a, b) Variability of ventral infraciliature (see also Fig. 1890-f). However, note that the number and
arrangement of the 18 frontal-ventral-transverse cirri is usually rather invariable; p 699.

Pleurotricha lanceolata sensu RAo (1928) is classified as a species of the Stylony-

chia mytilus complex because prominent caudal cirri are present. Furthermore, RAo
(1928) possibly misinterpreted the dorsal cilia as outer right marginal row (Fig. 1510).
The CIENKOWSKY (1855; Fig. 222m, n) and EDMONDSON (1906; Fig. 234k) populations
are insufficiently redescribed. Pleurotricha lanceolata sensu SMITH (1978, Fig. 92k) is
classified as Allotricha antarctica. Allotricha mollis STERK! is a valid species belonging to
the Oxytrichinae; it is thus excluded from the synonymy of P. lanceolata (BORROR
1972a).

M 0 r p hoi 0 g y and b i 0 log y: Size obviously rather variable, in life 100-300

times 50-120 J.lm: 173-208 J.lm (EHRENBERG 1838), 170-297 J.lm (STEIN 1859b), 100 to
165 J.lm (MANwELL 1928a), 170-300 J.lm and 200-250 J.lm (Fig. 188n; KAHL 1932),
225-235 x 88-94 J.lm (WANn & NIE 1933), 128-200 x 52-84 J.lm (JEFFRIES & MELLOTI
 Pleurotricha 705

i
.
: ~:.
.":CV'.:
/ ~ 1
, / \
/ ~ f\
/ / ,.
, /,::..
/ I , .., ; /
, ~.
I ./

I ~
I I
I ~

.
I ,
II " ,

,, I

,"
~

" 111_:
I I .:
' .. ' e

Fig. 189c-f: Pleurotricha lanceolata (c, d, from JEFFRIES & MELLOTT 1968; e, 1: after JEFFRIES & MELwTI
1968. c-f, protargol impregnation). c, d) Variability of ventral infraciliature (cp. Fig. 188s-z, 189a, b). e, f)
Ventral and dorsal infraciliature of representative specimens, sizes not indicated. CV = position of contractile
vacuole, RMRl, RMR2 = right marginal row 1 and 2, respectively; p 699.

1968), 200-250 x 100-120 Jim (MARriN-GoNzALEz et al. 1984). Elliptical to elongate

lancet-shaped, anterior end rounded or obliquely truncated, posterior usually narrowly
rounded, sometimes even slightly tapered. Body margins slightly to distinctly converging
posteriorly; left margin almost straight or convex, right usually much more convex. Mac-
ronuclear nodules after protargol impregnation about 15-29 11m long. 2-3 micronuclei,
about 2.5 11m across. Contractile vacuole opens dorsally between kineties 1 and 2, cy-
topyge between dorsal kineties 2 and 3 (JEFFRIES & MEllOTT 1968). Rapid movement,
rather similar to in Histriculus histrio (STEIN 1859b).
Both undulating membranes consist of zigzagging basal bodies. Arrangement of
frontal, ventral, and transverse cirri rather invariable (cp. Fig. 188f, i, n, 18ge, i) and al-
most identical to in Steinia sphagnicola (cp. Fig. 18ge and Fig. 173c). Bases of frontal,
ventral, and transverse cirri composed of rather differing numbers of basal bodies (Fig.
189j). Right marginal row 1 (= outer one) begins at level of distal end of adoral zone of
membranelles, terminates at posterior end of cell. Right marginal row 2 of rather varying
length, but always present. Row 3, if present at all, usually distinctly shortened anteriorly
and posteriorly. DRAGESCO (1972b) popUlation sometimes with a fourth right marginal
row with Up to 3 cirri (Fig. 189g). Cirri in all marginal rows about equidistant; an impor-
 706 SYSTEMATIC SECTION

.? " I f !II!fIg
;3'", 4~
•
• t

,;
II
;
;

; ;

,
,,
;

.;
; .;

I
#
i
I ,
t #
I ; ,.

. ••
, I
'f I
"
, I

••

Fig. 189g, h: Pleurotricha lanceolata (g, from DRAGESCO 1972b; h, from DRAGESCO & DRAGESCO-KERNEIS
1986). Ventral infraciliature after protargol impregnation, g = 180 )un, h = 125 11m; p 699.

tant difference to Coniculostomum monilata, where the cirri of the inner rows are dis-
tinctly more widely spaced than in the outer row. This is due to the different morphogene-
sis in Pleurotricha and Coniculostomum. Invariably 6 dorsal kineties, kineties 1-4 about
of body length, dorsomarginal rows (= kineties 5 and 6) only 113 of body length (Fig.
189f, Table 42). For details on the fibrillar system, see MARTiN-GONzALEz et al. (1984).
Resting cyst spherical; ectocyst with prominent, about 6 J.Lm long, spine-like projec-
tions (Fig. 1880, 189n-u, x). Cysts average 56 J.Lm in diameter (without spines; JEFFRIES
1956; Fig. 189p-x); however, after ENGELMANN (1862) up to 100 J.Lm across. PENN (1935)
investigated the effect of various factors (food, temperature, acidity, alkalinity, age of
hay-rye infusion, excretion products of different organisms, bacteria, crowding, urine,
ammonia, salts, extraction of water from the organism) on the rate of encystment. This
part of the life cycle was studied by MANwELL (1928a), who stated that the old macronu-
 Pleurotricha 707

-f'
\

. : ~:~: : .
·::tX7 ..::::t::;:::-
":::\\", \%2
.,.... .':". !·;. · .:· .~ :.· :. :. : .· :.· : .·:V ~4·
'1fG:3
····:::::::::·.::·.:.·.:::.V:·::·5

.0::::'
...•:. .::::. ,.:,:.:••:.0:.' .0::::
•••••• .....::::.
••::.:: 00

.:.:.:.:f:::o "0° 0::.::.°::::

0 0 :."

RMC LMC

Q
Q

Fig. 189i, j: Pleurotricha lanceolata (~ after MARTIN-GONzALEz et aI. 1984; j, from MARTIN-GoNzALEz et aI.
1984. i, j, silver carbonate impregnation). i) Ventral infraciliature, size not indicated. j) Diagram of different
cirri showing number and arrangement of basal bodies. FI = cirrus III, F2 = lI/3, F3 = III/3, F4 = II/2, FS =
VIl4, F6 = VI/3, F7 = lII/2, F8 = lV/4, LMC = left marginal cirrus, RMC = right marginal cirrus, Tl = 1II1, T2
= III/1, T3 = lVII, T4 = VII, T5 = VIII, VI = lVl2, V2 = V/4, V3 = V/3, V4 = VI2, V5 = VII2 (see Fig. 6a for
designation of cirri); p 699.

cleus and apparently 1 micronucleus is extruded during encystment. However, this could
not be confIrmed by JEFFRIES (l956), who studied mainly excystment (Fig. 189p-x). He
found that the optimal rate of excystment is obtained at 20-23 °C. The time required for
the completion of the excystment process averages 5-8 hoUTS. Citric acid is a very good
excysting agent (JEFFRIES 1959, 1962).
Morphogenesis of cell division and conjugation were studied by ENGELMANN (1862)
and MANwELL (l928a, b), however, only live and hematoxylin-stained material was used.
According to MANwELL (1928a) morphogenesis commences with the formation of an oral
primordium in about mid-body, and not adjacent to the left transverse cirrus as is the case
in many other Stylonychinae. However, this must be confirmed on protargol-impregnated
material. The inner right marginal row develops from part of a streak in the outer parental
right marginal row (BORROR & WICKLOW 1982). Regeneration and dorsi-conjugant forma-
tion were studied by HEwm (l914) and TCHANG & PANG (1979a).
708 SYSTEMATIC SECTION
 Pleurotricha 709

Mirror-imaged doublets can be obtained either by heat-shocking (41 cC, 17 min) ran-
dom cultures of cells in the "log phase" and sub-cloning or by sub-cloning surgically in-
duced longitudinal fragments of cells in the "log phase" of growth (GRIMES et al. 1980).

o c cur r e n c e and e colo g y: According to the literature common in detritus and

aufwuchs of freshwater; much more common than Pleurotricha grandis. I have never
seen this species and there is also no recent record from Austria (FOISSNER & FOISSNER
1988). Probably cosmopolitan.
Locus classicus of Pleurotricha lanceolata is a freshwater habitat near Berlin, Ger-
many, where EHRENBERG (1835, 1838) discovered it among algae and cyanobacteria. Lo-
cus classicus of the synonym, P. tchadensis, is a pond in a reservation in Chad (DRAGESCO
1972b; see also DRAGESCO 1980). STEIN (1859a, b) foundP. lanceolata together with P.
grandis in the locus classicus of the latter species (!;lee there), in a fish pond covered with
Lemna sp. near Tharand (Germany), and, like EHRENBERG, highly abundantly between
tufts of Oscillatoria and Spirogyra in a brook near the village ofNiemegk, Czechoslova-
kia. Further records substantiated by illustrations: environs of Leipzig, Germany (ENGEL-
MANN 1862); Madrid, Spain (MARTiN-GONzALEZ et al. 1984); abundantly among decaying
organic matter and aquatic plants in Lake Ho Hu, Nanking, China, during September
(WANG & NIE 1933); infusion of dry leaves, Lahore, India (GULAn 1925); culture of
leaves and swamp water from Orient Springs near Amherst, Massachusetts, USA (MAN-
WEll 1928a, b); freshwater near Providence, Rhode Island, USA (LIlLY 1942); Upper
Peninsula of Lake Michigan, USA (LUNDIN & WEST 1963, WEST 1953); stream near Car-
lisle and spring-fed stream on the campus of the University of North Carolina, USA (JEF-
FRIES 1956, JEFFRIES & MaLoTI 1968); Cameroon (DRAGESCO 1970).
Records from freshwater habitats not substantiated by illustrations: Vienna and Up-
per Austria and font of the cemetery in Innsbruck, Austria (DALLA TORRE 1891, RIEss
1840); Belgium (BERVOETS 1940); Denmark (NIELSEN 1960); near Perth, England
(CRAIGIE 1921); Ferrara and Sardinia, Italy (CANELLA 1954, PARONA 1883); Madrid and
Barcelona, Spain (FERNANDEZ-LEBORANS & ANTONIO-GARCiA 1988, MARGALEF LOPEZ
1945); Bern and near Basle, Switzerland (PERTY 1852, RIGGENBACH 1922); saline lakes
and various freshwater habitats in the USSR (EICHWALD 1844, GURWITSCH 1934, WEISSE
1848a, b); Palestine (BODENHEIMER 1937); at pH 3.0 in surface waters polluted by acid

~ Fig. 189k-x: Pleurotricha lanceolata (k, from WANG & NIE 1933; 1, after Gui.All 1925; m, from LUNDIN &
WEST 1963; n, from MANwELL 1928a; 0, after lLoWAlSKY 1926; p-x, from JEFFRIES 1956. k-m, o-x, from life; n,
haematoxylin stain). k-m) Ventral views, k = 240 !lm, I, m = size not indicated. n) A cyst containing 7 micro-
nuclei, several of which appear to be in mitosis, 80 !lm. 0) Cyst, diameter without spines about 70 !lm. p-x)
Successive stages of excystment. p, q) Resting cyst (P) and cyst after 50 min in excystment medium (q), con-
tractile vacuole and auxiliary fluid vacuoles present. r) Cyst inunediately after one of the early systoles of the
contractile vacuole. s) Organism now free in cyst membranes. t) Rupture of ectocyst, exposed endocyst with
enclosed organism. u) Endocyst and enclosed organism escaping ectocyst. v) Folded organism free in
endocyst. w) Trophozoite almost fully differentiated, macronuclear nodules and micronuclei discernible. x)
Empty ectocyst. CV = contractile vacuole, EC = ectocyst, Ma = macronucleus, SB = storage body; p 699.
710 SYSTEMATIC SECTION

Table 42 Morphometric data of Pleurotricha lanceolata (lines I, 2c, 2m, clone 1-20, clone II-8, and mixed
population II from JEFFRIES & MELWT 1968; lines 3, 4, population from Cameroon and Benin respectively,
from DRAGESCO & DRAGESCO-KERNEIS 1986; line 5, from DRAGESCO 1972b). All data from protargol-
impregnated specimens. All measurements in micrometres. ? = sample size not indicated; if only I value is
known it is listed in the mean column, if 2 values are available they are listed as Min and Max. CV = coeffi-
cient of variation (in %), Max = maximum value, mean = arithmetic mean, Min = minimum value, n = sample
size, SD = standard deviation

Character mean SD CV Min Max n

Body, length 12 105.1 90 120 >50
2c2 106.3 80 136 >50
31 140 165 ?
41 115 135 ?
51 130 185 ?
Body, width 12 51.5 40 66 >50
2c2 43.9 32 60 >50
Adoral membralielles, number 33 53.0 50 61 10
5 54 79 ?
Right marginal row 1, number 30.7 24 37 >50
of cirri 2c 30.8 15 36 >50
2m 33.4 19 42 >50
3 30 35 9
4 30 40 ?
5 33 37 ?
Right marginal row 2, number 1 14.9 8 23 >50
of cirri 2c 14.6 7 20 >50
2m 17.5 9 25 >50
3 19 25 ?
4 20 27 ?
5 25 30 ?
Right marginal row 3, number 3 2 12 ?
of cirri 5 12 21 ?
Left marginal row, number 25.1 18 39 >50
of cirri 2c 25.8 12 32 >50
2m 26.9 20 35 >50
5 21 27 ?
Basal body pairs in dorsal 1 28.5 22 36 >50
kinety I, number 2c 28.3 21 34 >50
2m 34.7 23 41 >50
Basal body pairs in dorsal 1 25.6 22 31 >50
kinety 2, number 2c 26.0 19 32 >50
2m 31.5 23 38 >50
Basal body pairs in dorsal 1 25.3 21 31 >50
kinety 3, number 2c 25.8 22 31 >50
2m 30.3 23 39 >50
Basal body pairs in dorsal 1 22.2 16 27 >50
kinety 4, number 2c 22.8 18 29 >50
2m 27.5 20 38 >50
Basal body pairs in dorsal 1 10.9 7 15 >50
kinety 5, number 2c 11.6 8 17 >50
2m 13.7 8 19 >50
 PIeurotricha 711

Table 42 Continued

Character mean SD CV Min Max n

Basal body pairs in dorsal 6.6 4 10 >50
kinety 6, number 2c 6.8 5 10 >50
2m 6.0 11 >50

1 After protargol impregnation.

2 After fIXation with Bouin.
3 It is not clear if these values are valid for the population of Cameroon or Benin.

mine drainage, Indiana, USA (LACKEY 1938c); at 20-24 °C and pH 6.5-7.0 in the Doug-
las Lake and in a small stream in this region (Michigan) and in the Cape Fear River,
North Carolina, USA (CAIRNS & YONGUE 1966, 1973a); spring-fed pond with bottom of
leaf and twig litter, silt and a white marly clay-sand mixture, North Carolina, USA
(YONGUE & CAIRNS 1971); Devil's Lake complex, North Dakota, USA (EDMONDSON
1920); pond in the botanical garden of the University of Pennsylvania, USA, during win-
ter (WANG 1928); Rock Creek River with medium hard water, Pennsylvania, USA (PAT-
RICK 1961); Reelfoot Lake, Tennessee, USA (BEVEL 1938); New River at 19 °C, pH 7.3,
8 mg J-l O2 (YONGUE & CAIRNS 1979) and pond in Virginia, USA (HENEBRY & CAIRNS
1980); Brazil (CUNHA 1913).
RICHARDS & CUTLER (1933) recorded Pleurotricha lanceolata from filters for the pu-
rification of waste waters from beet sugar factories.
Records from marine (EICHWALD 1852, FERNANDEZ-LEBORANS & NOVILLO 1993) and
terrestrial habitats (BODENHEIMER 1935, CHAUDHURI 1929, DIXON 1937, 1939, GOODEY
1911, HARTMANN 1956, LoUSIER & BAMFORTH 1990, NIKOUUK & GELTZER 1972, Rico FER-
RAT 1985, ROSA 1957a, SUDZUKI 1979, VARGA 1936) not documented by original illustra-
tions.
Feeds on bacteria, algae (diatoms), flagellates (Trachelomonas sp., Euglena gracilis,
Astasia klebsiz), and ciliates like Tetrahymena and Dexiostoma campylum (HALL 1943,
LILLY 1942, LILLY et al. 1949, RIGGENBACH 1922, WANG & NIE 1933). However, in
bacteria-free cultures P. lanceolata failed to grow on Tetrahymena sp. (LILLY 1953).
PENN (1935) obtained extraordinarily large specimens (300-350 /lm) by feeding the pleu-
rotrichs their own cysts. Cannibalism is also reported by LILLY (1942). Glaucoma scintil-
lans and Chilomonas paramaecium are inadequate as food, and the algae Stichococcus
bacillaris and Chlorella sp. and a number of single strains of bacteria did not support
continued growth of P.lanceolata (LILLY 1942, see also KIDDER 1951).
Average division rate 1-2.05 (GREENLEAF 1926, WOODRUFF 1905a, see also HAMMOND
1938). BAITSELL (1914) found a remarkably high rate of 4 divisions per day for a 40 d pe-
riod with hay infusion as culture medium (see also JENNINGS 1939, ZAiKA 1970). JEFFRIES
(1956) used a 0.15 % (w/v) extract ofCerophyl for cultivation. Sterile Cerophyl solutions
were added to sterile 60 mm petri dishes and inoculated with Enterobacter aerogenes. A
712 SYSTEMATIC SECTION

day or two later they were inoculated with Telrahymena sp. and set aside until large num-
bers of well-feed animals were present. Each of these cultures were seeded with several
specimens of P. lanceo/ala. MARTiN-GONzALEZ et aI. (1984) used the following culture
procedure: test tubes containing 10 ml sterile lettuce infusion (1.5 g of lettuce powder and
1.5 ml CaC03 per litre distilled water, pH 7.0) were inoculated with E. aerogenes and in-
cubated 24 h at 37°C; 1 ml of a rich Glaucoma scinlillans culture was added as food
24 h later, and 30 specimens of P. lanceolala from the original samples were put into
each tube. Populations were kept at an optimum by sub-culturing every 2-3 days.
According to PENN (1959) P. lanceo/ala may be cultured in a hay-rye infusion with
Colpidium or in a physiological medium with Chlorogonium as food. For experimental
work, the latter method is preferred. A 0.2 % hay and 0.2 % rye infusion is prepared by
boiling 1 gof hay and 1 g of rye in 600 ml of spring water for 8 min. After boiling and
cooling, there are about 500 ml of solution. Then half of the rye grains are removed, leav-
ing the other half with all the hay in the solution. This is then transferred to a 1 litre bat-
tery jar and left for 2 d in order to allow bacteria to grow. When the infusion is 2 d old
and contains many bacteria, it is inoculated with Colpidium (or Chilomonas). After 24 h
there are numerous colpidia present in the infusion. This is then inoculated with Pleura-
tricha. From time to time rich cultures of separately raised colpidia are added to the jar as
additional food supply.
When doing physiological work where bacteria and organic matter are to be avoided,
PENN (1959) recommends a physiological medium consisting of inorganic salts may be
prepared according to the following formula: CaCb 0.0008 N, NaN03 0.0003 N, MgS04
0.0002 N, K2HP04 0.0001 N, KH2P04 0.0001 N, NaN03 0.0008 N. Boveri dishes of
50 ml capacity provided with covers may be used for culturing Pleurotricha. Place 20 ml
of the medium in each Boveri dish. Add one pipette of concentrated culture of Chloroga-
nium, cultivated separately with the same medium, to each dish. Then transfer one or sev-
eral pleurotrichs into each dish. Cover and place the cultures in a well-lit part of the
room. Pleurotrichs so cultivated are morphologically and physiologically large and uni-
form. They divide four times a day. Some autecologicaI data are shown in Table 27.
It is somewhat strange that Pleurotricha lanceolala is not classified saprobiologi-
cally, whereas P. grandis, which occurs much more rarely, can be found in many lists.
According to the faunistic and ecological data, P. lanceolala probably prefers eutrophic
stagnant waters. LACKEY (l938a) found it in ponds, pools and still areas along stream
margins, polluted streams, and in trickling filters in the USA, indicating a distribution
over a wide saprobic range with a preference of beta- and alphamesosaprobic areas.

Pleurotricha grandis STEIN, 1859 (Fig. 190a, b)

1859 Pleurotricha grandis STEIN, Lotos, 9: 4 (no illustration).

1859 Pleurotricha grandis. STEIN - STEIN, Organismus der Infusionsthiere I, p 169 (Fig. 190a).
1882 Pleurotricha grandis, STEIN - KENr, Manual infusoria II, p 782.
1932 Pleurotricha grandis STEIN, 1859 - KAHL, Tierwelt Dtl., 25: 593.
 Pleurotricha 713

1935 Pleurotricha grandis STEIN 1859 - WANG & NlE, Sinensia, Shanghai, 6: 495 (Fig. 190b).
1979 Pleurotricha (pleurotricha) grandis - JANKOWSKI, Trudy zoo!. lnst., Leningr., 86: 62 (erroneously fIxed
as type of subgenus, see nomenclature of genus).
1991 Pleurotricha grandis STEIN, 1859 - FOISSNER, BLATIERER, BERGER & KOHMANN, lnformationsberichte
Bayer. Landesamtes filr Wasserwirtschaft, 1191: 275.

T a x 0 nom y: Detailed redescription necessary. Pleurotricha grandis sensu KOFFMAN

(1926) is only about 98 flm long and occurred in the soil of a garden. Since the size is not
even the half of STEIN'S (1859b) smallest specimens and since P. grandis is not reliably
recorded from terrestrial habitats (see FOISSNER 1987a), the identification is rather doubt-
fully. It is thus classified as an insufficient redescription (Fig. 226b), together with the
populations by GooDEY (1911; Fig. 223a), RAo (1928; Fig. 231a), and SCHMALL (1976;
Fig. 226g).

M 0 r p hoi 0 g y and b i 0 log y: After STEIN (1859b) in life 21~15 flm long, af-
ter WANG & NIE (1935) 300 x 140 flm on average. Broad elliptical, anterior end rounded,
posterior slightly tapered. According to STEIN (1859b) body firm, after WANG & NIE
(1935) however ''rather flexible". Usually 2 micronuclei. Rapid movement. 3-4 (usually
4) left and right marginal rows respectively. Rows become shorter at both ends from out-
side to inside. Inner right marginal row begins at about level of cytostome, terminates
near pretransverse ventral cirri, consists of only about 9 cirri. Inner left marginal row lo-
cated beside postoral ventral cirri, made up of3-5 cirri only. Dorsal pattern unknown.

o c cur r e n c e and e colo g y: Rare; in detritus of eutrophic (mesosaprobic) stag-

nant waters. Eurasia. Locus classicus is a trout-pond in the village of Tharand near Dres-
den, Germany, where STEIN (1859a, b) discovered Pleurotricha grandis among Calli-
triche sp. The highest number and the largest specimens occurred 8 days after sampling,
when the plants began to decompose. WANG & NIE (1935) found it in great abundance
among decaying organic matter and aquatic plants in Lake Ho Hu, Nanking, China.
Records from freshwater habitats not substantiated by illustrations: rather abundant
in a lake (Lansersee) in Tyrol, Austria (DALLA TORRE 1891); puddle in Estonia (JACOBSON
1928); polluted font in Germany with green, foul water (GERSCHLER 1910); caves and
bogs in Germany and Switzerland (GITTLESON & HOOVER 1969, GRIEPENBURG 1934, 1935,
1939, MESSIKOMMER 1954, SCHMIDT 1916); lake (GroBes Heiliges Meer) in Germany
(KEMPER 1930); Hamburg harbour, Germany (ROY 1938); Italy (ENRIQUEs 1913; see also
DINI et al. 1995); rare in May in the botanical garden of Basle, Switzerland (RIGGENBACH
1922); Ukraine (DoBROVLIANSKI 1914); periphyton of Lake Dong Hu, China (SHEN 1980);
infusion of dry leaves in Lahore, India (BHATIA 1920); pond near Lake Erie, Ohio, USA
(STEHLE 1923).
BOUTCHINSKY (1895) and BUTSCHINSKY (1897) found Pleurotricha grandis in saline
inland waters in the USSR, AzOVSKY (1996) in the Barents Sea. Records from terrestrial
habitats are unreliable (BHATIA 1936, CHAUDHURI 1929, KOFFMAN 1934, RAo 1928, SAN-
DON 1927, SHIBUYA 1927)_
714 SYSTEMATIC SECTION

Fig. 190a, b: Pleuro-

tricha grandis (a, after
STEIN 1859b; b, from
WANG & NIE 1935).
Ventral views from
life, a = 210-415 Jim,
b = 275 JIm; P 712.

Voracious feeder devouring

mainly diatoms and other unicellular
algae. Half digested remnants are usu-
ally found crowding the cytoplasm.
Biomass of 106 individuals about
1300 mg (FOISSNER et al. 1991). GAUSE
(1936a, b) found Pleurotricha grandis
at pH 6.3-7.9; at pH 7.6 he counted up
to 6000 indo 1.1• Hydrogen sulphide is
not tolerated (WETZEL 1928b).
FmSSNER (1988a), MAUCH (1976),
SLADECEK (1963), SLADECEK et al.
(1981), and WEGL (1983) classified P.
grandis as betamesosaprobic indicator
of water quality (b; b = 10, I = 5, SI =
2.0). However, according to the data
a by STEIN (1859b) and WANG & NIE
(1935) beta- to alphamesosaprobic or
alphamesosaprobic conditions were preferred. Thus, the saprobic classification has to be
verified on reliably determined populations (FOISSNER et al. 1991).
 Pleurotricha 715

•
• •
V2 ijt\Vl
RMC2 mm • TC

cc ..
a llJJi!!! • ~ b

Fig. 19la, b: Pleurotricha p/anensis, a species indetenninata (from FERNANDEZ-LEBORANS 1984. a, from life
(1); b, silver carbonate impregnation). a) Ventral view, about lOS J.lm. b) Ventral infraciliature, size not indi-
cated. Explanation of original labels: AM =anterior microtubule bundle, AZM = adoral zone of membranelles,
BC = buccal cirrus, CC = caudal cirri, FC = frontal cirri (including some frontoventral cirri), IMF = intennem-
branellar fibres, Kd = kinetodesmal fibre, LMC = left marginal cirral row, Ma = macronucleus, Mi = micronu-
cleus, PFI,2 = undulating membranes, PM = postcirral microtubule bundle, RMCI,2 = inner and outer right
marginal cirral row respectively, SOF =subOral fibres, TC = transverse cirri, VI,2 = pretransverse ventral cirri;
p 715.

Species indeterminata

Pleurotricha planensis FERNANDEZ-LEBORANS, 1984, J. nat. Hist., 18: 939 (Fig. 191a, b).
Remarks: In life (?) 102-108 x 57-60 J.lm. The description, the illustrations, and the mi-
crographs are of very poor quality. Postoral ventral cirri obviously absent (or overlooked
because of the low quality of the impregnation). Caudal cirri very prominent, conse-
716 SYSTEMATIC SECTION

quently classification in Pleurotricha certainly incorrect. I doubt that the infraciliature is

correctly shown. I thus avoid any speculation as to which taxon it could belong. The un-
dulating membranes are also described by FERNANDEz-LEBoRANs (1985). Locus classicus
is the coast of the Mediterranean Sea in the Morro del Gos area, Castellon, Spain.

Insufficient redescriptions

Pleurotricha grandis ? (STEIN) - GOODEY, 1911, Proc. R. Soc., 84: 169 (Fig. 223a). Re-
marks: Has, unequivocally, only 1 left and 1 right marginal row. According to size, infra-
ciliature, and edaphic habitat it could be a Sterkiella histriomuscorum, which is rather
common in soil. England?

Pleurotricha grandis ST. - KOFFMAN, 1926, Acta zool. Stockh., 7: 311 (Fig. 226b). Re-
marks: The small size (about 98 ~m long) and the edaphic habitat (garden near Stock-
holm, Sweden) indicate a misidentification (see taxonomy of this species). Perhaps a re-
drawing from STEIN (1859b).

Pleurotricha grandis - RAo, 1928, J. Indian Inst. Sci., IIA: Plate II, Fig. 8 (Fig. 23 1a).
Remarks: The figure shows only that it is a hypotrich ciliate. Soil in India.

Pleurotricha grandis STEIN 1859 - SCHMAll, 1976, Diplomarbeit, p 36 (Fig. 226g). Re-
marks: Possibly an Allotricha. 90-140 ~m, 3 times as long as broad. Very flexible. Three
right and 2 left cirral rows (including marginal ones). 32 adoral membranelles. 16-21 ~m
long transverse cirri protrude beyond posterior end of cell. Feeds mainly on sulphide bac-
teria (ciliates thus appear reddish), flagellates, single celled green algae, and diatoms
were also ingested. Pool in Bonn, Germany.

Pleurotricha lanceolata EHR. - EDMONDSON, 1906, Proc. Davenport Acad. Sci., 11: 99
(Fig. 234k). Remarks: Possibly a Histriculus because only one right marginal row is pre-
sent and caudal cirri are lacking. Freshwater, among algae. Iowa, USA.

Pleurotricha mim. 1 - IzQUIERDO, 1906, Protozoos, P 190 (Fig. 241f). Remarks: In life
280 11m long. Five conspicuous caudal cirri and thus certainly not a Pleurotricha. Fresh-
water in Chile.

Pleurolricha nUm. 2 - IzQUIERDO, 1906, Protozoos, p 190 (Fig. 241g, h). Remarks: In life
100-140 ~m long. Cyst 112 ~m across. Feeds on Arcella. Possibly a Pleurotricha lan-
ceo/ala. Freshwater in Chile.

P/eurotricha sp. - CHAUDHURI, 1929, AnnIs Protist., 2: Plate II, Fig. 31 ( Fig. 226c). Re-
marks: Soil in India.
 Rigidocortex 717

Pleurotricha sp. - CONN, 1905, Bull. Conn. st. geol. nat. Hist. Surv., 2: 59 (Fig. 226a).
Remarks: According to the arrangement of the cirri it could be classified as Pleurotricha
grandis, as recently suggested by ForssNER et al. (1991). However, because of the small
size (95 11m), I avoid such a synonymisation. Freshwater habitat in Connecticut, USA.

Pleurotricha sp. - NII<OUUK & GELTZER, 1972, Pocvennye prostejsie SSSR, 1972, P 132
(Fig. 225k). Remarks: Soil in USSR

Pleurotricha sp. - OLIVEIRA, KRAu & NASCIMENTO, 1954, Mems Inst. Oswaldo Cruz, 51:
409 (Fig. 2340). Remarks: In a cistern in Rio de Janeiro, Brazil.

Pleurotricha sp. - WEST & LUNDIN, 1963, Pap. Mich. Acad. Sci.,48: 108 (Fig. 220b).
Remarks: See also LUNDIN & WEST (1963, P 167, Fig. 7). Freshwater in Michigan, USA.

Stylonychia lanceolata E. - CIENKOWSKY, 1855, Z. wiss. Zool., 6: 301,306 (Fig. 222m,

n). Remarks: Very likely not a Pleuotricha lanceolata because the cyst has a smooth sur-
face.

Rigidocortex gen. nov.

C h a r act e r i sat ion: Undulating membranes in Oxytricha pattern. One right and 1
left row of marginal cirri, slightly separate posteriorly. More than six dorsal kineties.
Dorsal morphogenesis in Oxytricha pattern, however, kinety 3 not with simple, but with
multiple fragmentation.

T Y pes p e c i e s: Australocirrus octonucieatus ForsSNER, 1988.

T a x 0 nom y: BLATTERER & FOISSNER (1988) established Australocirrus with two spe-
cies, namely A. oscitans (type) and A. octonucieatus, because both show multiple frag-
mentation of dorsal kinety 3. However, they differ significantly in body flexibility. Fur-
thermore, in the very flexible Australocirrus oscitans cirrus V/3 (posterior postoral ven-
tral cirrus) is involved in primordia formation, supporting the classification in the Oxytri-
chinae. Conversely, the body of A. octonucieatus is rigid, suggesting that it is not the sis-
ter group of A. oscitans. I thus transfer A. octonucieatus to the Stylonychinae. It differs
from the rather similar Sterkiella cavicola in the higher number of macronuclear nodules
and the multiple fragmentation of dorsal kinety 3. The latter character is the autapomor-
phy of Rigidocortex. It differs from other taxa of the Stylonychinae with multiple frag-
mentation (Pattersoniella, Laurentiella, Onychodromus) in that it has the typical18-cirri
pattern.
718 SYSTEMATIC SECTION

Single species

Rigidocortex octonuc/eatus (FOISSNER, 1988) comb. nov. (Fig. 192a-l,

Table 43)

1988 Australocirrus octonculeatus FOISSNER, Stapfia, 17: 123 (the slide ofholotype specimens is deposited in
the OberOsterreichische Landesmuseum in Linz, Upper Austria).

T a x 0 nom y: See genus section.

M 0 r p hoi 0 g y and b i 0 log y: FOIsSNER (1 988b) investigated 4 populations, one

each from Kenya, Australia, Austria, and Germany. Because they agree very well in all
important characters, they were not separately described in the original description. In all
populations very small and very large specimens occurred; many characters thus show
high coefficients of variability.
Type material (population from Kenya) in life about 180 x 80 J-lm, populations from
Europe about 260-300 x 130 J-lm. Oval, that is, slightly converging anteriorly (Fig. 192c,
h) or margins almost in parallel (Fig. 192a, d). Body rigid and not contractile. Flattened
2-3:1 dorso-ventrally. Cross-section with strongly flattened lateral areas, ventral side
slightly concave, central dorsal part of cell distinctly arched (Fig. 192b, e). Macronuclear
nodules spherical to slightly ellipsoidal, longitudinally arranged in about median of cell;
number of nodules rather invariably on average: 7-9 (median = 8, n = 16; type material);
7-10 (median = 8, mean = 8.2, SD = 0.9, CV = 11.3 %, n = 13; population frOID Austria),
7-13 (median = 10, mean = 10.1, SD = 1.9, CV = 18.9 %, n = 15; population from Ger-
many). Macronuclear nodules in life about 15-18 x 14-15 J-lm, after protargol impregna-
tion 8-15 x 7-13 J-lm. 3-9, usually 5 micronuclei near macronucleus, in life about 4 J-lm
across. Contractile vacuole about in mid-body, during diastole with conspicuous collect-
ing canals; sometimes canals appear to consist of single vesicles (Fig. 192h). Cortical
granules lacking (checked in live specimens and with methyl green-pyronin stain). Pelli-
cle and cytoplasm colourless, latter densely filled with 2-10 J-lm sized, yellowish cyto-
plasmic crystals so that cells appear dark at low magnification and bright field illumina-
tion, sometimes many 4-6 J-lm sized, greasily shining globules in the marginal area (Fig.
192e). Moves hastily to and fro, stiff like a board.
Adoral zone of membranelles about 45 % of body length. Bases of largest mem-
braneIles in life 20-28 J-lm broad, cilia of membranelies up to 30 J-lm long. Buccal area
broad, proximal part deep. Undulating membranes distinctly curved, in small specimens
they intersect somewhat behind the middle (Fig. 192i), in large specimens they are ar-
ranged almost side by side and intersect in proximal quarter (Fig. 1921). Anterior and
posterior fifth of paroral consists of basal bodies arranged in zigzag, middle part made up
of short obliquely arranged kineties of 3--4 basal bodies each. Frontal cirri in life
30-35 J-lID long, usually arranged right of median, especially in large specimens. Com-
 Rigidocortex 719

Fig. 192a~ Rigidocortex octonucleatus (from FOISSNER 1988b. a-g, from life. a, b, f, g, type material from
Kenya; c, e, population from Germany; d, population from Austria). a, C, d) Ventral and dorsal views, a =
235 Jim. b) Right lateral view. e) Cross section. The small circles are globular inclusions in the marginal area
of the cell. f) 2-10 Jim sized cytoplasmic crystals. g) Resting cyst, 60 Jim across; p 718.

paratively often (3 of 11) specimens with 2 (Fig. 192k) or 4 frontal cirri occur. 3-7, usu-
ally 5, in life 35-40 ~m long, slightly fringed transverse cirri, arranged almost longitudi-
nally and sometimes subterminally, do not protrude (or only slightly) beyond posterior
end of cell. Marginal cirri in life about 25 ~m long, rows almost confluent posteriorly.
Two almost unshortened dorsal kineties, each with 1 caudal cirrus; other kineties slightly
to very distinctly shortened, including posterior fragment of"kinety 3" which also bears a
caudal cirrus (Fig. I92j). Some morphogenetic stages show that only 2 dorsomarginal ki-
 720 SYSTEMATIC SECTION

,,
,
I
I
,, ..'
, •
f

...
.-

h
I
I
, \

"
• •

Fig. 192b-j Rigidocortex octonucleatus (from FOISSNER 1988b, type material from Kenya h, from life; i, j,
protargol impregnation). h) Dorsal view showing contractile vacuole complex during diastole. i, j) Ventral and
dorsal infraciliature, 115 11m; p 718.

neties are fonned. Thus, the other shortened kineties probably originate by multiple frag-
mentation and/or by retention of parental kineties. All together about 9-13 dorsal kine-
ties.
Resting cysts spherical, 50-62 !lm (mean = 58 !lm; SD = 4.0 !lm; CV = 6.9 %; n = 8)
across, covered by a 5-6 !lm thick, irregularly folded wall (Fig. 1929).

o c cur r e n c e and e col 0 g y: Very common; rather certainly confmed to terres-

trial habitats, especially in litter of deciduous forests, where it is regularly associated with
Territricha stramenticola (own observations). Locus classicus is a deciduous forest in
Kenya, about 300 m from the shore of Lake Nakuru, where ForssNER (1988b) discovered
Rigidocortex octonucleatus in the upper soil layer (0-5 cm; brown-grey; high content of
humus; densely grown with grass). ForssNER (1988b) found it also in the litter of beech
forests near Munich, Gennany, and Salzburg city, Austria. BLATfERER & FOISSNER (1988)
found R octonucleatus in the upper soil layer with litter and roots of a Eucalyptus forest
in the Belair National Park near Adelaide, Australia. Further records: soil (0-8 cm,
pH 5.1) from blackwater inundation primary (?) rain forest on Anavilhanas archipelagos
 Rigidocortex 721

"~"~It
\ I

,aD r--. .-
~.
~

',-
cst)" .
,
,.., .., ..
~QQ~
~.,~/
.. • # # •

~
~
-.
Fig. 1921\, I Rigidocortex octonucleatus (from FOISSNER 1988b. k, I, protargol impregnation). Ventral infra-
ciliature of a large (k = 185 11m) and a very large (I = 275 11m) specimen of the type material from Kenya
Macronuclear nodules shown in outline only; p 718.

in the Rio Negro, Brazil (ForssNER 1997b); dark brown soil mixed with much leaf litter
(PH 7.1) from Amazonian rain forest near Iquitos, Peru (FOISSNER 1997b).
Feeds on ciliates (Colpoda stein ii, C. cucullus, Opercularia arboricolum, Vorticella
astyliformis, Leptopharynx costatus, hypotrichs, for example, Euplotes sp.) and hetero-
722 SYSTEMATIC SECTION

trophic flagellates. Sometimes bacteria, fungal spores, Euglena sp., and testaceans (Eu-
glypha sp.) can be found in the voluminous food vacuoles. Biomass of 106 individuals
about 900 mg (population from Europe).

Table 43 Morphometric data of Rigidocortex octonucleatus (from FOlsSNER 1988b). All data are based on
protargol-impregnated specimens. All measurements in micrometres. CV = coefficient of variation (in %),
Max = maximum value, mean == arithmetic mean, Min = minimum value, n = sample size, SO = standard de-
viation

Character mean SO CV Min Max n

Body, length 142.6 9.0 20.9 108 195 11
Body, width 71.2 16.5 23.1 52 110 11
Adoral zone of membranelles, length 65.1 16.6 25.5 47 95 11
Adoral membranelles, number 49.4 8.7 17.6 38 67 II
Right marginal row, number of cirri 27.4 3.4 12.3 22 33 11
Left marginal row, number of cirri 24.6 1.9 7.8 21 27 11
Dorsal kineties, number 10.2 1.3 12.9 8 13 lO

Onychodromus STEIN, 1859

1859 Onychodromus STEIN, Lotos, 9: 4 - Type (original designation): Onychodromus grandis STEIN, 1859.
1859 Onychodromus STEIN - STEIN, Organismus der Infusionsthiere I, p 143.
1882 Onychodromus, STEIN - KENT, Manual infusoria II, p 766.
1932 Onychodromus STEIN, 1859 - KAlIL, Tierwelt OtI., 25: 621.
1972 Onychodromus STEIN, 1859 - BoRROR, 1. Protozool., 19: 14.
1974 Onychodromus STEIN - STILLER, Fauna Hung., 115: 163.

C h a r act e r i sat ion: Undulating membranes in Stylonychia pattern. Number of

frontal-ventral-transverse cirri increased, that is, more than 18. One right and 1 left row
of marginal cirri, separate posteriorly. More than 6 dorsal kineties. Caudal cirri present.
Cytoplasmic processes (horns) on dorsal surface. Frontal-ventral-transverse cirra1 primor-
dia ofproter and opisthe originate independently. At least 1 dorsal primordium with mul-
tiple fragmentation and 2 or more dorsomarginal kineties.

Nom e n cia t u rea n d t a x 0 nom y: Incorrect subsequent spellings: Onchodro-

mus grandis STEIN (CONN 1905, legend to Plate XXVI; MAUCH 1976, p 432); Onyocho-
dromus (WRIGHT & LYNN 1997, p 334); Onichodromus grandis (STEIN) (MAoRAZO-
GARIBAY & L6PEZ-OCHOTERENA 1973, p 67); Onychdromus (WICKLOW 1996); Onychodo-
mus grandis (SONNEBORN 1964, P 696); Onychodroma grandis (ST.) (PROWAZEK 1910,
p 154); Onychodromas (MCGRADy-STEED & MORIN 1996, P 97); Onychodromas grandis
(for example, BAMFORTH 1963, P 133; SULLIVAN 1957, P 195); Onychodromis grandis
STEIN (CAIRNS & YONGUE 1973b, P 190); Onychodromous quadricornutus (EMBLEY et al.
1995, p 88); Onychodronus (LANGLOIS & COATS 1997, P 174); Onycodromus STEIN
 Onychodromus 723

(GRANDORI & GRANDORI 1934, p 292); Orynchodromus grandis (KARL) (SZTRANTOWICZ

1984, p 74); Oxychodromus grandis, STEIN (KENT 1882, P 766).
Onychodromus very probably belongs to the Stylonychinae because the body is rigid,
the adoral zone of membranelles is more than 40 % of body length, and cortical granules
are lacking (Fig. 25a). This position is supported by the Stylonychia pattern of the undu-
lating membranes (Fig. 193p, y, 195e, h; see chapter 2 in the general section) and mo-
lecular biological data which suggest that Onychodromus quadricornutus is more closely
related to Stylonychia pustulata and "Oxytricha nova" (a species of the Sterkiella his-
triomuscorum complex) than to Oxytricha granulifera (SCHLEGEL et al. 1991; see also
BERNHARD et al. 1995, EMBLEY et al. 1992, GREENWOOD et at 1991a, b, Hm.T et al. 1995,
LEIPE et al. 1994, SCHLEGEL 1991). Accordingly, the increased number of cirri per streak
(Onychodromus grandis, Onychodromus quadricornutus) and the increased number of
streaks (0. quadricornutus) evolved from the 18 frontal-ventral-transverse cirral pattern.
The synapomorphy of Onychodromus grandis and o. quadricornutus are the dorsal
cytoplasmic processes, a curious character which is, unfortunately, not mentioned in the
original description (STEIN 1859a), some revisions (for example, CuRDs et al. 1983, KARL
1932, KENT 1882), and redescriptions of the type species (JONES 1974, LUNDIN & WEST
1963, SMITH 1914, TUFFRAU & FLEURY 1994). The function of these processes is not com-
pletely understood, however, in O. quadricornutus these horns can prevent intraspecific
predation (WICKLOW 1988b). Aspidisca turrita (EHRENBERG, 1831) CLAPAREoE & LACH-
MANN, 1858 evolved a similar horn independently (FOIssNER 1994f, FOISSNER et al. 1991).
Reviewers of the original description of Onychodromus quadricornutus considered the
decision to ally this conspicuous species with o. grandis as being too conservative. They
suggested a new genus, which would, however, not increase the understanding of the phy-
logenetic relationships ofthis species group.
Onychodromus quadricornutus also shows some conspicuous agreements with
Kerona pediculus, the well-known ectocommensal on Hydra and on bryozoans: (i) Dor-
sal morphogenesis is more or less identical, that is, all 3 primordia show multiple frag-
mentation and 2 dorsomarginal kineties originate, resulting in a high number of dorsal ki-
neties (FOISSNER et al. 1987b). (ii) New marginal rows originate without involvement of
parental marginal cirri. This peculiarity is known only from these 2 species. (iii) The
number of cirri per streak is distinctly increased. However, I failed to construct a plausi-
ble cladogram including Onychodromus grandis, O. quadricornutus, and Kerona pedicu-
lus without assuming too many parallelisms or reductions.
According to CORLISS (1979, p 311), Peritromoides BHATIA, 1936 is a junior syno-
nym of Onychodromus. However, the rather superficially described type species, Peritro-
moides simplex BHATIA, 1936, is reminiscent of Hypotrichidium and was thus transferred
to this taxon by JANKOWSKI (1979). CURDS et al. (1983, p 432) proposed synonymy with
Himantophorus EHRENBERG, 1838, which is, however, very likely a euplotid because the
contractile vacuole is in the right posterior portion ofthe cell (EHRENBERG 1838).
 724 SYSTEMATIC SECTION

Key to species
1 2 dorsal processes; usually 4, sometimes up to 8 macronuclear nodules; 5 or 6 trans-
verse cirri; 1 contractile vacuole in about mid-body; frontoventral cirri not in highly
ordered rows (Fig. 193a, k, I, r) ........................... 0. grandis (p 724)
3 or 4 dorsal processes; 10-30, usually 11-19 macronuclear nodules; 9-16 trans-
verse cirri; several contractile vacuoles; frontoventral cirri in about 13 highly ordered
rows (Fig. 195a-c, h, i) .............................. O. quadricornutus (p 738)

Onychodromus grandis STEIN, 1859 (Fig. 193a-y, 194a-n, Table 44)

1859 Onychodromus grandis STEIN, Lotos, 9: 4 (no illustration; dorsal processes not mentioned).
1859 Onychodromus grandis STEIN - STEIN, Organismus der Infusionsthiere I, 145 (Fig. 193a-f; dorsal proc-
esses illustrated and mentioned).
1914 Onychodromus grandis STEIN - SMI1lI, Kans. Univ. Sci. Bull., 9: 165 (Fig. 1939).
1932 Onychodromus grandis STEIN, 1859 - KAHL, Tierwelt Otl., 25: 621 (Fig. 193h).
1934 Onycodromus grandis STEIN - GRANDORI & GRANDORI, Boll. Lab. Zool. agr. Bachic. R. 1st. sup. agr. Mi-
lano, 5: 292, Tavola XIII, fig. 280 (incorrect subsequent spelling of Onychodromus; very likely a re-
drawing).
1963 Onychodromus gram/is STEIN - LuNDIN & WEST, Free-living Protozoa, p 68 (Fig. 193i).
1972 Onychodromus grandis STEIN, 1859 - BORROR, J. Protozool., 19: 14.
1974 Onychodromus grandis STEIN, 1859 - JONES, Univ. South Alabama Monogr., 1: 41 (Fig. 193j).
1991 Onychodromus grandis STEIN, 1859 - GsCHWIND, Oiplomarbeit, p 43 (Fig. 193k-q).
1992 Onychodromus gram/is var. simplex SZABO, Europ. J. Protistol., 28: 358 (no illustration, see taxonomy).
1994 Onychodromus gram/is - TUFFRAU & FLEURY, Traite de Zoologie, 2: 142 (Fig. 194n).
1995 Onychodromus gram/is STEIN 1859 - SZABO & WILBERT, J. Euk. Microbiol., 42: 50 (Fig. 193t-y,
194a-1).

Nom e n c I a t u rea n d t a x 0 nom y: Incorrect subsequent spellings: Onychodro-

mus gradus (WALKER & GRIM 1973, p 566); Pnychodromus grandis var. simp/ex (SZABO
1996). The redescriptions by GSCHWIND (1991) and SZABO & WILBERT (1995) agree
largely with the STEIN (1859b) description, especially as concerns the dorsal processes.
However, STEIN'S (1859a, b) population had distinctly more frontal, buccal, and frontov-
entral cirri (16-28) and more postoral and pretransverse ventral cirri (15-21) than the
GSCHWIND (1991) and SZABO & WILBERT (1995) populations, which have 10-14 and 6-10
cirri respectively (Table 44). SZABO (1992) therefore established the variety simplex.
KAHL's (1932) illustration shows an intermediate number of 14 and 13 cirri respectively
(Fig. 193h), indicating that all populations belong to the same species (GSCHWIND 1991).
The dorsal processes are sometimes inconspicuous (GSCHWIND 1991, STEIN 1859b) and

Fig. 193a-f Onychodromus gram/is (from STEIN 1859b. a-t; from life). a, C, d) Ventral and dorsal (c) view, ~
80-350 11m. Arrows mark dorsal processes. Figure (a) shows a specimen with 6, (c) with 7, and (d) with 5
transverse cirri. b) Old, large specimen (300 x 210 11m) with 5 macronuclear nodules. e, f) Late and very late
morphogenetic stage.
 Onychodromus 725

193a
726 SYSTEMATIC SECTION

Fig. 1939-j Onychodromus grandis (g, after SMI11i 1914; h, from KAm. 1932; i, from LUNDIN & WEST 1963;j,
from JONES 1974. g-j, from life). Ventral views, g = 255 ~m, h = 250 ~m, i = size not indicated, j = 110 ~m.
The identification by JONES (1974) is somewhat uncertain (small size, short adoral zone of membranelIes, sa-
line water); p 724.

were thus very likely overlooked by some workers (JONES 1974, KAHL 1932, LUNDIN &
WEST 1963, SMITH 1914, TUFFRAU & FLEURY 1994).
Onychodromus grandis sensu BORGER (1980) and sensu CONN (1905) are insuffi-
ciently redescribed. The terrestrial (?) population ofMAupAS (1889; Fig. 181o-q) is pos-
sibly Sterkiella cavicola (see there).
The following chapter is based mainly on the redescription by GSCHWIND (1991), who
used the SCHLEGEL & STEINBROCK (1986) population (Fig. 193r, s).

M 0 r p hoi 0 g y and b i 0 log y: In life 120-235 /lm (mean = 185.9 /lID, SD =

36.6/lm, CV = 19.7 %, n = 11) x 60-110 /lm (mean = 82.4/lm, SD = 15.7/lm, CV =
19.1 %, n = 11; GSCHWIND 1991); according to STEIN (1859b) large specimens 350 times
170/lID, old cells broad, that is, 300 x 210 /lm (Fig. 193b), smallest specimens only
80/lm long. Elliptical to almost rectangular (that is, parallel margins), anterior end dis-
tinctly obliquely truncated, posterior end slightly tapered to broadly rounded. Body margin
thin and thus hyaline. Left margin straight, sometimes slightly indented at level of buccal
vertex, right margin convex. Flattened about 2-3:1 dorso-ventrally, ventral side flat to
 Onychodromus 727

Fig. 193k-il Onychodromus grandis (from GSCHWIND 1991. k-n, from life; 0, protargol impregnation). k)
Ventral view, 160 11m. \) Dorsal view showing dorsal horns (arrows) and contractile vacuole with canals. m)
Transverse cirrus with fringed end. 0) Right-lateral view. 0) Dorsal infraciliature of a specimen with 8 macro-
nuclear nodules, 195 11m. Arrowheads mark caudal cirri; p 724.

slightly concave, dorsal side distinctly arched centrally. Dorsally left of median invariably
2 plug-shaped processes, anterior one about at same site as contractile vacuole and thus
probably bearing the excretion pore, posterior one slightly in front of left transverse cirri;
processes usually turned to left margin of cell, sometimes inconspicuous (GSCHWIND 1991,
STEIN 1859b), rounded terminally and filled with a darkish, grained mass; in old specimens
anterior process or both processes lacking (STEIN 1859b). Body firm. Usually 4, sometimes
5, 6, 7, or 8 macronuclear nodules (STEIN 1859b), ellipsoid to spherical, slightly left of me-
dian, with many tiny chromatin bodies. Usually 4 micronuclei (DoGIEL 1929, Table 44), al-
most spherical, usually near macronuclear nodules. Contractile vacuole slightly behind
level of buccal vertex, about at same site as anterior dorsal process, during diastole with an
anterior and posterior canal. Pellicle fragile and colourless. Cortical granules lacking. Cy-
toplasm with some crystals, in well fed specimens with many 1-2 ~m sized, spherical to
lens-shaped bodies. Movement conspicuously jerky.
 728 SYSTEMATIC SECTION

..

."
".
".
,
" ,
,
,, , ". ,/
I

f_ ... ji

.
",
..
I

t, ....
I "
t , tit
t
"
'" p

Fig. 193p, q Onychodromus grandis (from GSCHWIND 1991. p, q, protargol impregnation). p) Ventral infra-
ciliature, 175 11m. The undulating membranes are almost straight and arranged in parallel as in Sty/onychia.
Usually they are inconspicuously bent to the right (cp. Fig. 193a, d). q) Dorsal infraciIiature and nuclear appa-
ratus, 175 11m. The great distance between some kineties is very likely due to the dorsal processes (cp. Fig.
193r, s). This specimen obviously has 3 dorsomarginal kineties, indicating that FOISSNER et al. (1987b, Table
ll) counted too many (namely 6) dorsomarginals; p 724.

Adoral zone of membrane lIes almost 50 % of body length, anteriorly arranged on a

prominent collar, posteriorly covered by a peristomiallip. Bases of largest membranelies
in life about 10-12 !lm broad. Buccal area flat and broad. Undulating membranes almost
straight, sometimes crossing posteriorly. Invariably 3 (n = 24) distinctly enlarged, in life
about 28 !lm long frontal cirri.' Arrangement of frontoventral, postoral, and pretransverse
ventral cirri as in Figures 193k, p, y. Anterior frontoventral and buccal cirri distinctly
larger than posterior one. Buccal cirri and frontoventral cirri arranged in 3 rows, left (=
buccal row) and middle row according to STEIN (I 859b) with 5-10 cirri each (see taxon-
omy according to different number of cirri). Postoral and pretransverse ventral cirri ac-
 Onychodromus 729

Fig. 193r, s Onychodromus grandis (original scanning electron micrographs kindly supplied by M. SCHLEGEL,
Leipzig; same population as Fig. 193k-q). Right lateral and dorsal view, 140 11m. Note the 2 rounded dorsal
processes near the left margin (arrows in r). Near the right margin is a further, however, less conspicuous ele-
vation which was obviously overlooked in live observations; p 724.

cording to STEIN (1859b) usually arranged in 4 rows (Fig. 193a): right row with 4-5 cirri,
next one with 6-9, next one with 2-3, and left row with 3-4 cirri. Transverse cirri in life
30-38 /lID long, distally fringed, hook-shaped arranged, protrude distinctly beyond poste-
rior end of cell (Fig. 193k, ID, p, y); according to STEIN (1859b) 5-7, usually 6 transverse
cirri (this requires an additional cirral primordium). Marginal rows distinctly separated
posteriorly, cirri in life about 20 IlID long. Dorsal cilia in life about 3 /lID long, arrange-
730 SYSTEMATIC SECTION

.. : . .-

,<~;:i~::'~·},>:"

193t
~
~,
~

\
\
.. VI
1

....
\
)
)
}
) ;
J
.,
;

,
I
J ,
I
)
'I,

",!

.. ~
 Onychodromus 731

ment and number of kineties somewhat variable (Fig. 1930, q, s), according to SZAB6 &
WILBERT (1995) usually (n = 15) 9 kineties, including 3 dorsomarginal kineties (Fig.
193x); according to FOISSNER et al. (1987b, Table II) 6 dorsomarginal kineties, which is,
however, not in agreement with Figures 1930, q, s. Invariably 3 caudal cirri (n = 24 and
25; GSCHWIND 1991, STEIN 1859b, SZAB6 & WILBERT 1995), stiff, in life 20-30 !lm long.
Obviously erroneously, CURDS et al. (1983) state that caudal cirri are lacking.
Some ultrastructural observations are reported by TUFFRAU et al. (1968), however,
identification is not supported by an illustration. TUFFRAU (1965b) found macrostome gi-
ants. Cyst contains chitin (BUSSERS & JEUNIAUX 1974).

M 0 r p hog e n e sis is described by STEIN (1859b; Fig. 193e, f) and SZAB6 & WILBERT
(1995; Fig. 194a-l); however, reinvestigation is recommended because some important
details (origin of primordia, relation of cirri to streaks) are very likely (at least partly)
misobserved. Formation of the oral primordium begins somewhat left of the left trans-
verse cirrus (Fig. 194a). Three frontal-ventral-transverse cirral primordia of the proter
originate from the posterior-most cirrus of the right frontoventral row (Fig. 194b, c), indi-
cating that this cirrus is homologous to cirrus IV/3 of some Stylonychinae with 18 cirri
(see Fig. 153g). Furthermore, each one primordium originates in the buccal row (Fig.
194c) or slightly left of it (Fig. 194d; ForSSNER et al. 1987b, Table II) and in the cirral row
behind the right frontal cirrus (Fig. 194c). However, it is uncertain if parental cirri are in-
volved or not (Fig. 194d). The origin of opisthe's primordia is uncertain. According to
Figures 194b-d only few (1 or 2) postoral ventral cirri are involved in primordia forma-
tion. The primordia I-III of the opisthe develop from the oral primordium, as in other

+- Fig. 193t-y Onychodromus grandis (from SZAB6 & WILBERT 1995. t, from life; u-y, protargol impregnation).
t) Ventral view, 190 ~m. u) Dorsal view showing the 2 dorsal processes (horns). v) Dorsal horn. w) Nuclear
apparatus. x, y) Dorsal and ventral infraciliature, x = 250 ~m, Y = 245 ~m. Cirri originating from same streak
are connected by a broken line; however, I doubt that the proposal is correct (cp. Fig. 194m). Explanation of
original labels: I-VI = frontal-ventral-transverse cirral streaks I-VI, 1-3 = ventral rows; p 724.

Fig. 194a--d Onychodromus grandis (from SzAB6 & WILBERT 1995. a-d, morphogenetic stages in ventral -+
view after protargol impregnation). a) Very early stage (260 ~m) showing the beginning of the oral primor-
dium formation (arrow). b, c) Early stages (b = 250 ~m, c = 255 ~m) showing 5 primordia each in both the
proter and the opisthe. One frontoventral cirrus splits into 3 primordia (arrow), identically to in Sty/onychia
pustulata and S. hifaria. Arrowhead marks beginning of marginal primordia formation. d) Middle stage,
265 ~m. Arrowheads denote beginning of cirral segregation and dorsomarginal kineties; p 724. Figures on
p 732.

Fig. 194e-h Onychodromus grandis (from SZAB6 & WILBERT 1995. e-h, morphogenetic stages with nuclear -+
apparatus in ventral view after protargol impregnation). Late stages showing migration of cirri, 240-255 ~m.
Cirri which originate from the same streak are connected by a broken line; however, I doubt that the proposal
is correct (cp. Fig. 194m). New structures black, parental white. I-VI = frontal-ventral-transverse cirral streaks;
p 724. Figures on p 733.
732 SYSTEMATIC SECTION

o
o
.,.
o

o
c
'" ~o
'" t;j.;;J:
&.
offl
of;

..
.."
b
194a

()
Q
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(1
<>
¢
J,
I
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. D·
 Onychodromus 733

::

-.. .-
<>
.. 0

00 CI
~<> " f

.
• 0
:<>
·0 .'"
...
.'" ......
-.:...
-...
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-;

~
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o ,,0 h
734 SYSTEMATIC SECTION

,e
\
\ ,
\ ,
,
,
wr
.1\

~\
,
,
,
" if:
~,
7J

I I
~ If

~, f" f
'J
,\ ,!.,
I', I

I I,. --
.
• I..

~~
\ I"'"
... ...
..

..
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, ....
.' :. " "

.... ~

.' Ie
: '. • 'i'.

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:f)

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4l
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 Onychodromus 735

Fig. 194m, n Onychodromus grandis

(m, modified from GsCHWIND 1991, same
as Fig. 193p; n, from TUFFRAU & FLEURY
1994. m, protargol impregnation; n, from
life?). m) Proposal for the relationships
of cirri to the frontal-ventral-transverse
cirral streaks I-VI, that is, cirri which
very likely originate from the same
streak are connected by a broken line.
Arrows mark cirri which are presumably
homologous to the 18 frontal-ventral-
transverse cirri of the typical oxytrichids.
n) Ventral view, 100-300 ~m; p 724.

oxytrichids. I doubt that SZABO & WILBERT (1995) recognised the relation of the cirri to
the primordia correctly (Fig. 193y, 194e, h, j). According to them, primordium II does
not produce a transverse cirrus, whereas primordium VI develops 2 prominent cirri which
is very unlikely. Unfortunately, the micrographs in SZABO & WILBERT (1995) do not show

<- Fig. 194i-1 Onychodromus grandis (from SZAB6 & WILBERT 1995. i-l, morphogenetic stages after protargol
impregnation). i, j) Ventral infraciliature of a very late stage (i = 250 ~m) and a postdivider (j = 200 ~m). Cirri
which originate from the same streak are connected by a broken line; however, I doubt that the proposal is cor-
rect (cp. Fig. 194m). New structures black, parental white. k, I) Dorsal infraciliature of a middle and a late
stage; parental kineties omitted. Large arrow in (k) marks multiple fragmentation of primordium 1, small arrow
denotes simple (= normal) fragmentation of primordium 3, arrowhead marks dorsomarginal kineties; p 724.
 736 SYSTEMATIC SECTION

Table 44 Morphometric data of Onychodromus grandis (1, from GSCHWIND 1991; 2, from SZABO & WILBERT
1995). Ail data are based on protargol-impregnated specimens. Ail measurements in micrometres. CV = coeffi-
cient of variation (in %), Max = maximum value, mean = arithmetic mean, Min = minimum value, n = sample
size, SD = standard deviation

Character mean SD CV Min Max n

Body, length 161.8 23.0 14.2 100 218 24
2 237.3 26.5 11.0 184 285 25
Body, width 96.9 1604 16.9 70 115 24
2 146.1 16.2 11.0 114 171 25
Macronucleus, number of nodules 4.5 1.0 2304 4 8 39
2 4.0 0 0 4 4 30
Macronuclear nodule, length 1 20.5 3.5 17.1 14 25 24
2 28.2 4.0 14.0 28 32 25
Macronuclear nodule, width 1 lOA 1.7 16.3 7 15 24
2 14.9 2.3 15.0 13 18 25
Micronuclei, number 1 3.5 0.7 2004 2 6 24
2 4.0 0 0 4 4 30
Micronucleus, length 1 4.9 0.5 10.1 3.5 5.5 24
2 5.6 0.4 7.0 4.5 5.6 25
Micronucleus, width 4.2 0.5 11.6 3.5 5 24
2 4.2 0.8 19.0 3.3 5.6 25
Adoral zone of membranelles, length 80.1 7.8 9.7 66 95 24
21 16404 904 6.0 154 180 30
Adoral membranelles, number I 52.6 6.0 9.7 41 59 24
2 64.3 3.9 6.0 54 67 30
Buccal cirri, number 3.1 0.7 23.7 2 4 24
24 3 4 ?
Frontoventra1 cirri, number' 1 6.2 0.6 9.2 5 7 24
Postoral ventral cirri, number 1 6.3 0.7 11.8 5 8 24
22 9.0 0 0 9 9 25
Pretransverse ventral cirri, number 1 1.9 0.3 14.7 I 2 24
Right marginal row, number of cirri 35.0 3.0 8.5 28 39 24
2 37.3 2.8 8.0 35 43 25
Left marginal row, number of cirri 1 24.7 2.1 5.0 21 31 24
2 26.3 0.8 3.0 25 28 25
Transverse cirri, number 5.0 0 0 5 5 24
2 5.0 0 0 5 5 25
I According to these data the adoral zone is almost 70 % of body length, which does not agree with the illus-
trations (Fig. 193y, 194a).
2 Very likely, pretransverse ventral cirri included.
3 Not mentioned in Table 1 ofSZAB6 & WILBERT (1995).

4 From text.

any details. I propose a different origin of cirri based mainly on the assumption that O.
grandis is a typical Stylonychinae with a slightly increased number of cirri per streak
(Fig. 194m).
Parental marginal cirri are involved in primordia fonnation of new marginal rows
(Fig. 194c-i), which is a significant difference to 0. quadricornutus. Dorsal morpho-
 Onychodromus 737

genesis begins with the fonnation of3 primordia. Primordium 1 (= near left body margin,
large arrow in Fig. 194k) shows multiple, and primordium 3 simple fragmentation. Three
(SZAB6 & WILBERT 1995; Fig. 194k) or 6 (FOISSNER et al. 1987b, possibly a misobserva-
tion) dorsomarginal kineties also occur. Caudal cirri originate at the posterior fragments
of primordia 1 and 3 and at posterior end of primordium 2 (Fig. 194k, I).

o c cur r e n c e and e colo g y: Rare in freshwater. Locus classicus is the Baum-

garten area near Prague, Czechoslovakia, where STEIN (1859a, b) discovered Onychodro-
mus grandis very abundantly in a marshy ditch in October and November. He found it to-
gether with Stylonychia mytilus, Tachysoma pellionellum, Paramecium aurelia, and As-
pidisca lynceus. According to KAlIL (1932), strong saprobity preferred.
Records substantiated by illustrations: Gennany? , exact sites not mentioned
(GSCHWIND 1991, who studied the SCHLEGEL & STEINBROCK 1986 population; KAlIL 1932);
temporary, shallow alkaline water bodies of Pentezug, HortobAgy National Park, Hun-
gary, with a dense growth of Glyceria maxima (SZAB6 & WILBERT 1995); pond water
from Kansas, USA (SMITH 1914); New York, USA (HAUSMAN 1934); freshwater in the
Upper Peninsula (counties Baraga and Marquette) of Michigan, USA (LUNDIN & WEST
1963); upper end of Mobile Bay, Alabama, USA, with a salinity of about 2 %0 (JONES
1974).
Records from limnetic habitats not substantiated by illustrations: Tyrol, Austria
(DALLA TORRE 1891); surface film and detritus of ditches regularly contaminated with
pesticides and untreated ditches near Hamburg, Gennany (CASPERS & HECKMAN 1982);
pond near Bonn, Gennany (SCHMIDT 1913, 1916); plankton of Lake Prester near Magde-
burg, Gennany (HONIGMANN 1909); freshwater(?) habitat (Ober-Warnow) near Rostock,
Gennany (LEVANDER 1892); cooling system ofa conventional power station at the Main
River, Gennany (BERNERTH 1982); with a dominance of 0.6-1.8 % in the plankton ofa
side-branch of the Danube River at G5d, Hungary, in September (BERECZKY 1991);
plankton of Tisza River, Hungary (J6SA 1974); Italy (MAnoNI & ULUHOGIAN 1997); north-
ern Russia (MEREscHKOWSKY 1877); rare in saline lakes near Odessa, Ukraine (BOUTClllN-
SKY 1895, BUTSClllNSKY 1897); Mexico (MAnRAZo-GARIBAY & L6PEz-OcHoTERENA 1973);
rarely at 10°C and pH 6.2 in a tundra pool in Alaska, USA (SULUVAN 1957); at up to
14.2 mg}"1 hydrogen sulphide in pools of the Silver Lake Bog and other sites in Iowa,
USA (HEMPSTEAD & JAHN 1940, SHAWHAN et al. 1947); south-eastern Louisiana, USA
(BAMFORTH 1963); in Sphagnum moss of Buell Bog, Minnesota, USA (BOVEE 1979);
planktonic in the Scioto River in Ohio, USA (LACKEY 1938b); Cleveland county, Okla-
homa, USA (GABEL 1927); Conestoga drainage basin, Pennsylvania, USA (CAIRNS
1965a); Mountain Lake region, Giles County and Potomac River, Virginia, USA (BOVEE
1960, PATRICK 1961); in clear, flowing water with abundant plant life in USA? (HAUSMAN
1917); Brazil (CUNHA 1913, PROWAZEK 1910); bottom area of test tubes, New Zealand
(STOUT 1956); mud surface of Blue Lake, Raoul Island, New Zealand (BROWN & PEART
1973). LACKEY (1938a) found Onychodromus grandis with low frequency in a trickling
filter in the USA.
 738 SYSTEMATIC SECTION

GRANDORI & GRANDORI'S (1934) illustration of a specimen from soil matches Figure
193b rather well, indicating that their illustration is not an original. All other records from
terrestrial habitats not substantiated by illustrations (JAMESON 1914, L6PEZ-OCHOTERENA &
ROURE-CANE 1970, NrrwUUK & GELTZER 1972, SANDON & CUTLER 1924, STOUT 1981,
SZTRANTOWICZ 1984); possibly confused with the quadrinucleate Sterkiella cavicola.
Feeds on bacteria, flagellates (for example, Astasia) and their cysts, testaceans (Ar-
cella vulgaris), ciliates like Dexiostoma campylum, Tetrahymena pyriformis, Parame-
cium aurelia, and in culture also on starch (FAURE-FREMIET 1961a, GSCHWIND 1991, SAX-
ENA & SAXENA 1978, STEIN 1859b, SZABO & WILBERT 1995). Also cannibalistic (STEIN
1859b). According to FAURE-FREMIET (1961a) 12°C was preferred in cultures. SAXENA &
SAXENA (1978) cultured Onychodromus grandis in CHALKLEY'S medium and hay infusion
previously inoculated with Aerobacter aerogenes. Culture medium was changed daily to
maintain the animals in exponential growth phase otherwise stationary phase followed.
They estimated a generation time of 20 h ± 35 min at 24 ± 2 °C (± SD?). SZABO & WIL-
BERT (1995) cultured it in mineral water enriched with rice grains. SZABO & WILBERT
(1995) found 0. grandis mainly in June and July together with Bryophyllum tegularum,
Spathidium amphoriforme, Loxophyllum meleagris, Calyptotricha pleuronemoides, Vor-
ticella nebulifera, V. convallaria, Balladinopsis sphaericus (a species indeterminata de-
scribed by VUXANOVICI 1963) at the following conditions: pH 7.1-7.8, 9-19°C, 6.2 to
8.5 mg I-I O2, 430-650 J.1S cm-I, 0.25~.5 %0 salinity.

Onychodromus quadricornutus FOISSNER, SCHLEGEL & PRESCOIT, 1987

(Fig. 195a-z, Table 45)

1985 Styx sp. - LIN & PREsCOTT, J. Protozool., 32: 144 (see nomenclature and taxonomy).
1987 Onychodromus quadricornutus FOISSNER, SCHLEGEL & PREsCOTT, J. Protozool., 34: 150 (Fig. 195a-v; 1
slide ofholotype specimens and 1 slide ofparatype specimens are deposited in the Oberosterreichische
Landesmuseum in Linz, Upper Austria).
1993 Onychodromus indica KAMRA & SAPRA, Acta Protozool., 32: 107 (Fig. 195w, x).

Nom e n c I a t u rea n d t a x 0 nom y: LIN & PRESCOTT (1985) named this species
Styx sp. without giving a description that would satisfies the criteria of the IcZN (1985).
This genus name is thus a nomen nudum needing no further consideration. In addition, it
is a junior homonym of Styx STAUDINGER, a butterfly, and Styx OPPENHEIM, a mollusc

Fig. 195a-g Onychodromus quadricornutus (from FOISSNER et aI. 1987b. a-c, from life; d-g, protargol im- -+
pregnation). a) Dorsal view showing the 4 dorsal horns (species name) and the contractile vacuoles which form
a C-shaped figure, 400 )lm. b, c) Ventral and right lateral view, 435 )lID, 400 )lm. d) Adoral membranelle. e, g)
Ventral and dorsal infraciliature and nuclear apparatus, e = 285 )lm, g = 320 )lm. Arrowhead marks left frontal
cirrus (cirrus Ill). Anterior arrow denotes buccal row, posterior ones mark the 2 pretransverse ventral cirri
which originate from the 2 rightmost cirral primordia and which are probably homologous with the 2 pretrans-
verse ventral cirri of the 18-cirri oxytrichids. 1) Each basal body pair of the dorsal side is surrounded by fibrils
forming a rhomboid figure.
 Onychodromus 739

~---"-'-..
! .,.//' ('

:,\ '",.,;s';"
~
~
!
'~"A" ..
i~l
:'\
'.,,.,.
~:
I .)~~. ': . . I

' \. .4l"f}< '. '. .

\ \e"
" 1-\.:.... . .. .
".:'
·t:"· ...

195a
c
b ,.' i

i " •• ' •••• ~ '." .'. ,

: : : . ' , '• • : . : : ' . ' : ' . ' ,: I \ ....

<'\'. '.
: ......... ).\ .......
.~~ .
~.j .

:}i i
.,'((tf :
"-I:".'"

':~~ :'~:.:. ~.: J;'?>

~¥){,
IJ~
WI
~.~':/
.,\\
9
740 SYSTEMATIC SECTION

Fig. 195h, i Orrychodromus quadricomulus (from FOISSNER et aI. 1987b. h, i, scanning electron micrographs).
Ventral and dorsal view showing the impressive cirral pattern and the 4 prominent dorsal horns. One caudal
cirrus is inserted near the tip of the caudal hom (arrow). Bars = 100 ~m; p 738.

(FOISSNER et al. 1987b). Incorrect subsequent spellings: Onychodromus quadricornatus

(HOFFMAN et al. 1995, p 1281); Onychodromus quadricornutis (EMBLEY et al. 1992,
p 1483); Onychodromus quedricomutus (LEE & KUGRENS 1992, P 539).
Synonymy of Onychodromus quadricornutus and 0. indica was already proposed by
SZABO & WILBERT (I995, P 59) because qualitative differences are present neither during
interphase nor during morphogenesis. They differ in some morphometric characters, in-
cluding the nuclear apparatus (Table 45). The main deviations are the number of micro-
 Onychodromus 741

Fig. 19Sj Onychodromus quadricornutus (from FOISSNER et aI. 1987b). Scanning electron micrograph show-
ing the posterior end of the cell. One caudal cirrus is inserted almost at the tip of the caudal hom, bar = 20 Jim.
CC = caudal cirri, Dc = dorsal cilia; p 738.

nuclei (4-26 against 3-4) and, according to I<AMRA & SAPRA (1993) more importantly,
their size (5 ~m against 6.4 ~m). However, it is well-known that size and shape and many
other characters of these species depend strongly on culture conditions (FOlSSNER et al.
1987b, I<AMRA & SAPRA 1994, WICKLOW 1988b, 1997). Very likely, they are geographical
races of the same species.
Onychodromus quadricornutus should not be confused with Laurentiella strenua,
which lacks dorsal processes and has a distinctly lower number of cirri (Fig. 196). Ony-
chodromus quadricornutus also shows some agreements with the ectocommensal Kerona
pediculus (see relevant chapter in genus section). The comparison of small subunit ribo-
somal RNA sequences assigns Onychodromus quadricornutus very close to Stylonychia
(SCHLEGEL et al. 1991; see also HAMMERsCHMIDT et al. 1996, LYNN 1996, RAGAN et al.
1996).

M 0 r p hoi 0 g y and b i 0 log y: Cells from spring-water cultures in life

370 x 200 x 70 ~m (n = 6) and thus considerably smaller than those from Pringsheim's
medium (Table 45; FOISSNER et al. 1987b), according to LIN & PREsCOTT (1986) and
WICKLOW (1988b) cannibalistic giants in life up to 1 mm long. Chlorogonium-fed indi-
viduals of the synonym, O. indica, in life 205 x 136 (n = 10; I<AMRA & SAPRA 1993),
bacteria-fed specimens after protargol impregnation 145-191 x 56-81 ~m, cannibals
285-376 x 153-205 ~m (I<AMRA & SAPRA 1994). Cell length frequencies in replicates of
a well-fed clone of 0. quadricornutus show normal distributions (Fig. 195y; WICKLOW
742 SYSTEMATIC SECTION

q
f
~ ~
.. '
·. . ~=-
I~
~
§§:'
0
q
f
,
~ · · J~/
I :::- •• /Y"
l(
• •
q .1:'. f •
0:-.'
0".;'
t:

•
'.'
0 0:
••
....•.
f CJ
0
•• • •
, , , {J

CJ
••
. ·...
I
CJ
•• • CJ
••
••
0 • •
••

J
• •
•• ••
Cl
It
0 a
•
.. ••
.•
t:J
It
q •
'
It
~t::l •

2/m
() o
0°00 •• '
o .'
000 ._.
'. ..... m
.. , , , ,
"
()
()

, , , #
,
()
() , , ,
()

()
"
, q

, , , , #

, 1
,
. .
\a .~

~
•"
~

,
~

,,
~
I Fig. 195k-n Onychodromus
.. , quadricornutus (from FOIss-
, , NER et al. 1987b. k-n, ventral
infraciliature of morphoge-
, , • netic stages after protargol im-
, • , pregnation). k-m) Develop-

, • , ment of the oral primordium.

• , Note the round clusters of ba-
, sal bodies near the upper trans-
• ", verse cirri (arrow). Bars =
• ,
, 30I1m, lOOI1m, 100 11m. 0)
Three streaks split off from the
0
anterior end of the oral primor-
f
0 dium. Cirral primordia origi-
• •
f
o
•ot nate within the parental ventral
•
f

t. rows (except for the 2 right-

" t, most rows and the buccal row;
00

'. 't, ••• n

but see text) by disintegration
of some cirri (arrow). Bar =
90 11m; p 738.
 Onychodromus 743

o
o
c
o
o
o
o
o
• 0 <:> 0°
•••• ()I:I 0°
•••• 0
o
Fig. 1950, P Onychodromus quadricornutus (from FOISSNER et aI. 1987b. 0, p, ventral infraciliature ofmor-
phogenetic stages after protargol impregnation). Middle stages showing primordia formation within the paren-
tal rows and differentiation of new cirri within the cirral streaks. The left frontal cirrus splits from the primor-
dium of the undulating membranes identically to in other hypotrichs (arrow). Bars = 90 JIm, 100 JIm. New
structures black, parental white; p 738.

1988b); starvation followed by intraspecific predation, however, induces cells within a

clone to split into two size classes: smallianceolate cells and cannibal giants. The same
behaviour was observed by KAMRA & SAPRA (1994) on the synonym, 0. indica.
Body shape in life and in growing cultures rather variable, outline elliptical to lan-
ceolate, sometimes distinctly truncated posteriorly. Many undersized individuals of very
variable shape develop in old cultures. Body stiff, only a little flexible under the cover
glass, conspicuously (2-3:1) flattened dorso-ventrally, especially the rims. Invariably 4 (n
= 50; FOISSNER et al. 1987b) prominent cone-shaped processes (horns, spines) on the dor-
sal surface (1 median hom in anterior half of body, 2 horns subterminally right and left of
median, and 1 smaller caudal hom; Fig. 195a, c, g, i, j, x) and 3 (n = 10) in Chlorogo-
 744 SYSTEMATIC SECTION

••
• •

• •
• •
•• ..
, ,"
• • •f •• • •
f •, •
•• ••
• f

'0

r
Fig. 195q, r Onychodromus quadricornutus (from FOISSNER et al. 1987b. q, r, ventral infraciliature of mor-
phogenetic stages after protargol impregnation). Late and very late stage showing positioning of new cirri, q =
350 llm, r = 345 llm. Arrow in (q) marks the 2 new dorsomarginal kineties of the proter. Large arrows in (r)
point to the anteriorly migrating rightmost ventral rows, which are thus very likely homologous to the migra-
tory cirri (cirri VY3 and VI/4) of the typical (l8-cirri) oxytrichids. The tiny arrow in (r) denotes the rest of the
fibrils of the parental oral apparatus. New cirri black, parental ones white; p 738.

nium-fed cells of the synonym, O. indica (KAMRA & SAPRA 1993); however, the synonym
also has a small elevation in the mid anterior region which becomes hom-shaped after
turning cannibal (Fig. 195x; KAMRA & SAPRA 1993). Horns immotile, tips sometimes bi-
furcated. Number of horns did not change with culture conditions and culture age; how-
ever, length and shape of spines is regulated by intraspecific predator-released morpho-
gens (WrcKLow 1990, 1991, 1993; see also FUNT et al. 1993). Induction experiments indi-
cate that a substance released by cannibal giants stimulates hom growth in clone- and
non-clonemates within 24 h (WrcKLow 1988b). Furthermore, O. quadricornutus cells ex-
 Onychodromus 745

I
\

\ \
\

t
Fig. 1955, t Onychodromus quadricornutus (from FOISsNER et al. 1987b. s, t, dorsal infraciliature and nuclear
apparatus of morphogenetic stages after protargol impregnation). Early and middle stage showing formation of
3 primordia each for the proter and the opisthe and multiple fragmentation of each primordium. Furthermore, 2
dorsomarginal kineties are formed in both filial products (Fig. 195q). Parental kineties have been omitted to
make details clearer (cp. Fig. 195v); P 738.

posed to the predacious ciliate Lembadion magnum also develop hypertrophied spines.
Selection experiments show that conspecific giants prey on cells with undeveloped spines
(<20 J,lm in length) to a much greater extent than on cells with developed spines (>40 J,lm
in length). Furthermore, WICKLOW (1988b) proposed that transformation of a popUlation
of similarly sized o. quadricornutus cells into 2 different size classes may function to in-
crease the range of potential prey sizes available to the 0. quadricornutus population; hy-
pertrophied spines appear to function as an inducible defence against intermittent preda-
tors appearing in the system, including conspecific giants (Fig. 195z).
Macronuclear nodules ellipsoid, connected by a fine strand, arranged like a question
mark left of median (Fig. 1959). Nodules do not increase in number during cannibalistic
growth, but become much larger through extra rounds of DNA replication without nu-
clear division (LIN & PRESCOTI 1986). Micronuclei about 5.~.5 J,lm across, most near
 746 SYSTEMATIC SECTION

Fig, 195u, v Onychodromus quadricomutus (from FOISSNER et a1. 1987b. U, v, scanning electron
micrographs). Ventral and dorsal view of late stages. The right parental marginal row and the rightmost paren-
tal ventral row are still unchanged. The oral apparatus of the proter shows clear symptoms of some reorganisa-
tion (cp. Fig. 195h), for example, lack of the cilia of the undulating membranes. The triangles in (v) mark the
developing new dorsal kineties, which originate by multiple fragmentation of single main streaks (cp. Fig.
195s, t). Bars = 100 11m; p 738.

macronucleus, some scattered in cytoplasm (ForSSNER et al. 1987b, KAMRA & SAPRA 1993,
LIN & PRESCOTT 1986; further literature on nuclear apparatus: HOFFMAN et al. 1995, PRES-
COTT 1994, SAPRA et al. 1985). About 4-6 contractile vacuoles along left and anterior
 Onychodromus 747

NC 1 "" "
NC 2 0 D
#I
II
II
0

" " "0

"
0 4)
11
/I (J 0
RV1 " 0 "
Q (I "
o 0 C o it
o ",
RV2 Q 0
" • ", D

0
g 0
" ~
""
0 (J

". " " <>

(I 0 II
RMC "
" ""
0 «I
II 0 Q
I'
"
0)
o 0
Q "

Fig. 195w, x Onychodromus quadricornutus (from KAMRA & SAPRA 1993. W, X, protargol impregnation).
Ventral and dorsal infraciliature, W = 230 11m, x = 215 11m. Arrowheads in (x) mark dorsal horns at the poste-
rior end of the cell and an inconspicuous elevation in the median of the anterior third. Explanation of original
labels: AZM = adoral zone of membranelles, CI- 7 = crowned frontal-ventral-transverse cirraI rows, CCI _l =
caudal cirri, ruM = endora!, L = left marginal row of cirri, NC1,2 = non-crowned frontal-ventral-transverse CiT-
ra! rows, GUM = parora!, RMC = right marginal row of cirri, RVI ,2 = right frontal-ventral-transverse cirraI
rows, Tcirri = transverse cirri, VI ,2 = pretransverse ventral cirri; p 738.

body margin (Fig. 195a). Pellicle very brittle, sharp-edged fragments can be broken out
by touching cell with a hair or fine needle. Cortical granules lacking. Cytoplasm colour-
less, filled with numerous greasily shining globules, 1-5 ~m across. No cytoplasmic crys-
tals. Movement strikingly slow and jerky, grazing along substratum, leaving it only with
great difficulty following vigorous shaking (ForSSNER et aI. 1987b, KAMRA & SAPRA
1993).
Adoral zone of membranelles almost 50 % of body length, posterior portion covered
by pellicle. Bases of largest membranelles in life about 25 ~m broad, details as in other
hypotrichs. Buccal area large, considerably deep, a small bulge at its right side. Undulat-
ing membranes straight and in parallel, cilia in life about 12 ~m long.
 748 SYSTEMATIC SECTION

0> 100
30 C
day 1 5 80
20 .~

10 ::J 60 "
til "
'" ,
Or---~=-------~~------- c 40
+-'
'" , ,
Q) ,
u ,
30 L 20
...... 20 day 2
Q)
0.... 0 t
t-
...... 10
0 4 8 12 16 20 24
>. Time (hrs) z
~ O~~~------------~~~~---­
v Fig. 195y, z Onychodromus quadricor-
5- 30 nutus (from WICKLOW I 988b). y) Cell
v
tt 20 day 3 length frequency distribution of 2 repli-
cates (black and white circles) of cells
10
cultured with abundant Chlorogonium
(day I) followed by 3 days of Chlorogo-
nium deprivation (days 2-4). z) Suscep-
30
tibility of different morphotypes to pre-
20 day 4 dation by cannibal giants. Populations of
cells with undeveloped spines, <20 J.lm
10
long (black and white triangle), were
preyed on to a much greater extent than
730 810 populations of cells with prominent
Y spines, >40 J.lm long (black and white
circle). P <0.001, i. Control popula-
tions of both morphotypes cultured with-
out giants remained at, or slightly above 100 % survival. At about 18 h (arrow), in the presence of giants, cells
initially with undeveloped spines attained spine length>30 J.lm, thereby decreasing susceptibility to predation;
p 738.

Cirral pattern rather invariably, on average 10-11 frontal-ventral-transverse cirral

rows, that is, no clustered cirri, except pretransverse and transverse cirri (Fig. 195e, h, w).
Cirral rows elongate from right to left, except for rightmost one, which is almost as long
as body. Thus, an inconspicuous suture is produced at right part of ventral ciliature field
(Fig. 195e). First 2-4 cirri of leftmost rows conspicuously enlarged ("crowned" rows in
Fig. 195w), about 25 J.1m long, form 2-3 arched "rows" along anterior body margin
(these arched rows are, however, not homologous to the corona of Kerona pediculus,
which originates only from one streak!). Marginal cirri in life about 20 J.1m long, those of
right row sometimes irregularly distributed. Invariably 2 slightly enlarged pretransverse
ventral cirri (n = 25; ForssNER et al. 1987b). Transverse cirri conspicuously enlarged, in
life only about 25 J.1m long, only posterior-most ones project slightly beyond posterior
end of cell, rather motionless during movement of ciliate.
Dorsal cilia in life about 4 J.1m long, arranged in many (>20) rows invariably includ-
ing 2 dorsomarginal kineties, most rows more or less shortened (Fig. 1959). This shorten-
 Onychodromus 749

ing and the small distances between rows prevent exact counting of kineties. All kineties
appear composed of densely packed basal body pairs, only anterior ones being ciliated.
Each pair surrounded by fibrils forming a rhomboid figure (Fig. 195f, x). Invariably 3
fine caudal cirri (n = 25, ForssNER et al. 1987b; n = 10, KAMRA & SAPRA 1993), left one
constantly inserted near tip of caudal hom (Fig. 195j; ForssNER et al. 1987b); conversely,
according to Figure 195x the posterior hom is situated between the middle and the right
caudal cirrus in the synonym, Onychodromus indica.

M 0 r p hog en e sis (Fig. 195k-v). Morphogenesis of cell division is both described

for the type material (ForssNER et al. 1987b) and the synonym, Onychodromus indica
(KAMRA & SAPRA 1993). The following paragraphs are from ForssNER et al. (l987b) un-
less otherwise indicated.
A few basal bodies develop at the left border of the first one to three transverse cirri,
which appear intact (Fig. 195k). Basal bodies increase in number, fonning a narrow patch
that extends anteriorly and reaches the proximity of the buccal vertex. The resulting oral
primordium is club-shaped, rounded anteriorly, and pointed posteriorly. About 50 % of
the dividing cells show 1 or 2 rounded clusters of basal bodies at the posterior end of the
oral primordium (Fig. 1951-n). From the anterior end of the oral primordium, 3 streaks
separate; the left one becomes the primordium of the undulating membranes, the middle
one the buccal row, and the right one develops the joining ventral row. The buccal row of
the proter is fonned by an anlage that develops near the parental row, whose cirri are re-
absorbed at the end of the division process (FOISSNER et al. 1987b). Conversely, in the
synonym 0. indica some buccal cirri are involved in the fonnation of primordium n (Fig.
4 and 5 in KAMRA & SAPRA 1993). Some parental cirri of the ventral rows disintegrate and
rearrange their basal bodies into the ciliary streaks appearing next, which average 13 for
each tomite (Fig. 195n, 0). Those of the proter evolve a little later than those of the opis-
the. On average there are 2 frontal-ventral-transverse anlagen more in the filial products
than ventral rows in the interphase individuals, suggesting that some reduction occurs in
later division phases. This reduction probably begins after the new transverse cirri have
been separated, because the number of transverse cirri of the non-dividers is on average
the same as the number of frontal-ventral-transverse cirral anlagen. No anlagen develop
within the two or three rightmost ventral rows, which remain unchanged during the whole
division process. These rows are produced by splitting of some of the rightmost frontal-
ventral-transverse cirral streaks in both the proter and the opisthe (Fig. 1950).
Development of cirral primordia is accomplished by growth, in both length and
width, of thin streaks and by posterior fragmentation in cirri, which progresses as an an-
teroposterior wave, transverse cirri being the last and largest, except for the frontal cirri
that derive from the approximately 8 leftmost cirral streaks (Fig. 195o-r). While this hap-
pens, the rest of the oral primordium differentiates into membranelles. Alignment of
membranelles starts at the anterior right edge of the primordium. The primordium of the
undulating membranes of the opisthe develops as a long anarchic field of basal bodies
parallel to the right border of the oral primordium and remains connected with streaks II
750 SYSTEMATIC SECTION

Table 45 Morphometric data of Onychodromus quadricornutus (1, spring water culture from FOISsNER et aI.
1987b; 2, from KAMRA & SAPRA 1993). All data are based on protargol-impregnated specimens. All measure-
ments in micrometres. CV = coefficient of variation (in %), Max = maximum value, mean = arithmetic mean,
Min = minimum value, n = sample size, SD = standard deviation

Character mean SD CV Min Max n

Body, length 11 272.5 48.5 17.8 170 361 50
12 423.3 112.8 26.7 222 694 50
2 205.3 24.3 11.8 180 226 10
Body, width 11 157.4 33.7 21.4 83 230 50
12 235.6 71.5 30.4 124 375 50
2 136.6 15.5 11.4 119 166 10
Macronucleus, number of nodules 19.6 4.8 24.7 11 28 25
2 11.4 1.3 11.1 10 13 10
Nuclear apparatus, length 192.4 34.6 18.0 143 270 25
Macronuclear nodule, length 18.3 4.8 26.2 11 28 25
Macronuclear nodule, width 1 9.2 2.0 22.2 6 14 25
Micronuclei, number 1 9.6 4.9 51.2 4 26 25
2 3.3 0.5 14.6 3 4 10
Micronucleus, length 5.2 0.7 13.3 4 6 25
23 6.4 0.5 8.3 6 7 20
Micronucleus, width 1 4.9 0.6 13.0 4 6 25
Adoral zone of membrane lies, 134.3 20.2 15.0 92 165 25
length 2 100.4 12.2 12.1 89 124 10
Adoral membranelles, number 130.5 21.7 16.6 89 180 25
2 82.0 8.1 9.8 66 98 25
Ventral cirral rows, number 10.6 1.3 12.4 8 13 25
C rows, number" 2 7.0 0.5 6.7 6 8 10
NC rows, number' 2 1.9 0.3 16.6 1 2 10
FVT cirri, number 2 147.8 7.2 4.9 136 159 10
Buccal row, number of cirri 5.6 1.2 20.5 3 8 25
2 5.0 0.7 13.4 4 6 25
C3 row, number of cirri4 2 7.1 0.7 9.9 6 8 25
RV I row, number of cirri4 2 15.4 1.4 8.8 13 18 10
RV2row, number of cirri4 2 18.8 1.6 8.6 16 21 10
Right marginal row, number of cirri 57.7 7.1 12.2 45 74 25
2 47.2 3.3 7.1 43 53 10
Left marginal row, number of cirri 43.5 5.3 12.3 34 54 25
2 35.4 2.2 6.3 32 39 10
Transverse cirri, number 1 12.2 1.5 12.7 9 16 25
2 10.0 0.7 7.3 9 12 25
Dorsal kineties, number I >20.0 ?
2 21.8 2.0 9.1 19 25 10

I From spring water culture.

2 From Pringsheim's solution culture.

3 Diameter.

4 For of simplicity, the original labelling of KAMRA & SAPRA (1993) is used. C and NC mearIS crowned and

non-crowned frontal-ventral-trarISverse cirri rows respectively (see Fig. 195w).

5 Including all cirri, except marginal and caudal cirri.
 Onychodromus 751

and III until the start of organisation of the endoral and the first (leftmost) frontal cirrus
(Fig. 195o--r). Thus, the frontal-ventral-transverse cirral system of the opisthe derives
from usually 13 primordia: three of these come from the oral primordium and originate
by proliferation and rearrangement of basal bodies from old ventral cirri. KAMRA & SAPRA
(1993) found the following origins of the frontal-ventral-transverse cirral primordia for
the proter (see Fig. 195w for designation of structures): primordium I from undulating
membranes, II from C2 (buccal row), III from C3, IV from C4, V from Cs, VJ-IXIXII from
or between C6-1!, NC 1 sometimes NC2• For the opisthe KAMRA & SAPRA (1993) found:
I-III from oral primordium, IV from C4, V from Cs, VI-IXlXII from or between ~,
NC 1 and often NC2•
The paroral and endoral, the buccal area, and some fibrillar structures of the proter
undergo a partial disorganisation. The basal bodies temporarily lose their linear arrange-
ment and later rearrange and reform the paroral and endoral and a new left frontal cirrus.
The buccal area appears smaller, less deep, and the fibrils in the peristomial vertex disap-
pear (Fig. 195q, r). The parental adoral zone of membrane lIes is inherited without anyap-
preciable changes. Thus, the frontal-ventral-transverse cirral system of the proter evolves
from 1 paroral primordium and, on average, 12 primordia which originate by prolifera-
tion and rearrangement of basal bodies from parental ventral cirri (Fig. 195o--r).
All cirral streaks of the proter develop without participation of the primordia of the
opisthe, as indicated by the high number of unchanged parental cirri and the large gap be-
tween the 2 anlagen fields in the early and middle stages of division (Fig. 195n-q).
New marginal cirri appear a little later than the frontal-ventral-transverse cirral anla-
gen and are built from streaks always formed close to the right of the old rows at 2 sites
on each side, one anterior and the other posterior to the future fission plane. No parental
marginal cirri are involved in this process, as indicated by the unchanged parental right
marginal row in the early and late division stages (Fig. 1950, q, r, u). This kind of mar-
ginal row formation is only known from the ectocommensal Kerona pediculus (HEM-
BERGER & WILBERT 1982; Fig. 208t). Furthermore, both species have the same, rather
complicated dorsal morphogenetic pattern, indicating a close relationship. Dorsal mor-
phogenesis commences with the formation of 3 primordia each in the proter and the opis-
the (Fig. 1955). Later, all these primordia show multiple fragmentation and each
posterior-most (rightmost) fragment bears a caudal cirrus (Fig. 195t, v). Furthermore, 2
dorsomarginal kineties are formed in each filial product (Fig. 195q, r).
All cirri and dorsal kineties which do not participate in morphogenesis disappear af-
ter cytokinesis. The dorsal horns develop at the end of the fission process. The macronu-
clear nodules fuse to a rod-like mass during middle stages. Later, this mass segregates
into the typical nodulated macronucleus of the interphase individuals (Fig. 195q, s, t).
Morphogenesis during reorganisation is similar to that of cell division, but, of course,
only one set of primordia is formed (KAMRA & SAPRA 1993).

o c cur r e n c eand e colo g y: Not very common in freshwater. Locus classicus

of Onychodromus quadricornutus unknown because the population was obtained from a
752 SYSTEMATIC SECTION

freshwater aquariwn at a local pet shop in Boulder, Colorado, USA, where tropical
aquariwn fish were sold, indicating that it is probably from some tropical climate (FOISS-
NER et al. 1987b, LIN & PRESCOTT 1985). CORLISS & PANG (unpublished) probably found
the same species in Shanghai, China (FOISSNER et al. 1987b, footnote 2; WICKLOW 1988b).
Locus classicus of the synonym, 0. indica, is Delhi, India, where KAMRA & SAPRA (1993)
discovered it in the Yamuna River at a point where a large amount of ash was discharged
by a thermal power station. Surprisingly, two records are from Asia, indicating that O.
quadricornutus is possibly confined to this continent. Unfortunately, the locus classicus
of O. quadricornutus is unknown, however, it cannot be excluded that the aquariwn was
filled with material from tropical Asia.
Feeds on algae, heterotrophic flagellates, ciliates, and wheat starch (FOISSNER et al.
1987b, KAMRA & SAPRA 1993). Cannibalistic when starved (FOISSNER et al. 1987b, LIN &
PRESCOTT 1985, WICKLOW 1988a, b). FOISSNER et al. (1987b) cultured O. quadricornutus
at room temperature in bottled spring water (Eau de Volvic, France) enriched with wheat
grains, small ciliates (Chilodonella uncinata, Cyclidium glaucoma, Tetrahymena thermo-
phila), and heterotrophic flagellates as food, and in Pringsheim's solution with Chlorogo-
nium elongatum as food; mediwn was changed every two weeks. KAMRA & SAPRA (1993)
cultured the synonym, O. indica successfully at about 23°C on mixed bacteria, Tetrahy-
mena thermophila, Stylonychia lemnae and on Chlorogonium elongatum. Generation
time of the synonym under optimal conditions 9 ± 0.5 h (± SD?; KAMRA & SAPRA 1993).
Onychodromus quadricornutus is one of the most volwninous hypotrichs at about
2 x 106 !lm3 (spring water culture) and 5 x 106 !lm3 (pringsheim's culture) and reaching
peak values of up to 20 x 106 !lm3 (FOISSNER et al. 1987b).

Insufficient redescriptions

Onychodromus grandis STEIN? - BORGER, 1908, An. Univ. Chile, 122: 186, Lamina XV,
Fig. 4. Remarks: In life 126!lm long, 7 macronuclear nodules. Feeds on diatoms. Fresh-
water lagoon of San Crist6bal, Chile.

Onychodromus grandis STEIN - CONN, Bull. Conn. St. geol. nat. Hist. Surv., 2: 59, Plate
XXVI, Fig. 247. Remarks: The illustration shows too many cirri to accept the identifica-
tion. The cirral rows and the body shape are somewhat reminiscent of Laurentiella.
Freshwater in Connecticut, USA.

Laurentiella DRAGESCO & NJINE, 1971

1966 Laurentia macrostoma DRAGESCO, Protistologica, 2: 85 - Type (original designation): Laurentia mac-
rostoma DRAGESCO, 1966.
1971 Laurentiella DRAGESCO & NJINE, Annis Fac. Sci. Univ. fed. Cameroun, 7-8: 125 (see nomenclature).
1972 Laurentia DRAGESCO, 1966 - BORROR, J. Protozool., 19: 14.
1974 Laurentia DRAGESCO - STILLER, Fauna Hung., lIS: 109.
 Laurentiella 753

1979 Laurentiella (Laurentiella) JANKOWSKI, Trudy zool. Inst., Leningr., 86: 57 - Type (original designation):
Laurentia macrostoma DRAGEsco, 1966.

C h a r act e r i sat ion: Undulating membranes in Stylonychia pattern. 5-7 frontal-

ventral-transverse cirral rows. One right and 1 left row of marginal cirri. Caudal cirri pre-
sent. Primordium II of proter originates from oral primordium. Dorsal kineties primordia
1 and 3 with multiple fragmentation, more than 2 dorsomarginal kineties.

Nom e n c I at u rea n d t a x 0 nom y: See relevant chapter in single species. For a

detailed description of the Stylonychia pattern mentioned in the characterisation, see
chapter 2 in the general section.

Single species

Laurentiella strenua (DINGFELDER, 1962) BERGER & FOISSNER, 1989

(Fig. 196a-v, Table 46)

1962 Paruro/eptus strenuus DINGFElDER, Arch. Protistenk., 105: 617 (Fig. 196a).
1966 Parauroleptus strenuus DINGFElDER, 1962 - MAns, Acta Fac. Rerum nat. Univ. comen., Bratisl., 13:
238 (Fig. 196b; incorrect subsequent spelling of Paruroleptus).
1966 Laurentia macrostoma DRAGESco, Protistologica, 2: 85 (Fig. 196c-e; new synonym).
1972 Laurentia macrostoma DRAGESCO, 1966 - BORROR, J. Protozool., 19: 14 (Fig. 196f; somewhat modified
redrawing from DRAGESCO 1966b).
1974 Paruroleptus strenuus DINGFElDER - STILLER, Fauna Hung., 1I5: 80.
1974 Laurentia macrostoma DRAGESCO - STILLER, Fauna Hung., 1I5: 1I0.
1975 Onychodromus grandis - JARENO, J. Protozool., 22: 82A (misidentification).
1976 Paruroleptus strenuus DINGFELDER, 1962 - CZAPIK & JORDAN, Acta Protozool., 15: 284 (Fig. I 96g).
1976 Laurentia acuminata FEDRIANI, MARTIN & PEREZ-SILVA, Boln R. Soc. esp. Hist. nat., 74: 67 (Fig. 196h,
i).
1977 Onychodromus acuminatus FEDRIANI et coIl. 1976 - JARENO, Protistologica, 13: 188.
1979 Laurentiella (Laurentiella) macrostoma (DRAGESCO, 1966) - JANKOWSKI, Trudy zool. Inst., Leningr., 86:
57.
1983 Laurentiella acuminata - MARTIN, FEDRIANI & PEREZ-SILVA, J. Protozool., 30: 519.
1987 Laurentiella acuminata FEDRIANI et aI. 1976 - SZABA & WILBERT, Debreceni Agrartudomanyi Egyetem
Tudomanyos KOzIemenyei, 27: 439 (Fig. 196j, k).
1989 Laurentiella strenua (DINGFELDER, 1962) - BERGER & FOISSNER, Bull. Br. Mus. nat. Hist. (Zool.), 55: 29
(Fig. 1961, m; morphometric data and line drawings of ventral and dorsal infraciliature of type popula-
tion of Laurentia acuminata, kindly supplied by J. MARTIN, University of Cordoba, Spain).

Nom e n c I a t u rea n d t a x 0 nom y: Laurentia acuminaba in FEDRlANI et al.

(1976b, p 73) is an incorrect original spelling. Incorrect subsequent spellings: Onyccho-
dromus acuminatus (PEREZ-PRIETO et al. 1981, p 290); Paruroleptus strenua (BERGER &
ForSSNER 1989a, p 29); strenuus DlNGFELDER (Puraropeltus) (STILLER 1974b, P 3).
Synonymy of Paruroleptus strenuus and Laurentia acuminata was already suggested
by HEMBERGER (1982, P 196) and BERGER & FOISSNER (1989a). Laurentia macrostoma is
754

'0 "
', ,, ,
. .I ':
.' :

.. . " ,' .,
0 ,' ,'
f:

.... : . : ! t
'0 ""

e
 Laurentiella 755

also evidently a further junior synonym of L. strenua, although there are some differences
between the three original descriptions, as in the number of macronucIear nodules and
cirral rows. However, both characters show a high variability (Table 46); furthermore,
DINGFELDER (1962) made only live observations, indicating that he underestimated the
number of cirral rows. Conversely, DRAGESCO (1966b) had only low quality protargol
slides and thus presumably also did not recognise the exact arrangement of the cirri (Fig.
196e). Furthermore, neither DRAGESCO (1966b) nor FEDRIANI et al. (1976b) compared
their species with P. strenuus, very likely because they overlooked DINGFELDER'S (1962)
paper. BORROR (1972a) also overlooked DINGFELDER'S paper and diagnosed Laurentia er-
roneously with 2 rows each of right and left marginal cirri.
Laurentia monilata DRAGESCO & NJINE, 1971 has the 18 frontal-ventral-transverse
cirral pattern of the "typical" oxytrichids and was thus transferred to a separate taxon,
Coniculostomum NJINE, 1979 (see there). Laurentiella strenua is possibly closely related
to Coniculostomum monilata and the Stylonychia mytilus complex because cirral streak
II of the proter originates from the oral primordium (Table 4) in all these taxa. Further-
more, the prominent oral apparatus is almost identical. This implies that in L. strenua the
increased number of cirri per streak, the multiple fragmentation of some dorsal primordia,
and the increased number of dorsomarginal kineties are apomorphies. Laurentiella stre-
nua can be easily separated in life and at low magnification from both, about equal sized,
taxa (Coniculostomum, Stylonychia) by the non-elongated caudal cirri, the pointed poste-
rior end of the cel~ and from the very common S. mytilus also by the increased number of
macronucIear nodules.
HEMBERGER (1982, p 196), in his unpublished thesis, transferred Paruroleptus
strenuus to Onychodromus (0. strenuus); however, Onychodromus has dorsal horns as
autapomorphy. Furthermore, in Onychodromus spp. streak II of the proter originates in-
dependently from the oral primordium (FOISSNER et al. 1987b, SZAB6 & WILBERT 1987).
Pattersoniella has curved and intersecting undulating membranes and a distinctly higher
number of cirral streaks which, however, segregate only 3 or 4 cirri each (Fig. 197).
The following chapter includes some unpublished observations kindly supplied by
W. ForssNER (Fig. 196n-v) and own observations, mainly concerning the live aspect.

M 0 r p hoi 0 g y and b i 0 log y: In life 100-500 ~m long: 103-326 ~m (DING-

FELDER 1962), 250-500 ~m (CZAPIK & JORDAN 1976b), 250-320 x 100-150 ~m (FEDRIANI
et al. 1976b), 250-320 x 100-140 ~m (SZAB6 & WILBERT 1987), 250-300 times
120-160 ~m (own observations), 300 x 160 ~m (W. FOISSNER, pers. comm.). Body stiff
as a board, almost triangular, that is, anterior end broad, posterior very narrowly rounded

+- Fig. 196a-£ Laurentiella strenua (a, from DINGFELDER 1962; b, from MATIS 1966; c-e, from DRAGESCO 1966b;
f, after DRAGESCO 1966b from BORROR 1972a. a, b, from life; c-f, protargol impregnation). a, b) Ventral view,
a = 280 11m, b = 250 11m. c, d) Variability of body shape and macronucleus. e, 1) Ventral infraciliature,
200 11m. In BORROR'S redrawing the arrangement of some cirri is changed, for example, in his figure (1) a dis-
tinct second left marginal row is recognisable which is unequivocally lacking in the original figure (e); p 753.
 756 SYSTEMATIC SECTION

...

k
Fig. 1969-k Laurentiella strenua (g, from CZAPIK & JORDAN 1976b; h, i, from FEDRIANI et al. 1976b;j, k, from
SzAB6 & WILBERT 1987. g-k, protargol impregnation). g, h, j, k) Ventral infraciliature, g = size not indicated, h
= 290 flm, j, k = 250 flm. Arrowheads in (h) mark a discontinuity in the rightmost frontal-ventral-transverse
cirral row. Arrows in (h, j) denote an enlarged cirrus which is very likely not a true transverse cirrus (that is,
the posterior-most cirrus of a cirral streak) but a pretransverse ventral cirrus, assuming that each cirral row pro-
duces 1 transverse cirrus (otherwise a "surplus" streak must have produced a transverse cirrus but no ventral
cirri, which is very unlikely). i) Somewhat deviating arrangement of transverse cirri in the population from the
"Parque del Retiro" (Madrid) of the synonym, Laurentia acuminata. 1-5 = numbering of frontal-ventral-
transverse cirral rows for morphometrical analysis (see Table 46; this numeration does not coincide with the
one usually used, that is, 1 corresponds to streak 11); p 753.
 Laurentiella 757

.. ...... • •, ~
~
.. ..
.. • , ~
. •• ~
... ...
~
~
... . ..' ~ ,

.
t' ,
, , ...
.....
• • •, .
~
..
..
I • ' III

.. , , "-
I • III
..
,
• It
.....

Fig. 1961, m Laurentie/Ia strenua (from BERGER & FOISSNER 1989a I, m, protargol impregnation). Ventral and
dorsal infraciliature (250 11m) of the type population of L. acuminata, drawn from slides kindly supplied by J.
MARTIN, University of Cordoba, Spain. Arrowhead marks caudal cirri. The dorsal infraciliature is rather com-
plicated in non-dividing specimens and originates by multiple fragmentation of2 dorsal kineties and formation
of 6 new dorsomarginal kineties. Possibly some parental basal body pairs are still retained after fission, as indi-
cated by the great distance between them (left margin of figure); p 753.

or tapered. Anterior end of cell distinctly turned dorsally, posterior quarter very strongly
flattened and thus translucent as in Stylonychia mytilus (Fig. 196n-r). Number of macro-
nuclear nodules rather variable (Table 46); according to FEDRIANI et al. (1976b) usually 4,
sometimes - possibly depending on physiological state - 5 or even up to 12 nodules pre-
sent in non-dividers; nodules ovoid to spherical, in life of rather different size, for exam-
ple, 20 x 15 /lm, 30 x 12 /lm, 15 x IS /lIn, 47 x 15 /lm, 38 x 20 /lm (own observations);
nodules usually arranged like the figure 7, according to FEDRIANI et al. (1976b) connected
by fine threadlike structures. Usually 7-9 globular micronuclei (FEDRIANI et al. 1976b),
according to DINGFELDER (1962) about 3.0-3.5 /lIn, according to own observations in life
4-5/lm across. For the macronuclear cycle stages the following approximate lengths
were estimated: G1 = 7.5 h, S = 1 h, division = 1 h, G2 = very short or lacking. Times for
micronuclear cycle: G1= 7.5 h, S = 1 h, division = 1h; G2 = absent. Total macronuclear
758 SYSTEMA TIC SECTION

Fig. 1%0, 0 Laurentiella strenua (n, 0, original scanning electron micrographs kindly supplied by W. FOIss-
NER). Ventral view showing the almost triangular outline, the giant oral apparatus with the straight paroral and
endoral (arrow), and the arrangement of the cirri; p 753.

DNA content per cell (average = 154.65 times micronucleus) found in cells with 4 mac-
ronuclear nodules was not significantly different from that found in cells with 5 nodules
(average = 157.93 times micronucleus; TORRES et al. 1979b). One contractile vacuole in
about mid-body, that is, about at level of buccal vertex (Fig. 196a, b), during diastole
with an anterior and posterior canal (W. FOISSNER, pers. comm.). Cortical granules lack-
ing. Cytoplasm usually dark at low magnification due to numerous cytoplasmic crystals.
Creeps on the substratum or swims like a moderately fast rotating board.
 Laurentiella 759

Adoral zone of membranelles about 50 % of body length, distal end distinctly over-
lapping on right body margin. Membranelles in life up to 22 /lm broad. Buccal area very
broad and deep (bowl-shaped), often with conspicuous fold (Fig. 196p), similar to in the
Stylonychia mytilus complex (Fig. 148i). Undulating membranes 120-180/lm long
(CZAPIK & JORDAN 1976b). Cilia of paroral in life about 25 /lm long, organised in slanting
pairs resulting in a zigzag pattern; endoral composed ofa single row of basal bodies with
postciliary microtubules (TORRES et al. 1986). Invariably 3 distinctly enlarged frontal cirri
(n = 11, BERGER & FOISSNER 1989a; n = 15, SZAB6 & WILBERT 1987), in life about 40 /lm
long (W. FOISSNER, pers. comm.). Buccal cirri (cirral row 1 in Fig. 196j) and cirri of cirral
row behind right frontal cirrus (row 2 in Fig. 196j) somewhat larger than cirri of remain-
ing frontoventral rows (MARTIN et al. 1983; Fig. 1961). Arrangement of frontal-ventral-
transverse cirral rows, see Figures 196h, j, I, o. Rightmost cirral row (= row 5 in Fig.
196j) often with a discontinuity (Fig. 196h), strongly indicating that it is homologous to
cirral row VI of the typical oxytrichids, where 2 cirri (= migratory cirri) are widely sepa-
rated from thepretransverse ventral cirrus (V1/2) and transverse cirrus (VIII). According
to DINGFELDER (1962) only 3 cirral rows (with 12-18 cirri each), which is, however, very
likely an underestimation due to live observation (Fig. 196a). The ventral infraciliature of
the synonym Laurentia macrostoma (Fig. 196e) shows either a somewhat deviating
specimen or includes some inaccuracies because of low quality impregnation. Transverse
cirri in life about 40-45 /lm long, slightly fringed, distinctly enlarged, arranged almost
longitudinally in median of cell, the anterior-most inserting rather close to buccal vertex
(Fig. 1961), and thus only posterior-most cirrus distinctly protruding beyond posterior end
of cell (Fig. 196a, n). Marginal cirri in life about 30 /lm long, marginal rows only slightly
separated posteriorly (Fig. 1961), right one running parallel to body margin, left one be-
gins close to buccal vertex and reaches left body margin asymptotically. Dorsal cilia short
(DINGFELDER 1962, FEDRIANI et al. 1976b), that is about 4/lm (own observations; W.
ForssNER, pers. comm.), only the anterior basal body of a pair is ciliated (TORRES et al.
1986). Arrangement of kineties and kinety fragments rather complicated due to mUltiple
fragmentation during morphogenesis (see below). Invariably 3 caudal cirri (BERGER &
FOISSNER 1989a, DINGFELDER 1962, FEDRIANI et al. 1976b, SZAB6 & WILBERT 1987), in life
about of same length (30 /lm) as marginal cirri and thus rather inconspicuous. The ultra-
structure of the oral apparatus and ventral and dorsal infraciliature shows only minor dif-
ferences to other hypotrichs (TORRES et al. 1986).
Resting cysts spherical, about 80-100/lm (85 ± 0.77 /lm [mean ± SD?], n = 123)
across, spines about 4-8 /lm long. Cyst belongs to the kinetosome-resorbing type (Fig.
196s-v). Cyst volume about 1/3 of vegetative cell, contains only one disc or band-shaped
macronucleus and several (3.6 ± 0.12 [mean ± SD?], n = 180) micronuclei which are
morphologically similar to the micronuclei of the vegetative cell (GUTIERREZ & PEREZ-
SILVA 1983b, GUTIERREZ et al. 1981; W. FOIssNER, pers. comm.). Resting cyst wall, as is
usual, composed of 4 layers; ectocyst about 0.3 /lm, mesocyst about 3.3 /lIn, and endo-
cyst about 0.4 /lm thick, granular layer of rather varying thickness (GUTIERREZ et al.
1983a). For detailed description of ultrastructure and chemical composition of cyst wall
 760 SYSTEMA TIC SECTION

Fig. 196p--r Laurenliella slrenua (p--r, original scanning electron micrographs kindly supplied by W. FOISS-
NER). p) The oral apparatus consists of a very prominent adoral wne of membranelles, straight undulating
membranes, and a very large and deep, bowl-shaped buccal area traversed by a conspicuous fold in scanning
electron microscopy preparations. q) Left lateral view showing the distinctly dorsally curved anterior part and
the very flat posterior quarter of the cell. r) Dorsolateral view showing the almost perpendicularly erected ante-
rior end of the cell and the complicated dorsal infraciliature; p 753.
 Laurentiella 761

see GUTIERREZ et al. (1980a, b, 1983a, 1984). Encystment is faster at 37°C (70-80 % af-
ter 6-10 h) than at 20 °C (10 % after 6-10 h). At about 4 °C and 45°C vegetative cells
failed to encyst and eventually died (GUTIERREZ et al. 1981).
M 0 r p hog e n e sis of cell division, reorganisation, and regeneration are de-
scribed by MARTIN (1982) and MARTIN et al. (1983), however, no line drawings are given.
Morphogenesis commences in usual manner in that some basal bodies appear immedi-
ately left of the left transverse cirrus (= anterior-most one). Number of basal bodies in-
creases, forming a club-shaped oral primordium. From its anterior end, a group of basal
bodies diverges and parallels the right peristomial margin and fmally segregates as a line
of basal bodies, the anterior part of which, after separating, becomes the ciliary streak II
of the proter. 1bis matches Coniculostomum and the Stylonychia mytilus complex (see
Table 4 and taxonomy). The posterior part becomes the ciliary streak II of the opisthe
and two more primordial streaks arise, one at each side of its enlarged base: the left one
becomes the primordium of the undulating membranes, the right one becomes the ciliary
streak III of the opisthe. Some parental cirri disintegrate and rearrange their basal bodies
into the ciliary streaks appearing next, usually 3 for each tomite. Some of these streaks
initially are continuous with those corresponding to the other tomite. Sometimes more
than 6 streaks develop. Frequently no transverse cirri or no ventral cirri arise from these
"surplus" primordia, explaining the variation and discordance between number of trans-
verse cirri and number of cirral rows. The frontal-ventral-transverse streaks of the opisthe
usually derive from 6 cirral primordia; three of them coming from the oral primordium
and 3 (sometimes more) originate by proliferation and rearrangement of basal bodies
from parental ventral cirri.
The parental adoral zone of membranelles and the endoral are inherited by the proter
without any appreciable change. The paroral undergoes a partial disorganisation of its an-
terior tract; the basal bodies of this tract temporarily lose their linear arrangement and,
later on, rearrange and reform the paroral and a new left frontal cirrus (cirrus Ill).
The frontal-ventral-transverse cirral streak II of the proter is the anterior part of the
row of basal bodies extending from the oral primordium. Basal bodies of the remaining
streaks result, as in the opisthe, from disintegration of parental frontal cirri. These primor-
dia are also arranged in a variable number (usually 6) of parallel streaks.
FOISSNER et al. (1987b, p 157, footnote 3) studied protargol-impregnated specimens
of the synonym Laurentia acuminata (supplied by J. MARTIN, Spain). The reinvestigation
clearly showed multiple fragmentation of dorsal primordia. Dorsal kinety primordium 1
(the primordium closest to the left marginal row) fragments into several pieces as in Ony-
chodromus quadricornutus, the middle primordium does not fragment, and primordium 3
has a simple fragmentation, like kinety 3 in the Oxytricha pattern. Furthermore, the poste-
rior fragment ofkinety 3 (= "kinety 4" in, for example, the Oxytricha pattern) shows some
multiple fragmentation at its anterior end. Six dorsomarginal kineties complete dorsal mor-
phogenesis. Caudal cirri originate in principle as, for example, in the Oxytricha pattern
(Fig. 24a), however, in non-dividing specimens their origin is not recognisable (Fig.
196m). All parental frontal-ventral-transverse cirri. marginal cirri. and dorsal kineties
 762 SYSTEMA TIC SECTION

Fig. 196s-v Laurentiella strenua (s-v, originals kindly supplied by W. FOISSNER. S, t, from life; u, v, scanning
electron micrographs). s) Young cyst with still smooth ectocyst. t, v) Old cysts with long spines. Arrowhead
marks fused macronucleus. u) Middle-aged cyst; p 753.

which do not participate in morphogenesis disappear after cytokinesis (MARTIN et al.

1983).
Morphogenesis during excystment is described by JARENO & TUFFRAU (1979). JARENO
(1977, 1981b) described a process referred to as joint encystment, that is, encystment
takes place after pairing of two cells, whereby cytoplasm fuses in a manner similar to to-
tal conjugation. Nevertheless, this process is not a sexual one since it lacks macronuclear
degeneration and neither meiosis nor caryogamy could be detected. Excystment generates
doublets which are capable of fission: the anterior part of a doublet gives rise:to 2 single
cells, whereas the posterior part gives rise to 1 doublet. The singlet condition is recovered
 Laurentiella 763

after a varying number of divisions. Joint encystment may be held as one step in a life-
cycle because it generates very vigorous lines and a strain is maintained without any
symptom of senility through an alternation of growth periods and starvation periods dur-
ing which joint encystment occurs. Further literature on the synonym, Laurentia acumi-
nata, mainly concerning the cyst: GUTIERREZ et al. (1980c--e, 1982a, b, 1983b), JARENO
(1979, 1980, 1981a, 1984, 1985, 1987b, c, 1988, 1992), JARENO & TUFFRAU (1978),
NooucIll & MATSUSAKA (1989), TORRES et al. (1979a, b, 1982).

o c cur r e n c e and e colo g y: Common in temporary waters. According to DING-

FEWER (1962) beside Colpoda magna, Nassula plantiformis, and Maryna cardioides a
key species for track and meadow puddles.
Locus classicus is a lane south of the city Forchheim, Bavaria, Germany, where
DINGFEWER (1962) discovered Laurentiella strenua in puddles with hay. He also found it
in other track and meadow puddles, especially in putrid sites at pH 6.8-7.4. Locus classi-
cus of the synonym Laurentia macrostoma is a small saprobic puddle near the Lake of
Geneva, Thonon-Ies-Bains, France, where DRAGESCO (1966b) found it in summer. Locus
classicus of the synonym Laurentia acuminata is Seville, Spain, where FEDRIANI et al.
(1976b) discovered it in a sample of water from the "Parque de M." Luisa". They also
found it in samples of water collected in the "Estaci6n Biol6gica de Donana", Huelva,
Spain, and in the "Parque del Retiro" in Madrid.
Records substantiated by illustrations: eutrophic meadow puddle with cyanobacteria
near a lake (Seewaldsee) in Salzburg, Austria (own observations); abundantly in some
meadow pools in S1. Georg-Moorwald near Bratislava, Czechoslovakia, in November
(MATIS 1966); in the plant community "Agrosti-Bachmannietum erucaeformis" in the
Hortobagy National Park, Hungary, together with the ciliates Colpoda cucullus, Ble-
pharisma lateritium, B. steini, Dileptus anser, Halteria grandinella, Strombidium sulca-
tum (SZAB6 & WILBERT 1987); very abundantly in September at 8-12 °C in a small
meadow puddle near the Institute of Zoology of the University Jagiellone, Krakow, Po-
land (CZAPIK & JORDAN 1976b); desiccated brook in Hawaii, USA (W. ForssNER, pers.
comm.); Australia (W. FOISSNER pers. comm.; Fig. 196n-v).
Records not substantiated by illustrations: with about 2 % frequency in submerged
mosses from standing and running waters in Czechoslovakia (TIRJAKOVA & MATIS 1987a);
with 5.9 % frequency in paddy fields in Japan (TAKAHASIll & SUHAMA 1991). SZAB6 &
WILBERT (1987) stated, obviously erroneously, that Laurentia acuminata is known from
water samples in Cameroon (very likely confused with L. monilata, now Coniculo-
stomum monilata).
Feeds on bacteria, cyanobacteria, small algae, diatoms, flagellates, and ciliates like
Halteria and Tetrahymena (CZAPIK & JORDAN 1976b, DINGFEWER 1962, own
observations). FEDRIANI et al. (1976b) and MARTIN et al. (1983) cultivated it in Pringsheim
medium at about 20°C and fed it daily with Colpidium sp., grown in the same medium
with integral flour added and previously filtered, or with Chlorogonium sp. that had been
concentrated by centrifugation. W. FOISSNER (pers. comm.) cultivated Laurentiella Stre-
764 SYSTEMATIC SECTION

Table 46 Morphometric data of Laurentiella strenua (1, from DINGFELDER 1962; 2, from DRAGESCO 1966b; 3,
from CZAPIK & JORDAN 1976b; 4, from FEDRIANI et aI. 1976b; 5, from SzAB6 & WILBERT 1987; 6, type popula-
tion of the synonym Laurentia acuminata from BERGER & FOISSNER 1989a). All data are based on protargol-
impregnated specimens unless otherwise indicated. All measurements in micrometres. ? = sample size not indi-
cated; if only 1 value is known it is listed in the mean column, if 2 values are available they are listed as Min
and Max. CV = coefficient of variation (in %), Max = maximum value, mean = arithmetic mean, Min = mini-
mum value, n = sample size, SD = standard deviation

Character mean SD CV Min Max n

Body, length I' 103 326 ?
2 200.0 ?
3' 250 500 ?
4 250 320 ?
5 288.5 21.0 7.3 250 320 15
6 268.6 28.7 10.7 240 325 11
Body, width 4 100 150 ?
5 122.4 12.0 9.8 100 140 15
6 159.6 16.6 10.4 150 196 11
Macronucleus, number of nodules 8.0 ?
2 3 6 ?
3 4.0 ?
4 4.0 ?
5 5.9 1.1 18.6 5 7 15
6 4.6 1.8 38.6 2 7 11
Macronuclear nodule, diameter I' 20.0 ?
66 36.1 11.7 32.5 20 56 11
61 22.4 6.6 29.3 II 32 II
Micronuclei, number 1 8.0 ?
2 3 6 ?
4 3 21 ?
5 5.9 1.1 18.6 5 7 15
6 7.5 7.1 94.7 2 24 11
Adoral zone of membranelles, length 3' 180 200 ?
6 134.3 9.5 7.1 120 150 11
Adoral membranelles, number 1 50 60 ?
2 50 60 ?
3 60 70 ?
4 60.0 ?
5 63.5 1.7 2.7 62 66 15
6 65.8 1.9 2.9 64 70 11
CirraI row 1, number of cirri2 44 5.0 4 6 ?
55 3.5 0.5 14.3 3 4 15
CirraI row 2, number of cirrj2 44 7.0 6 7 ?
55 5.0 0 0 5 5 ?
Cirral row 3, number of cirri 2 4 7 8 ?
5 10.0 0 0 10 10 15
Cirral row 4, number of cirrj2 4 6 8 ?
5 10.0 0 0 10 10 15
Cirral row 5, number of cirri 2 43 6 11 ?
5 14.0 0 0 14 14 15
 Laurentiella 765

Table 46 Continued

Character mean SO CV Min Max n

Right marginal row, number of cirri 41.0 ?
2 36 44 ?
3 40 44 ?
4 40 45 ?
5 45.8 1.2 2.6 40 49 15
6 39.7 1.9 4.8 37 42 11
Left marginal row, number of cirri 1 20 23 ?
2 20 23 ?
3 22 23 ?
4 26 30 ?
5 27.2 0.5 1.8 25 28 15
6 28.8 1.7 5.8 27 32 II
Transverse cirri, number 5 6 ?
2 6 10 ?
3 2 6 ?
4 5 6 ?
5 6.0 0 0 6 6 15
6 5.6 0.8 14.4 5 7 11

1 From life.
2 For numbering of cirral rows, see Figure 196j.
3 Anterior part with 3-5, posterior one with 3-6 cirri (see Fig. 196h).

• Anterior-most enlarged cirrus (frontal cirrus) included.

S Anterior-most enlarged cirrus (frontal cirrus) not included.

6 Posterior macronuclear nodule, length.

7 Posterior macronuclear nodule, width.

nua in Eau de Volvic with Tetrahymena rostrata as food. The cell cycle of L. strenua re-
quires 9.5 ± 1 h (mean ± SD?) at about20 °C (TORRES et al. 1979b, 1980b).
Laurentiella strenua captured the vaccine virus if it was added to the culture
medium. The medium was completely decontaminated and the virus was inactivated in
the ciliate; however, in the cytoplasm a proliferation of membranous cisternae occurred
2 h after culture inoculation by vaccine virus (JARENO 1987a, JARENO et al. 1985, 1986,
PEREZ-PRIETO et al. 1981, 1983).
MARTIN et al. (1983) induced full reorganisation of the ciliature by (i) starvation for
more than 24 h, after which cells ceased motion, sank, altered their shape, and began
physiological reorganisation, (ii) treating starved cells (having no food in cytoplasm) with
urea (3 % [w/v] fmal concentration) for 30 s, causing lysis of most of the adoral zone of
membranelIes and subsequent synchronous regeneration, and (iii) micro-surgical removal
of a cell fragment.
DINGFELDER (1962) observed L. strenua in track puddles at temperatures of up to
43°C. SZAB6 & WILBERT (1987) found it in water of the "B-limno, Na-HC03-S04 type" at
766 SYSTEMATIC SECTION

20.5 DC, pH 7.2-7.6,6.5 mg 1- 1 0 2 (70.6 % saturation), and 0.075 % salinity. Small doses
ofUV-radiation (about 240 and 360 J m-2) produced a considerable increase of the repro-
ductive rate, that is, the growth constant was about 11-16 % higher than in the control
(GUTIERREZ & PEREZ-SILVA 1980, 1983a); LDso was between 720 and 1200 J m-2 and
LDlOo was about 2400 J m-2 (mean time of death was 95 h). About 88 % of the resting
cysts obtained from vegetative cells irradiated with doses of 720-960 J m-2 lacked the
characteristic spines and their cyst walls were thicker (6.5 Jlm) than those of the normal
cysts (3.7 Jlm).

Pattersoniella FOIssNER, 1987

1987 Pattersoniella FOISSNER, Zoo!. Beitr., 31: 207 - Type (original designation): Pattersoniella vitiphila
FOISSNER, 1987.

C h a r act e r i sat ion: Undulating ± membranes in Oxytricha pattern. Number of

frontal-ventral-transverse cirri distinctly increased. One right and 1 left row of marginal
cirri, almost confluent posteriorly. More than 6 dorsal kineties. Caudal cirri present.
Number of frontal-ventral-transverse cirral streaks distinctly increased (9-11 against 6 in
''typical'' oxytrichids); however, only 3 or 4 cirri originate per streak. Dorsal morpho-
genesis in Oxytricha pattern; however, kinety 3 not with simple, but with multiple frag-
mentation and very likely some parental kineties retained after division.

T a x 0 nom y: See single species. For a detailed description of the Oxytricha patterns
mentioned in the characterisation, see chapter 2 in the general section.

Single species

Pattersoniella vitiphila FOIssNER, 1987 (Fig. 197a-u, Table 47)

1987 Pattersonie/la vitiphi/a FOISSNER, Zoo!. Beitr., 31: 207 (Fig. 197a-s; type slides are deposited in the
OberOsterreichische Landesmuseum in Linz, Upper Austria).
1994 Pattersonie/la vitiphi/a - FOISSNER, Kataloge des O. O. Landesmuseums, Linz, 71: 192 (Fig. 197t).

T a x 0 nom y: The ventral cirral pattern, especially the "bicorona" and the ventral cirri
which are arranged in pairs, is reminiscent of the holostichids (Fig. 1979). However, frag-
mentation of dorsal kinety 3 assigns Pattersoniella unequivocally to the Oxytrichidae
(Fig. 197q). Furthermore, morphogenesis of the ventral infraciliature proceeds rather
similarly to in typical (I8-cirri) oxytrichids, that is, the frontal-ventral-transverse cirri
originate from longitudinal streaks (Fig. 197k, I); FOISSNER (l987e) hence classified it in
the superfamily Oxytrichoidea. The rigid cortex, the lack of cortical granules, and the
large oral apparatus (about 50 % of body length) assigns Pattersoniella to the Stylonychi-
 P attersoniella 767

b c

Fig. 197a--e Pattersoniella vitiphila (a-d, from FOISSNER 1987e; e, original kindly supplied by W. FOISSNER.
a--e, from life). a) Ventral view ofa nonnal specimen, 200 11m. b) Ventral view ofa very large specimen (mac-
rostome fonn, 390 11m) showing the oral apparatus and the contractile vacuoles. c) Right lateral view showing
convex dorsal and concave ventral side. d, e) Resting cysts, diameter about 50 11m. (d) from type material, (e)
from Australian population. The macronucleus is fused in the resting cyst (W. FOISSNER, pers. comm.) and not
fragmented in nodules as shown in (d); p 766.
768 SYSTEMATIC SECTION

#
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 Pattersoniella 769

nae (Fig. 25a), implying that the number of frontal-ventral-transverse streaks has secon-
darily increased (Fig. 197u). However, the relationship to the other members of this
group are uncertain. Pattersoniella differs from Onychodromus spp. and Laurentiella
strenua, among other things, in that the number of cirri originating per streak is not in-
creased and in the shape of the undulating membranes (Oxytricha against Stylonychia
pattern). Territricha stramenticola (Fig. 219), Holosticha camerounensis DRAGESCO,
1970, and H. macrostoma (DRAGESCO, 1970) BORROR, 1972 have only 3 distinctlyen-
larged frontal cirri, 2 macronuclear nodules, and 1 contractile vacuole.

M 0 r ph 0 log y and b i 0 log y: In life 140-330 x 70-150 J.1m (FOIsSNER 1987e).

Australian (Fig. 197e, t) population in life 300-350 x 125-140 J.1m. Outline broadly ellip-
tical, left margin usually more convex than right, often distinctly narrowed at level of
buccal vertex, both ends broadly rounded. Body firm, flattened about 2:1 dorsa-ventrally,
ventral side slightly concave, dorsal side distinctly convex (Fig. 197c, t). In cultures
FOISSNER (1987e) occasionally found very large specimens with a very expansive oral ap-
paratus and a rather high number of ventral cirri (Fig. 197h; Table 47). This macrostome
form shows all transitions to the normal form and was not observed in raw cultures. The
following description refers only to the normal form. Macronuclear nodules ellipsoid, in
life about 20 J.1m large, arranged in median or slightly left of it, nodules with many small
chromatin bodies. Micronuclei spherical to ellipsoid, located close to macronuclear nod-
ules. Several contractile vacuoles near left body margin and in buccal area (Fig. 197b),
during diastole connected by canals. Cytopyge at posterior end of cell, slightly right of
median. Pellicle rigid, shining, cortical granules lacking. Endoplasm colourless, coarsely
granulated, in well-fed specimens dark at low magnification due to many 7-40 J.1m sized
food vacuoles and 2-4 J.1m sized, yellowish crystals. Movement rapidly gliding, clumsily
swimming like a rotating board.
Adoral zone of membranelIes about 50 % of body length, formed distinctly like a
question mark, widely overlapping on right body margin and thus only slightly separate
from anterior end of right marginal row. Bases of largest membranelles in life about
17-25 J.1m broad. Pharyngeal fibres short, in life inconspicuous. Buccal area broad but
flat (Fig. 197t). Undulating membranes curved and intersecting, middle section of paroral
consists of short kineties with 3 basal bodies each; cilia of undulating membranes in life
about 25 J.1m long. Frontal cirri (usually 6) and cirri right of buccal cirrus (usually 4) dis-
tinctly enlarged and thus very prominent in life, forming a so called "bicorona", frontal
cirri in life 25-30 J.1m long (Fig. 197a, t). Usually 6 frontoventral cirri, namely 2 migra-

~ Fig. 197C-i Pattersoniella vitiphila (from FOISSNER 1987e. f-i, protargol impregnation). C, g) Ventral and dor-
sal infraciliature of a normal specimen, 240 11m. See Figure 197u for detailed explanation of ventral ciliature.
Arrows in (t) mark cirri of the anterior and posterior corona, together forming the bicorona Arrowhead in (g)
marks a kinety with widely spaced basal body pairs, indicating that it is a retained parental kinety. h) Ventral
infraciliature ofa "macrostome" specimen, 370 11m. i) Ventral infraciliature of the posterior part ofa very early
morphogenetic stage, bar = 17 11m. Stomatogenesis commences with formation of basal bodies immediately
left of the anterior-most transverse cirri (arrows); p 766.
770 SYSTEMATIC SECTION

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197J k

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 Pattersoniella 771

tory cirri near anterior end of right marginal row and 2 pairs of cirri. Three postoral ven-
tral and 2 pretransverse ventral cirri, identical to, for instance, in Histriculus or Sterkiella;
ventral cirri in life about 20 Ilm long. Transverse cirri in life 25-40 Ilm long, fringed, dis-
tinctly enlarged, only posterior-most reach posterior end of cell (Fig. 197a, t). Marginal

. .
0 0
Q ....
..:.. -
Q :-
Q 0
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t:lQ '" 0 0
:a
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C:J

Fig. 197n, 0 Patlersoniella vitiphila (from FOISSNER 1987e. n, 0, ventral infraciliature of morphogenetic stages
after protargol impregnation). Late and very late stage, n = 185 11m, 0 = 265 11m. Arrows mark dorsomarginal
kineties originating at anterior end of right marginal primordia. Cirri originating from the same streak are con-
nected by a broken line. New structures black, parental ones white; p 766.

+- Fig. 197j-m Patlersoniella vitiphila (from FOISSNER 1987e. j-m, ventral infraciliature of morphogenetic
stages after protargol impregnation). j) Very early stage, bar = 55 11m. The figure shows a specimen with an in-
creased number of postoral ventral cirri (cp. Fig. 197f). Arrows mark newly formed basal bodies immediately
left of left postoral ventral cirri. k) Early stage, 200 11m. Arrowhead marks migratory cirri, that is, the anterior-
most 2 cirri of the rightmost streak which are never involved in primordia formation. Arrow denotes posterior
postoral ventral cirrus which also does not participate in primordia formation in the Stylonychinae. I, m) Mid-
dle and late stage, I = 265 11m, 1= 230 11m. Arrow in (I) marks posterior postoral ventral cirrus which is not in-
volved in primordia formation. Primordia of proter and opisthe originate independently. Note the complete re-
organisation of the parental undulating membranes; p 766.
772 SYSTEMATIC SECTION

.",

, '

, '

'..,' q
197p

r
 Pattersoniella 773

cirri in life about 20-25 !lm long. Dorsal cilia in life 3 !lm long, arranged in 4 kineties of
body length and in several short kineties (see morphogenesis for further details); alto-
gether about 15 kineties on average. Invariably (n = 28) 3 caudal cirri, arranged optically
rather exactly in the gap between the marginal rows, marginal and caudal cirri thus form-
ing a rather homogeneous seam (Fig. 197a, t).
Resting cyst about 50 !lm across (Table 47). Wall about 2-3 !lm thick, finely folded
(Fig. 197d, e). Macronucleus fuSed (W. FOISSNER, pers. comm.) and not divided in nod-
ules as shown in Figure 197d. Micronuclei not fused.
M 0 r ph 0 g e n e sis commences with the proliferation of basal bodies very close
to the leftmost (= anterior-most) transverse cirri which are retained unchanged (Fig. 197i;
FOISSNER 1987e). Number of basal bodies increases to form a narrow oral primordium.
The anterior postoral ventral cirri are modified to primordia (Fig. 197j-I), that is, the
posterior-most postoral ventral cirrus is not involved in primordia formation as this is
typical for all Stylonychinae! Simultaneously the cirri of the posterior corona (cirri left of
buccal cirrus) and the frontoventral cirri (except migratory ones) are involved in primor-
dia formation of the proter (Fig. 197k-m). Both in the proter and in the opisthe, some pri-
mordia split so that altogether 9-11 streaks each occur, including streak I, which forms
the undulating membranes (Fig. 1971-n). The primordia of the proter and the opisthe
originate independently, whereas, for example, in Laurentiella strenua streak II of the
proter originates from the oral primordium. In middle stages the parental undulating
membranes are completely reorganised, including the formation of the left frontal cirrus
(cirrus Ill; Fig. 1971-n). Parental pharyngeal fibres are reabsorbed and new ones appear
immediately prior to cytokinesis. The 9-11 frontal-ventral-transverse cirral streaks segre-
gate the following number of cirri (from 6 specimens): 1, 3, 3, 3, 3, 3, 3, 4, 4 (3 times); 1,
3, 3, 3, 3, 3, 4, 4; 1, 3, 3, 3, 3, 3, 4, 4, 4; 1, 3, 3, 3, 3, 3, 3, 3, 4, 4. This pattern agrees
largely with the well known pattern in typical oxytrichids having 18 frontal-ventral-
transverse cirri, except that 3 surplus streaks (compared with the most common case hav-
ing 9 streaks) with each 3 cirri are formed. I thus have no doubt that Pattersoniella is a
Stylonychinae with a secondarily increased number of streaks (Fig. 197u). Morphogene-
sis of marginal rows proceeds as usual (Fig. 1971-0).
Dorsal morphogenesis proceeds primarily as in the Oxytricha pattern (Fig. 24a) with,
however, important modifications. Primordia formation occurs in kineties which corre-
spond to the kineties 1-3 of the Oxytricha pattern (Fig. 197p). Later "kinety 3" divides
not by simple, but by multiple fragmentation (Fig. 197q-s), similarly to in Laurentiella
strenua, Onychodromus quadricornutus, and Territricha stramenticola. Furthermore, in-
variably 2 dorsomarginal kineties occur (Fig. 197n, 0, s). Consequently, usually about 9
kineties and kinety fragments are newly formed. However, most non-dividing specimens

+- Fig. 197p-s Pattersoniella vitiphila (from FOISSNER 1987e. p-s, dorsal infraciliature of morphogenetic stages
after protargol impregnation). Early, middle, and late stage, p = 175 11m, q = 182 11m, r = 260 11m, S = 265 Ilffi·
Arrows in (q) mark beginning mUltiple fragmentation in kinety 3. Arrows in (s) mark dorsomarginal kineties,
arrowheads denote new caudal cirri. Detailed description, see text; p 766.
774 SYSTEMATIC SECTION

Fig. 197t Pattersoniel/a vitiphi/a (from FOISSNER 1994a Scanning electron micrograph; Australian popu-
lation). Ventral infraciliature ofa specimen with a slightly reduced number offrontoventral cirri, 165 J-lm. Note
the short (3 J-lm) dorsal cilia on the left body margin; p 766.
 Pattersoniella 775

Fig. 197u Pattersoniella vitiphila (same as Fig. 1971';

slightly modified). Ventra1 infraciliature. This figure illus-
trates the origin of Pattersoniella from "typical" oxytri-
chids with 18 frontal-ventral-transverse cirri (cp. Fig. 6a).
These 18 cirri, originating from 6 streaks, are connected
by broken lines, that is, the streaks I-VI of Figure 6a very
likely correspond to streaks 1, 2, 6, 8, 9, 10 of Patterso-
niella. Additional cirri are coruiected by dotted lines; cor-
responding additional transverse cirri are marked with ar-
rows. The anterior-most 2 cirri of streak 10 are the migra-
tory cirri, the anterior-most cirri of streaks 1--6 form the
anterior corona (= frontal cirri), the middle cirri of streaks
2-5 form the posterior corona (both coronas form the "bi-
corona" of Pattersoniella). 1-10 = frontal-ventral-trans-
verse cirral streaks; p 766.

have about 15 kineties (Fig. 1979). This can be

explained only in that parental kinety frag-
ments are retained after fission, which is also
indicated by the widely spaced basal body
pairs (Fig. 1979). Retention of parental kine-
ties is also known in Coniculostomum. Caudal
cirri occur, identically to in the Oxytricha pat-
tern, at the posterior end of kineties 1 and 2
and the posterior-most part originating from
the multiple fragmentation of kinety 3 (Fig.
197s).
The nuclear apparatus of Pattersoniella vi-
tiphila shows no peculiarities, that is, in the
middle stages of morphogenesis the macronu-
clear nodules fuse and divide in later stages
(Fig. 197p-s).

o c cur r e n c e and e colo g y: Possibly confined to terrestrial habitats of the

southern hemisphere. Locus classicus is Suva on the Fiji Island Viti Levu, South Pacific
Ocean, where FOISSNER (1987e) discovered Pattersoniella vitiphila in the soil (PH 5.4;
collector D. J. PATIERSON) of the rain forest. FOISSNER (1987e) cultivated it in Eau de Vol-
vic; some salad medium and squeezed wheat grains were added to support growth ofbac-
teria and small protozoa. W. ForssNER (pers. comm.) found P. vitiphila in an Australian
soil (Fig. 197e, t).
Feeds on a wide variety of food like heterotrophic flagellates, testaceans (Trinema
Iineare), green algae, soil particles, ciliates (Co/poda spp., Tetrahymena pyriformis), fun-
gal spores and hyphae. In cultures even starch and Paramecium aurelia were ingested.
Biomass of 106 specimens about 700 mg.
776 SYSTEMATIC SECTION

Table 47 Morphometric data of Pattersoniella vitiphila (from FOISSNER 1987e. line I, nonnal specimens; line
2, giant specimens, that is, macrostome fonn). All data are based on protargol-impregnated specimens unless
otherwise indicated. All measurements in micrometres. CV = coefficient of variation (in %), Max = maximum
value, mean = arithmetic mean, Min = minimum value, n = sample size, SD = standard deviation

Character mean SD CV Min Max n

Body, length I 230.5 31.0 13.5 170 295 25
2 348.0 336 364 3
Body, width 108.8 16.6 15.3 70 140 25
2 161.7 150 180 3
Nuclear apparatus, length 1 172.4 26.5 15.4 120 230 25
2 255.0 240 280 3
Macronucleus, number of nodules 15.6 1.1 7.2 13 18 25
2 17.3 16 19 3
Macronuclear nodule, length 17.4 4.3 24.6 11 27 25
2 28.0 28 28 3
Macronuclear nodule, width 1 13.2 2.1 16.1 9 20 25
2 20.3 17 24 3
Micronuclei, number 1 6.0 3.0 50.0 2 12 25
2 17.7 15 21 3
Micronucleus, diameter 1 4.1 0.5 12.3 3.5 5.6 25
2 4.7 4 6 3
Adoral zone of membranelles, length 107.4 21.1 19.6 70 140 25
2 197.0 190 205 3
Adoral membranelles, number I 60.4 8.4 14.0 44 83 25
2 105.7 98 119 3
Frontal cirri, number2 I 5.8 0.7 1l.5 5 8 25
2 7.3 7 8 3
Ventral cirri, numbecl 14.6 1.2 8.2 13 18 25
2 18.7 18 20 3
Right marginal row, number of cirri 1 31.5 2.5 8.0 27 36 25
2 41.0 36 47 3
Left marginal row, number of cirri 29.1 2.9 9.9 22 34 25
2 38.7 34 42 3
Transverse cirri, number 1 9.0 0.8 9.4 8 11 25
2 10.0 9 11 3
Distance 11 1 22.4 5.1 22.9 11 32 25
2 34.3 15 60 3
Resting cyst, diameter (in life) 49.2 4.5 9.1 43 60 17

I Distance from base of posterior-most transverse cirrus to posterior end of cell.

2 The anterior corona of the bicorona (see Fig. 197u).
3 All cirri excluding frontal (anterior corona) and transverse cirri.
 Ancystropodium 777

Taxa of Unknown Position within the Oxytrichidae

Apoamphisiella, Gastrostyla, Kerona, Paraurostyla, Parentocirrus, Pseudouroleptus,
and Territricha have at least one fragmenting dorsal kinety and are thus included in the
Oxytrichidae. In Gastrostyla, Paraurostyla, Parentocirrus, and Pseudouroleptus dorsal
morphogenesis is in the Oxytricha pattern which is very likely the plesiomorphic state
(Fig. 24a). Territricha (and Apoamphisiella?) shows not simple, but multiple fragmenta-
tion in kinety 3, and in the ectocommensal Kerona pronounced multiple fragmentation
occurs in kineties 1-3. However, not only dorsal, but also ventral morphogenesis of these
taxa provides some evidences that they belong to the Oxytrichidae, as already suggested
in previous classifications (see Table 13). The little-known Ancystropodium is also in-
cluded, because the ventral ciliature is reminiscent of Gastrostyla. EIGNER (1997) classi-
fied Gastrostyla, Kerona, Paraurostyla, and Parentocirrus in the Parakahliellidae.
Since I could not find out where these taxa branch off within the Oxytrichidae, I have
arranged them at the end of the book as taxa of unknown position within Oxytrichidae. I
prefer this arrangement to highly speculative ones, such as Paraurostyla is a ''primitive''
member of the Oxytrichidae (that is, branches off at the base of the cladogram), which is
of course not impossible. However, I could just as well assume that it branches of within
the Oxytrichinae having an increased number of frontal-ventral-transverse cirri as apo-
morphy. Very likely, as in Onychodromus, molecular biological data are needed to clar-
ify the situation.

Ancystropodium FAURE-FREMIET, 1907

1907 Ancystropodium FAURE-FREMIET, C. r. Seanc. Soc. BioI., 63: 377 - Type (by monotypy): Ancystropo-
dium maupasi FAURE-FREMIET, 1907.
1909 Ancystropodium FAURE-FREMIET, Arch. Protistenk., 13: 121.
1932 Ancystropodium FAURE-FR., 1909 - KARL, Tierwelt Dt\', 25: 598.
1972 Ancystropodium FAURE-FREMIET, 1907 - BORROR, 1. Protozoo\., 19: 14.
1974 Ancystropodium FAURE-FREMIET- STILLER, Fauna Hung., 115: 110.
1983 Ancystropodium FAURE-FREMIET, 1909 - CURDS, GATES & ROBERTS, British and other freshwater ciliated
protozoa, p 428.

Pro vis ion a I c h a r act e r is at ion: Adoral zone ofmembranelles formed like a
question mark. Body irregularly shaped, roughly like an upturned pyramid borne on a
contractile stalk-like process. Transverse cirri at posterior end of process, used to attach
to substrate. Most frontoventral and postoral ventral cirri form a continuous, slightly
oblique (frontoventral) row. One right and 1 left row of marginal cirri. Caudal cirri at
base of stalk-like process.

T a x 0 nom y: See single species, Ancystropodium maupasi.

778 SYSTEMATIC SECTION

Single species

Ancystropodium maupasi FAURE-FREMIET, 1907 (Fig. 198a-i)

1870 Stylonychiapustulata- TATEM, Mon. microsc. J., 4: 314 (misidentification; Fig. 198h, i).
1907 Ancyslropodium maupasi FAURE-FREMIET, C. r. Stanco Soc, BioI., 63: 377 (no illustration).
1909 Ancyslropodium maupasi FAURE-FREMIET, Arch. Protistenk., 13: 121 (Fig. 198a-g).
1910 Ancystropodium maupasi - MENzEL, K1einwelt, I: 34 (short German translation ofFAURE-FREMIET's pa-
per without own observations).
1932 Ancystropodium maupasi FAURE-FR., 1909 - KARL, Tierwelt Dtl., 25: 598 (incorrect date).
1974 Ancyslropodium maupasi FAURE-FREMIET- STILLER, Fauna Hung., 1I5: 1I2.
1983 Ancystropodium maupasi FAURE-FREMIET, 1909 - SHEN, Protozoa of the Tibetan Plateau, p 209, Plate
XLIV, Fig. 369 (see nomenclature and taxonomy; incorrect date).

Nom e n c I a t u rea n d . t a x 0 nom y: FAURE-FREMIET (1907) established this spe-

cies without illustration but with a sufficient description. 1907 is thus the correct year of
publication (BORROR 1972a, CORLISS 1979, JANKOWSKI 1979, TUFFRAU 1987, TUFFRAU &
FLEURY 1994).
Ancystropodium is one of the most curious species of hypotrichs, which has,
however, never been redescribed (except the misidentified Stylonychia pustulata sensu
TATEM 1870!). Thus, all available drawings, including SHEN'S (1983), are from FAURE-
FREMIET (1909). The ventral cirral pattern and its proposed morphogenetic origin indicate
a close relationship to Gastrostyla (Fig. 198e), whereas the prominent caudal cirri at the
base of the retractable tail-like process are reminiscent of Stylonychia (FAURE-FREMIET
1909, MENZEL 1910). Ancystropodium maupasi was thus always classified in the Oxytri-
chidae (BORROR 1972a, CORLISS 1979, CURDS et aL 1983, KAHL 1932, TUFFRAU 1987, TUF-
FRAU & FLEURY 1994). JANKOWSKI (1979, P 78) established the subfamily Ancystro-
podiinae.
The body shape, especially the tail-like process, is reminiscent of Psammomitra,
which has, however, a narrowed anterior end and very likely an Oxytricha-like cirral pat-
tern with only 18 frontal-ventral-transverse cirri (KAHL 1933, 1935). When feeding, this
marine psammobiont species is also attached to the substrate by a stalk-like process
(KAHL 1932, PARDucz 1938). KAHL (1932) and HEMBERGER (1982) supposed synonymy of
Oxytricha longicirrata (Fig. 46a, b) and A. maupasi. However, O. longicirrata has 18

Fig. 198a-g Ancystropodium maupasi (a-e, g, after FAURE-FREMIET 1909; f, from FAURE-FREMIET 1909. a-g, ~
from life). a) Ventral view of an attached specimen, size not indicated. b) Ventral view of a swimming speci-
men, 110 11m. c, d) Dorsal and ventro-iateral view of attached specimens, sizes not indicated. e) Arrangement
of cirri in a specimen with a retracted process. Cirri which originate during morphogenesis from the same
streak are connected by a broken line (exception: transverse cirri llIl-VlI). 1) Frontal view. g) Macronuclear
nodules and micronucleus, bar = 10 11m. CC = caudal cirri, TC = left transverse cirrus llIl, I-VI = frontal-
ventral-transverse cirri streaks I-VI.
Ancystropodium 779

" OJ
780 SYSTEMATIC SECTION

frontal-ventral-transverse cirri and, furthennore,

KAHL (1932) did not describe a retractable process
(the most important character of Ancystropodium) in
this species. All speculations about the validity of
Ancystropodium can only be resolved by a detailed
redescription.

M 0 r p hoI 0 g y and b i 0 log y: Size without

process in life 110 x 45 !lm. Body club-shaped. Flat-
tened about 1.5:1 dorso-ventrally (Fig. 198a, e, t: h).
Process about 90 !lm long, 2-3 !lm broad, posterior
end club-shaped, with distinct fibres at attachment
site (Fig. 198a, c, d). Invariably 2 spherical, about
15 !lm sized macronuclear nodules with large spheri-
cal nucleoli; invariably 1 spherical, about 6 J.1ID sized

~h
micronucleus between nodules, stains intensely with
methyl green-pyronin (Fig. 198g); CURDS et aI.
(1983), obviously erroneously, mentioned several
Fig. 198h, i Ancystropodium maupasi macronuclear nodules. Contractile vacuole at about
from life (after TA1EM 1870). Ventral level of cytostome. During feeding specimens attach
views (sizes not indicated) of a speci- to substrate by means of transverse cirri at end of
men with extended and retracted proc-
tail-like process; according to FAURE-FREMIET (1909),
ess; p 778.
A. maupasi can even make stepping movements in
this state. Swims rapidly with retracted process (Fig.
198c, d, i).
Adoral zone of membranelies about 40 % of body length (without process), proceeds
distally on prominent collar and makes a three-quarter turn (Fig. 198e, f). Membranelles
about 25 !lm long. Buccal area more or less funnel-shaped. Cirri on frontal area, and
transverse and caudal cirri distinctly larger than remaining cirri (Fig. 198e). Frontal cirri
Ill, II13, and nIl3 about 45 !lm, other cirri on frontal area about 25 !lm long. Invariably 5,
about 25 !lm long, very motile transverse cirri and only I pretransverse ventral cirrus at
posterior end of process. Ventral and marginal cirri about 20 !lm long. Cirri of streaks III,
IV, and V fonn a slightly obliquely arranged frontoventral row (Fig. 198e; more detailed
observations are needed to confinn these data). Only right marginal row extends to poste-
rior end of process; thus, in contracted specimens right marginal cirri more closely ar-
ranged posteriorly than anteriorly (Fig. 198a--e). Three conspicuous (like in Stylonychia
mytilus), about 35 !lm long caudal cirri at base of process (Fig. 198a, e). During morpho-
genesis 1,3,3,5,6,3 cirri originate from streaks I-VI (Fig. 198e; FAURE-FREMIET 1909,
1961b).

o c cur r e n c e and e colo g y: Extremely rare, probably confmed to freshwater.

Locus classicus is Rambouillet near Paris, France, where FAURE-FREMIET (1907, 1909)
 Apoamphisiella 781

discovered Ancystropodium maupasi in a peat-bog near a pond (etangs de Pontras aux

Essarts-Ie-Roi) in September. TATEM (1870) did not mention the collecting site of "Sty-
lonychia pustulata", presumably it is in the USA. GRABACKA (1973) recorded Ancystro-
podium maupasi from a fishpond in Poland, used for the treatment of sugar factory
wastes. WANG (1977) and SHEN (1983) found it in various freshwater habitats of the Ti-
betan Plateau. Feeds on bacteria and small flagellates (FAURE-FREMIET 1909).

Apoamphisiella FOISSNER, 1997

1997 Apoamphisiella FOISSNER, BioI. Fertil. Soils, 25: 335 - Type (original designation): Onychodromopsis
lihanyiensis GIiLLERT & TAMAs, 1958.

C h a r act e r i sat ion: Adoral zone of membranelles formed like a question mark.
Undulating membranes almost in Cyrtohymena pattern. Number of distinct frontoventral
cirri rather low (2 cirri), arranged in oblique row. One or more slightly enlarged postoral
ventral cirri. Five or more transverse cirri. Two frontoventral rows. One right and 1 left
row of marginal cirri. Caudal cirri present. Both frontoventral rows participate in anlagen
formation. Dorsal morphogenesis likely roughly in Oxytricha pattern, but probably not
with simple but with multiple fragmentation.

T a x 0 nom y: FOISSNER (1997b) suggested that Apoamphisiella tihanyiensis is an am-

phisiellid because of the postoral (= postperistomial) cirrus. He diagnosed Apoamphi-
siella as follows: "Amphisiellidae (?) with two long ventral cirral rows, a short, oblique
row of transverse cirri and at least one cirrus left of anterior end of ventral rows. Postperi-
stomial and caudal cirri present. With dorsomarginal kineties producing irregular field of
dorsal bristles near left anterior end. Both ventral rows participate in anlagen formation
during ontogenesis, and the postperistomial cirrus obtains its characteristic position by
distinct anlagen migration during cytokinesis." I classify Apoamphisiella in the Oxytri-
chidae as indicated by the fragmentation of dorsal kinety 3. Thus, I had to modify the
characterisation which is of course only provisional because detailed morphogenetic data
are lacking.
When I wrote the Apoamphisiella section I found that Holosticha hymenophora
STOKES, 1886, previously classified in Paraurostyla, is very similar to the type species,
Apoamphisiella tihanyiensis. I have great problems to find differences and thus separate
them by the cortical granules (present or absent). However, this character has to be
checked in detail in further populations from the type locations (for details, see A. hy-
menophora).
Onychodromopsis has a different pattern of undulating membranes, distinct pretrans-
verse ventral cirri, and 2 or more left marginal rows (Fig. 143d). In Parentocirrus hortu-
alis the rightmost frontoventral row is not involved in primordia formation (Fig. 218h).
Paraurostyla has more than 2 frontoventral rows (for example, Fig. 212r).
782 SYSTEMATIC SECTION

Key to species

1 Cortical granules present (Fig. 198.la-g, and Fig. 49 in FOISSNER 1997b) ........ .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . " A. tihanyiensis (p 782)
Cortical granules lacking (Fig. 215a~) ............... A. hymenophora (p 786)

Apoamphisiella fihanyiensis (GELLERT & TAMAs, 1958) ForSSNER, 1997

(Fig. 198.1a-g, Table 47.1)

1958 Orrychodromopsis tihanyiensis GELLERT & TAMAs, Annis Inst bioI. Tihany, 25: 230,240 (Fig. 198.le).
1997 Apoamphisiella tiharryiensis (GELLERT and TAMAS 1958) - FOISSNER, BioI. Fertil. Soils, 25: 335 (Fig.
198.la-d, f, g; two neotype slides ofprotargol-impregnated specimens are deposited in the Oben')ster-
reichische Landesmuseum in Linz, Upper Austria).

Nom e n c I a t u rea n d t a x on 0 my: "Pleurotricha tihanyensis" in FERNANDEZ-

LEBORANS & NovII.W (1993, P 216) is an incorrect subsequent spelling and a new combi-
nation, which is, however, not founded. This species is not mentioned by BORROR (1972a)
and STII.LER (1974b).
The long cirral rows of Apoamphisie/la tihanyiensis are reminiscent of Onycho-
dromopsis jlexi/is and Allotricha spp. However, Onychodromopsis jlexi/is (Fig. 143a~)
has short and straight undulating membranes and Allotricha (Fig. 91c, 92h) has the 18-
cirri pattern of the typical oxytrichids.HEMBERGER (1982, p 223) assumed that GELLERT &
TAMAs (1958) had overlooked some ventral cirri and thus classified it as senior synonym
of Pleurotricha variabilis REUTER (Fig. 91s), a supposed synonym of Allotricha mollis.
The FOISSNER (1997b) population from Peru (Fig. 198.1a~ f, g) is very similar to
the European type population, indicating a cosmopolitan distribution. There are,
however, also some differences discussed in detail by FOISSNER (1997b). The most impor-
tant one is the yellowish colour of the Peruvian population due to the cortical granules.
GELLERT & TAMAs (1958) did not mention such granules, very likely because they did not
make careful live observations, but studied mainly specimens prepared with opal blue.
Thus, they might simply have overlooked the cortical granules, which are sometimes less
distinctly yellowish (Brazilian populations mentioned by ForssNER 1997b). There are also
several differences in the cirral pattern of the European and Peruvian population, namely,
2 postoral ventral cirri against 1 cirrus and the length of the left frontoventral row (cp.
Fig. 198.1 e, t). Furthermore, the type population has distinct pretransverse ventral cirri
(arrow in Fig. 198.l.e) which is reminiscent of Onychodromopsis (Fig. 143d). As with
the cortical granules, FOISSNER (1997b) and I (preliminary) interpret these differences as
being a result of the different methodology used by the Hungarian workers. Detailed re-
description of further populations recommended. Synonymy with Apoamphisiella hy-
menophora cannot be excluded (see there).
 Apoamphisiella 783

b C

Fig. 198.1a-e Apoamphisiella tihanyiensis (a--d, from FOISSNER 1997b; e, from GELLERT & TAMAs 1958. a-d,
in life; e, opal blue preparation after BRESSLAU). a) Ventral view, 175 Ilm. b) Right lateral view. c) Cytoplasmic
crystals. d) Arrangement of cortical granules, which have a yellow colour. e) Ventral ciliature of type popula-
tion, 220 Ilm. Arrow marks some "pretransverse ventral cirri" (possibly a misobservation) not present in Fig.
198.1 f. DB = dorsal bristles, RM = right marginal row; p 782.

M 0 r p hoi 0 g y and b i 0 log y: Hungarian population: In life (?) about 220 Jl.m
long. Elliptical (Fig. 198.1e). Two macronuclear nodules, each with a micronucleus. No
cortical granules mentioned, very likely overlooked because probably not studied in life.
About 50 adoral membrane lIes. Undulating membranes strongly curved. Three distinctly
enlarged frontal cirri, 1 buccal cirrus, 2 frontoventral cirri, 2 postoral ventral cirri, 3 pre-
transverse ventral cirri. According to text 5 fringed transverse cirri which protrude
 784 SYSTEMATIC SECTION

j. . . ,

,
"
f'
r
, c:
#
, #
,
, ,
#
, I
, f'
,
r
#

,
,

r
.
, , ,
,
,
,
,
, f....,
, ,"
,
.•-
•
•..•.. 9

Fig. 198.lf, g Apoamphisiella lihanyiensis (from FOISSNER 1997b. f, g, protargol impregnation). C, g) Ventral
and dorsal infraciliature, 151 Jim. Arrow in (t) marks single postoral ventral cirrus (= postperistomial cirrus in
FOISSNER 1997b). Arrow in (g) marks caudal cirri at posterior-most fragment of dorsal kinety 3. Arrowhead de-
notes field of disorganised dorsal bristles due to mUltiple fragmentation in dorsal kinety 3 and/or parental bris-
tles; p 782.

beyond posterior end of cell, in the illustration, however, they do not!. Marginal rows
converge posteriorly, indicating that caudal cirri are present.
FOISSNER (1997b) population: In life 150-2lO x 50-80!-lm, dorsoventrally flattened
up to 2:1, very flexible but acontractile. Elliptical, both ends broadly rounded, right mar-
gin almost straight, left slightly convex (Fig. 198.la). Distinctly yellowish at low magnifi-
cation (:::;;IOOx) due to cortical granules. Invariably (n = II) 2 macronuclear nodules
 Apoamphisiella 785

rather close together in central third of body slightly left of mid-line, after protargol im-
pregnation 23-30 x 10-15 !lm (mean = 26.4 x 13.0 !lm, n = 11), with many 0.5-2.0 !lm
sized nucleoli. 4-8 (mean = 6, n = 11) spherical to slightly ellipsoidal (after protargol im-
pregnation 4.0 x 3.7 !lm, n = 11) micronuclei close to macronuclear nodules, compact
and thus easy to recognise in life (Fig. 198.1a, g). Contractile vacuole distinctly above
mid-body, with 2 lacunar collecting canals. Cortical granules 0.5-1;0 !lm across, con-
spicuous because brightly citrine, as, for example, in Holosticha multistilata, arranged in
rather widely spaced, longitudinal stripes, including cirral and bristle rows (Fig. 198.1d);
do not stain with protargol. Cytoplasm colourless, with many 2-4 !lm sized, yellowish
crystals (Fig. 198.lc) and food vacuoles. Glides quickly on slide and on soil particle.
Adoral zone of membranelles about 37 % of body length, proximal half slightly
spoon-like broadened with bases of largest membranelles about 12 J.lm wide and of con-
ventional fine structure; proximal end covered by buccal lip. Buccal cavity moderately
wide but deep and semicircularly curved anteriorly, posterior half covered by bay-like
projecting buccal lip (Fig. 198.la, f). Undulating membranes distinctly curved, especially
paroral, both very likely composed of narrowly spaced dikinetids; endoral optically inter-
sects paroral in mid-portion and traverses buccal cavity, forming bow-string to strongly
curved anterior portion of paroral (Fig. 198.1f); this arrangement strongly resembles the
Cyrtohymena pattern (Fig. 199). Pharyngeal fibres short, inconspicuous.
Invariably (n = 11) 3 frontal cirri, 1 buccal cirrus, 2 cirri left of anterior end of right
frontoventral row, and 1 postoral ventral cirrus (Fig. 198.lt: arrow); some specimens
from Brazil with 2 postoral ventral cirri (ForSSNER 1997b). Frontal cirri distinctly en-
larged, in life about 20 J.lm long, rightmost cirrus very close to distal end of adoral zone
of membranelies. Cirri left of anterior portion of right ventral row and isolated postoral
ventral cirrus slightly enlarged. Cirri offrontoventral rows in life about 15 J.lm long, right
row almost as long as body, begins near distal end of adoral zone of membranelies, termi-
nates in front of rightmost transverse cirrus; posterior-most cirrus slightly enlarged (pos-
sibly it is homologous with the pretransverse ventral cirrus VII2 of the typical
oxytrichids; Fig. 6a). Left frontoventral row in parallel with right but distinctly separate,
begins slightly in front of buccal vertex, extends to transverse cirri. Transverse cirri dis-
tinctlyenlarged, in life about 30 J.lm long and with frayed distal end, project above poste-
rior body margin, form slightly curved, oblique row slightly left of median. Marginal cirri
in life about 25 J.lm long, become gradually thinner posteriorly. Left marginal row J-
shaped, that is, terminates in median of cell, the posterior-most cirri thus easily confused
with caudal cirri. Right marginal row commences at level of buccal cirrus, terminates
about at level of posterior-most transverse cirrus, thus being separated from left marginal
row by small but distinct break.
Dorsal cilia in life about 3 J.lm long. Dorsal kineties 1-3 almost of body length; re-
maining kineties (posterior fragments of kinety 3 and dorsomarginal kineties) forming
conspicuous field of scattered bristles indicating that multiple fragmentation occurs
and/or parental kineties are retained (Fig. 198.1g). Ten caudal cirri on average on
786 SYSTEMATIC SECTION

Table 47.1 Morphometric data of Apoamphisiella tihanyiensis (from FOlssNER 1997b). All data are based on
protargol-impregnated specimens. All measurements in micrometres. CV = coefficient of variation (in %),
Max = maximum value, mean = arithmetic mean, Min = minimum value, n = sample size, SO = standard de-
viation

Character mean SO CV Min Max n

Body, length 163.1 13.8 8.5 147 190 II
Body, width 62.f 8.8 14.0 47 76 II
Adoral membranelles, number 56.5 3.3 5.8 52 62 II
Adoral zone of membranelles, length 60.8 4.1 6.7 55 65 II
Left marginal row, number of cirri 38.4 2.8 7.2 33 43 II
Right marginal row, number of cirri 35.4 4.2 11.9 28 43 II
Anterior end to left frontoventraI row, distance 41.6 1.9 4.5 40 45 11
Anterior end to right frontoventraI row, distance 20.2 2.0 10.1 15 22 II
Left frontoventraI row, number of cirri 24.2 1.9 8-0 21 27 II
Right frontoventraI row, number of cirri 33.4 2.9 8.6 27 37 II
Transverse cirri, number 6.6 0.8 12.5 5 8 11
Caudal cirri, number 10.0 3.3 32.9 3 14 II

posterior end of kineties 1 and 2, and probably on end of posterior-most fragment of ki-
nety 3; caudal cirri about of same length as marginal cirri.
FOISSNER (1997b) found in Brazilian freshwater populations that, as in Apoamphi-
siella hymenophora (GRIMES & L'HERNAULT 1978), both ventral rows develop cirral anla-
gen and the parental ventral infraciliature is completely resorbed.

o c cur r e n c e and e colo g y: Locus classicus is the eastern shore of Lake Bala-
ton, Hungary, where GELLERT & TAMAs (1958) discovered Apoamphisiel/a tihanyiensis in
drift material. GEllERT & TAMAs (1959b, 1960) and TAMAs & GELLERT (1959, 1960) ob-
served it in other habitats (for example, hydropsammon) of Lake Balaton. FOISSNER
(1997b) found A. tihanyiensis in a soil sample (light brown mixed with much leaf litter,
pH 6.2) from the Amazonian rain forest near the town of Iquitos, Peru (about lat 4°S,
long 74°W); he found it also in two freshwater sites in Rio de Janeiro and Praira do
Forte, Brazil. The marine record from the sub-littoral of the Bay of Biscay, Atlantic
Ocean, Spain, is questionable (FERNANDEZ-LEBORANS & NoVIILo 1993).
Feeds on diatoms and small ciliates (GELLERT & TAMAs 1958), according to FOISSNER
(1997b) on coccal cyanobacteria, fungal spores and hyphae, heterotrophic flagellates, na-
ked amoebae, testate amoebae (Trache/eug/ypha sp., Trinema lineare, Eug/ypha rotun-
da, E. compressa), ciliates (Blepharisma hyalinum, Vorticella swarmers), and even on
cysts or eggs of gastrotrichs.

Apoamphisiella hymenophora (STOKES, 1886) comb. nov. (Fig. 215a-d)

1886 Holosticha hymenophora STOKES, Proc. Am. phil. Soc., 23: 25 (Fig. 2ISa).
 Apoamphisiella 787

1888 Holosticha hymenophora, STOKES - STOKES, 1. Trenton nat. Hist. Soc., 1: 282 (Fig. 21Sa).
1932 Holosticha hymenophora STOKES, 1886 - KAlIL, Tierwelt Otl., 25: 586 (see nomenclature).
1963 Holosticha hymenophora STOKES - LUNDIN & WEST, Free-living protozoa, p 68 (Fig. 215b).
1972 Paraurostyla hymenophora (STOKES, 1886) - BORROR, 1. Protozool., 19: 10.
1978 Paraurostyla hymenophora (STOKES) - GRIMES & L'HERNAULT, 1. Protozool., 25: 65 (Fig. 21Se, d; mor-
phology and morphogenesis after electron microscopy).

Nom e n c I a t u rea nd ta x 0 nom y: Holosticha himenophora STOKES 1886 in

GRISPINI (1938, P 153) is an incorrect subsequent spelling. I<AHL (1932) divided Holosti-
cha into several subgenera; the correct name in his revision is thus Holosticha (Holosti-
cha) hymenophora.
BORROR (1972a) proposed synonymy with Uroleptus ellipticus WANG & NIE, 1933,
which, however, .lacks prominent transverse cirri. Onychodromopsis flexilis has more
scattered cirri right of the adoral zone of membranelles and 2 distinct pretransverse ven-
tral cirri, indicating that it is not synonymous with the species described by STOKES
(1886).
When I made the layout of the book, I found that "Paraurostyla hymenophord' be-
longs to Apoamphisiella (thus the illustrations are not arranged within the description!). It
is even rather difficult to find a difference between the type species, A. tihanyiensis, and
A. hymenophora, and synonymy thus cannot be excluded. STOKES (1886), who made de-
tailed live observations (in contrast to GELLERT & TAMAs 1958, the authors of the type
species, who used likely opal blue stained material only!) did not mention a colour or
granules, indicating that his species lacks cortical granUles. This is the only (uncertain!)
difference between A. tihanyiensis (with cortical granules according to the redescription
by FOISSNER 1997b) andA. hymenophora. GRIMES & L'HERNAULT (1978) also do not men-
tion cortical granules. However, some of their TEM-micrographs (their Figures 8-10)
show conspicuous structures close to the cirri, indicating that such granules are present.
The GRIMES & L'HERNAULT (1978) population differs in some further details from the
original description (left frontoventral row longer, undulating membranes curved, more
[6 against 5] transverse cirri). Thus, further redescriptions, including live observations
and detailed line drawings and biometry of protargol-impregnated specimens are needed
to ascertain if A. tihanyiensis is the junior synonym of A. hymenophora.

M 0 r p hoi 0 g y and b i 0 log y: In life 170-205 J.lm long, three to four times as
long as broad. Elliptical, slightly narrowed anteriorly and slightly curved towards left-
hand side. Soft and flexible and somewhat extensible. Two ellipsoid macronuclear nod-
ules. Two contractile vacuoles in mid-body (probably the main vacuole and the posterior
canal). Cytopyge subterminally, presumably dorsal.
Adoral zone of membrane lies about 35 % of body length. Undulating membranes ac-
cording to STOKES (1886; Fig. 215a) and LUNDIN & WEST (1963; Fig. 215b) almost
straight, according to GRIMES & L'HERNAULT (1978; Fig. 215c, d) distinctly curved (as in
A. tihanyiensis!). Three prominent frontal cirri, 1 buccal cirrus, and 1 prominent cirrus
behind right frontal cirrus (STOKES 1886), according to GRIMES & L'HERNAULT (1978)
788 SYSTEMATIC SECTION

anterior-most cirrus of left frontoventral row also distinctly enlarged. Two frontoventral
rows, according to original description and LUNDIN & WEST (1963) left one beginning at
buccal vertex, right one near distal end of adoral zone of membranelles. According to
GRIMES & L'lIERNAULT (1978; Fig. 215c) left frontoventral row begins about at same
level as right row (however, Fig. 1 in GRIMES & L'lIERNAULT 1978 gives the impression
that the left row is distinctly shortened anteriorly, identical as in Fig. 198.1t). Both rows
terminate immediately in front of right transverse cirri. Usually 1, sometimes up to 4
postoral ventral cirri (Fig. 215c). Five (STOKES 1886, LUNDIN & WEST 1963) to 6 (some-
times 7; GRIMES & L'HERNAULT 1978) moderately enlarged transverse cirri, only right one
slightly protruding beyond posterior end of cell. Marginal rows obviously not distinctly
separate posteriorly because they are described as confluent in all descriptions (very
likely, the inconspicuous caudal cirri are inserted in the gap between.the marginal rows).
Dorsal cilia short, immobile, arranged in 6 kineties; kinety 3 fragmented (GRIMES &
L'HERNAULT 1978) as in A. tihanyiensis. Ultrastructural details of ciliature, see GRIMES &
L'lIERNAULT (1978). Ultrastructure of cyst similar to that of 18-cirri oxytrichids, includ-
ing the lack of basal bodies (GRIMES & L'HERNAULT 1978).
Morphogenesis was studied by GRIMES & L'HERNAULT (1978) after protargol impreg-
nation and scanning electron microscopy. However, the line drawings are to inaccurate to
recognise important details. Thus, reinvestigation necessary. Frontal-ventral-transverse
primordia obviously develop independently in the proter and the opisthe. Both frontoven-
tral rows participate, as in Apoamphisiella tihanyiensis, in anlagen formation. The pres-
ence of 6 or 7 transverse cirri indicates that 7 or 8 anlagen are present (anlage I does not
make a transverse cirrus!).

o c cur r e n c e and e colo g y: Rare. Locus classicus not defined, but very likely
near Trenton, New Jersey, USA, where STOKES lived and worked; he found it in shallow
pools during early spring. Records documented by illustrations: freshwater from the Up-
per Peninsula of Michigan, USA (LUNDIN & WEST 1963, WEST 1953); Meadowbrook, a
small stream, where it crosses Hempstead Turnpike in Uniondale, New York, USA
(GRIMES & L'HERNAULT 1978; identified by A. C. BORROR). Records not documented by
illustrations: 16 indo cm-2 in running water in northern Italy (MAnoN! & GHETTI 1977); in a
well in Italy (GRISPINI 1938); Conestoga drainage basin, Pennsylvania, USA (CAIRNS
1965a). According to CURDS (1975) also in activated sludge.
GRIMES & L'HERNAULT (1978) maintained their population on phosphate-buffered
Cerophyl inoculated with Enterobacter aerogenes 2 d before use, and diluted 50 % (v/v)
with the Chapman-Andresen modification of Pringsheim's solution. Tetrahymena pyri-
formis was added to the above solution immediately before use.
 Gastrostyla 789

Gastrostyla ENGELMANN, 1862

1862 Gastrostyla ENGELMANN, Z. wiss. ZoO\., 11: 383 - Type (by monotypy): Gastrostyla steinii ENGELMANN,
1862.
1866 Nothopleurotricha DIESING, Sber. Akad. Wiss. Wien, 52: 521.
1866 Nothopleurotricha DIESING - DIESING, Sber. Akad. Wiss. Wien, 53: 117 - Type (by monotypy): Pleura-
tricha setifera ENGELMANN, 1862.
1882 Gastrostyla, ENGELMANN - KENr, Manual infusoria n, p 783.
1932 Gastrostyla ENGELMANN, 1861- KAm., Tierwelt Ot\., 25: 593.
1972 Gastrostyla ENGELMANN, 1862 - BORROR, J. Protozoo\., 19: 14.
1974 Gastrostyla ENGELMANN - STILLER, Fauna Hung., 115: 108.
1994 Gastrostyla ENGELMANN, 1862 - EIGNER & FOISSNER, J. Euk. Microbiol., 41: 260 (see taxonomy).

C h a r act e r i sat ion: Adoral zone of membranelles formed like a question mark.
Undulating membranes in Oxytricha pattern. Number of frontoventral and postoral ven-
tral cirri distinctly higher than 7. All frontoventral and postoral ventral cirri (usually ex-
cept of III12 and IVl2) form a more or less continuous, slightly oblique (frontoventral)
row. Two pretransverse ventral and 4 or 5 transverse cirri. One right and 1 left row of
marginal cirri. Five or 6 dorsal kineties. Caudal cirri present. Frontoventral cirral row
originates from primordia IV-VI. Dorsal morphogenesis in Oxytricha pattern.

Add i t ion a I c h a r act e r s: Both ends more or less broadly rounded, flattened
about 2:1 dorso-ventrally. Contractile vacuole in about mid-body or slightly in front ofit.
Cytoplasm colourless. Rapid movement. Adoral zone of membranelIes almost 40 % of
body length. except in Gastrostyla stenocephala, where it is only 25 %. Frontal cirri,
buccal cirrus, frontoventral cirrus behind right frontal cirrus (that is, cirrus 11112), and
transverse cirri usually distinctly enlarged. Frontoventral row at least half of body length,
composed of cirri of streaks VI, IV, and V (from anterior). Dorsal cilia short, that is,
3-5 /lm, except in G. stenocephala where they are 16/lm long.

Nom e n c I at u rea n d t a x 0 nom y: KAlIL (1932) and TUFFRAU & FLEURY (1994,
p 143) misdate Gastrostyla with 1861. Incorrect subsequent spellings: Gasterostyla
steinii (ENGLM.) (TCHANG 1931, p 71); Gasterostyla steini ENGELMANN (MAUCH et al.
1985, p 47); Gastostyla steini (HoRvATII 1950, p 153); Gastrostila (SERNOV 1929, P 557);
Gastrostylus (ALLISON 1924, p 345); Gastrosyla sp. ENGELMANN (SMITII 1978, P 27); Gas-
trosyla stenocephala (BORROR, 1963) (ALAnRO LUBEL et al. 1988, p 439); Gastrotyla ste-
nocephala (BORROR) (ALAnRO LUBEL 1985, P 28, 56).
For a detailed description of the patterns mentioned in the characterisation, see chap-
ter 2 in the general section. A relationship of Gastrostyla to the oxytrichids was already
supposed by several authors (for example, BORROR 1972a, CORLISS 1979, HEMBERGER
1982, STILLER 1974b, TUFFRAU & FLEURY 1994). In my opinion, Gastrostyla is unequivo-
cally an oxytrichid because of the fragmentation of dorsal kinety 3. Furthermore, the mor-
phogenetic pattern of the ventral infraciliature proceeds identically to in typical (I8-cirri)
oxytrichids, except that the 3 rightmost primordia IV-VI produce a higher number of
790 SYSTEMATIC SECTION

cirri to fonn a more or less continuous frontoventral row. However, it is difficult to estab-
lish the exact position of Gastrostyla within the oxytrichids. At least 2 possibilities exist:
(i) Gastrostyla is, simply, the sister-group of the typical (I8-cirri) oxytrichids, that is, the
18-cirri pattern evolved from a Gastrostyla-like pattern by a reduction of the number of
cirri originating from primordia IV-VI. This "primitive" position is generally supposed,
for example, by KARL (1932), MARTIN (1982), and WIRNSBERGER et al. (1986). (ii) Gas-
trostyla evolved from flexible 18-cirri oxytrichids (that is, Oxytrichinae, for example,
Oxytricha) by an increase of the number of cirri originating from primordia IV-VI. A re-
lationship to the Stylonychinae is unlikely because species with cortical granules exist
(for example, Gastrostyla pulchra). Furthennore, the body is usually not as rigid as in the
Stylonychinae because there is only one layer of (longitudinally oriented) subpellicular
microtubules (GRIM 1972). Because I can not decide for either of the two hypothesis, I
classify Gastrostyla as taxon of uncertain position within the oxytrichids.
Conversely, EIGNER & FOIssNER (1994) considered the possession of a frontoventral
row as sufficient to assign Gastrostyla to the amphisiellids. However, they failed to con-
struct a plausible cladogram indicating an uncertain classification. Furthennore, this hy-
pothesis assumes that fragmentation of dorsal kinety 3 evolved twice, which is very un-
likely. Moreover, Gastrostyla spp. can be separated from most amphisiellids (unfortu-
nately not from the type species, Amphisiella marioni; GOURRET & ROESER 1888, WICK-
LOW 1982a, b) not only by the dorsal pattern but also by the usually prominent, obliquely
arranged transverse cirri. Possibly, oxytrichids and amphisiellids are more closely related
than at present assumed.
The ventral infraciliature of Gastrostyla is reminiscent of Ancystropodium maupasi
(Fig. 198e). However, detailed redescription of this curious species is needed for further
speculations on phylogenetic relationship.
Synonymy of Gastrostyla and Nothopleurotricha was already proposed by KENT
(1882); according to CORLISS (1979, P 208) the junior synonym is a nomen oblitum.
STERKI (1878, p 57) proclaimed two new Gastrostyla species, which were, however,
never described.

Key to species

1 Marine ............................................................ 7
Limnetic or edaphic .................................................. 2
2 2 macronuclear nodules ............................................... 4
More than 2 macronuclear nodules ...................................... 3
3 4 macronuclear nodules (Fig. 199a, d) ...................... G. steinii (p 791)
8, rarely only 6 macronuclear nodules (Fig. 205a) .......... G. muscorum (p 808)
4 (2) Transverse cirri arranged in 2 distinctly separated groups with 3 and 2 cirri each
(Fig.204a-c) ......................................... G. setifera (p 816)
Transverse cirri arranged in a rather oblique, J-shaped pattern (e.g., Fig. 202b) .. 5
5 3 caudal cirri (Fig. 203a, b, e) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. 6
 Gastrostyla 791

7-12, usually 10 caudal cirri (Fig. 202a, g) .............. G. dorsicirrata (p 803)

6 Pyriform to ovoid, that is, anterior third of cell narrowed, posterior end broadly
rounded; frontoventral row terminates near transverse cirri (Fig. 203a, b) ........ .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. G. mystacea (p 808)
Prolate ellipsoidal, that is, widest in mid-body; frontoventral row terminates dis-
tinctly in front of transverse cirri (Fig. 203e, f, h) ............ G. minima (p 810)
7 (1) Anterior portion of cell cephalised, that is, distinctly narrowed; transverse cirri
terminally (Fig. 207a, f) ............................... G. stenocepha/a (p 822)
Anterior portion not cephalised; transverse cirri extremely displaced anteriad (Fig.
206a, b,e, h) .............................................. G. pu/chra (p 818)

Gastrostyla steinii ENGELMANN, 1862 (Fig. 2a, b, 7a, b, 199a-z, 200a-d,

201a-p, Tables 3, 48)

1862 Gastrostyla steinii ENGELMANN, Z.wiss. Zool., II: 383 (Fig. I 99a).
1882 Gastrostyla steinii, ENG. - KENr, Manual infusoria II, p 784.
1886 Gastrostyla vorax NUSSBAUM, Arch. mikr. Anat., 26: 494 (Fig. 1999; new synonym).
1899 Gastrostyla steinii ENGELMANN - ScHUSTER, Sci. Gossip, 5: 203 (Fig. 201m).
1901 Gastrostyla steinii ENGLM. - Roux, Mem.lnst. nato. genev., 19: 100 (Fig. 199h).
1906 Gastrostyla steinii ENG. - EDMONDSON, Proc. Davenport Acad. Sci., 11: 99 (Fig. 201k).
1914 Gastrostyla steinii ENGELMANN - SMI1ll, Kans. Univ. Sci. Bull., 9: 165 (Fig. 201p).
1924 Gastrostyla steinii ENGELMANN - BRAVO, Revta mex. BioI., 4: 165 (Fig. 201 n).
1926 Gastrostyla steinii, ENGELM. - KOFFMAN, Actazool., Stockh., 7: 312 (Fig. 2010).
1930 Gastrostyla steinii ENGELMANN - WEYER, Arch. Protistenk., 71: 139 (Fig. 199x).
1931 Gastrostyla philippinensis SHIBUYA, Proc. imp. Acad. Japan, 7: 126 (Fig. 1991, m; new synonym).
1932 Gastrostyla steini ENGELMANN, 1861 - KAIn.., Tierwelt Dtl., 25: 595 (Fig. 199k; incorrect date and in-
correct subsequent spelling).
1933 Gastrostyla steini ENGELM. - HORVArn, Arch. Protistenk., 80: 290 (Fig. 199i, j; incorrect subsequent
spelling).
1945 Gastrostyla steini ENG. - SOKOLOFF, An. Esc. naco Cienc. bioI., Mtx., 4: 41 (Fig. 199p; incorrect subse-
quent spelling).
1948 Gastrostyla steini ENGELM. - STELLA, Riv. BioI., 40: 144 (Fig. 2011; incorrect subsequent spelling).
1966 Histrio lemani DRAGESCO, Protistologica, 2: 89 (Fig. 199u; new synonym).
1970 Gastrostyla steinii - GRIM, Trans. Am. microsc. Soc., 89: 486 (Fig. 199r, s).
1972 Gastrostyla steini ENGELMANN - BICK, Ciliated protozoa, p 184 (Fig. 199q; incorrect subsequent spell-
ing).
1972 Gastrostyla steinii ENGELMANN, 1862 - BORROR, J. Protozool., 19: 14.
1972 Gastrostyla philippinensis SIllBUYA, 1931 - BORROR, J. Protozool., 19: 14.
1972 Histriculus lemani (DRAGESCO, 1966) - BORROR, J. Protozool., 19: 14.
1973 Gastrostyla steinii - WALKER & GRIM, J. Protozool., 20: 566 (Fig. 199t,200a-d).
1982 Gastrostyla steinii ENGELMANN, 1862 - FOISSNER, Arch. Protistenk., 126: 71 (Fig. 199b-f; detailed rede-
scription; slides are deposited in the Oberosterreichische Landesmuseum in Linz, Upper Austria).
1985 Gastrostyla parasteinii SMALL & LYNN, Ciliophora, p 455 (Fig. 199t; for Gastrostyla steinii of WALKER
& GRIM 1973; new synonym).
1991 Gastrostyla steinii ENGELMANN, 1862 - FOISSNER, BLATIERER, BERGER & KOHMANN, lnformationsberich-
te des Bayer. Landesamtes fur Wasserwirtschaft, l/91: 272.
792 SYSTEMATIC SECTION

t#
t#
t#
t#
t#
t#
t#
II
t#
t# II

•
t#
II
t#
t# II
t# ~ ~

t#
I "
,
I
I
I

•
#

Fig. 199a--c Gastrostyla stein;; (a, after ENGELMANN 1862; b, c, from FOISSNER 1982. a, b, from life; c, protar-
gol impregnation). a, b) Ventral views, a = 240 11m, b = 170 11m. Gastrostyla steinii usually has only 4 trans-
verse cirri and 4 macronuclear nodules. c) Ventral infraciliature, 90 11m. Arrow marks postoral ventral cirrus
which is homologous with cirrus IVI2 of the 18-cirri oxytrichids; p 791.

Nom e n cIa t u rea n d t a x 0 nom y: Incorrect subsequent spellings: the most

common is "GastrostyLa steint' (for example, ANDRE 1912, P 134, 1916; BURKOVSKY
1984, p 21; HORvArn 1933; KAHL 1932; PUYTORAC et al. 1994, p 3; SOKOLOFF 1930,
p 262; STILLER 1974b, p 108); Gastrostyla steinni (BORROR & WICKLOW 1982, P 285A).
Gastrostyla vorax, G. phi/ippinensis, and G. parasteinii were established mainly be-
cause of minor differences in the number and arrangement of the frontal, ventral, and
transverse cirri. However, the available data show that these characters are rather variable
in Gastrostyla steinii (Fig. 200a-d), indicating that all these species are synonyms.
DRAGESCO (1966b) compared Histrio lemani only with the marine "Histriculus simi/is"
(now Sterkiella quadrinucleatus, Fig. 186b). The number and the arrangement of the ven-
tral cirri, however, suggest that it is G. steinii. The only conspicuous difference is the
long adoral zone of membrane lIes, which is about 50 % of body length in Histrio Lemani.
 Gastrostyla 793

\
":";.i" ::;~~~~;
I'
r :. :"~":}~
....
I
'"

.... ;.:>

r.:L

e
Fig. 199<1-( Gastrostyla steinii (from FOISSNER 1982. d, protargol impregnation; e, t: from life). d) Dorsal in-
fraciliature, 90 11m. e, f) Dorsal (showing contractile vacuole) and left lateral view; p 791.

The populations by BALDENSPERGER (1927, Planche VI, 202) and SCHNEIDER (1930; Fig.
233h) are insufficiently redescribed.

M 0 r p hoi 0 g y and b i 0 log y: Size rather variable, that is, in life

140-320 x 45-100 /lm; 150-320/lm (ENGELMANN 1862), up to 300 x 130/lm (Nuss-
BAUM 1886), about 200 x 100 /lm (SHIBUYA 1931), 130-160 /lm long (DRAGESCO 1966b),
about 140-200 x 50-80 /lm (FOISSNER 1982), according to STELLA (1948; Fig. 2011) only
70 /lm long. Elliptical, often with parallel margins, sometimes posteriorly slightly broader
than anteriorly. After ENGELMANN (1862) and FOISSNER (1982) only slightly, according to
NUSSBAUM (1886) and HORvArn (1950) rather flexible. Usually 4, rarely 6, in life about
15 x 10 /lm sized macronuclear nodules, bounded by a common macronuclear envelope
with a narrow connecting isthmus between Gt interphase nuclei (WALKER & GOODE
1975). Usually 3-6, rarely up to 14 (WALKER & GOODE 1975) micronuclei; according to
WEYER (1930) the number of micronuclei varies between 1 and 7 within clones. Contrac-
tile vacuole with 2 long collecting canals; takes about 43 min at 23°C to excrete the
amount of water corresponding to cell volume (BRAND 1935). Cortical granules lacking.
Posterior half of cell often packed with reddish-yellow, about 2-5 11m long crystals and
794 SYSTEMATIC SECTION

Fig. 1999-k Gastrostyla steinii (g, after NUSSBAUM 1886; h, from Roux 1901; i, j, from HORVAlli 1933; Ie, af-
ter ENGELMANN 1862 from KAHL 1932. g, from life, arrangement of cirri after osmium fixation and pikrocar-
mine staining; h,j, Ie, from life; i, osmium toluidinblue stain). g-i, k) Ventral views, g, i = size not indicated, h
= 245 ~m, k = 240 ~m. j) Right lateral view; p 791.

with some 2-7 Jlm sized, shiny globules. Food vacuoles up to 35 Jlm in diameter. Defeca-
tion vacuoles with loose, granulated content, when ejected at right margin of posterior
third of cell (WEYER 1930). Usually glides, never stands still.
Bases of largest membranelles in life about 12 Jlm wide. Buccal area moderately
broad and rather deep. Undulating membranes consist of2 rows of basal bodies each, dis-
tinctly curved (KNAIPPE et al. 1989); intersect about halfway. All cirri rather strong. Fron-
tal and transverse cirri about 25 J.1m, marginal cirri about 20 J.1m long. Arrangement of
frontoventral cirri rather invariable, usually 5 or 6 frontoventral cirri and 9-11 postoral
and pretransverse ventral cirri. Usually 4, sometimes 5 (very rarely 6) transverse cirri;
however, it can not be excluded that pretransverse ventral cirri were counted as trans-
verse cirri. Dorsal kineties 1 and 3 distinctly, kineties 2 and 4 only slightly shortened an-
teriorly. Kineties 5 and 6 (= dorsomarginal rows) strongly shortened posteriorly (Fig.
199d). For details on fibrillar system and ultrastructure of cortex, see TUFFRAU (1965a)
and GRIM (1971, 1972).
Resting cysts spherical with strongly wrinkled surface, in life about 40-50 Jlm in dia-
meter (FOISSNER 1987a, HAUSMANN & FOISSNER 1986, NUSSBAUM 1886; Fig. 199v-z), ac-
 Gastrostyla 795

Fig. 1991--q Gastrostyla steinii (I, m, from SmsUYA 1931; n, 0, from HORvArn 1942; p, after SOKOLOFF 1945;
q, after BICK 1972a I, m, p, q, from life; n, 0, carmine acetic-acid staining). I, p, q) Ventral views, I, p = size
not indicated, q = 265 ~m. SOKOLOFF (1945, [p]) draws none and BICK (19723, [q]) 2 postoral ventral cirri. m)
Buccal area n, 0) Doubling (n) of the number of macronuclear nodules by means of micro-0peration; p 791.

cording to WALKER et al. (1980) only 27-291lm in diameter. Cyst wall 1.4-2.0 Ilm thick,
composed of 4 distinct layers (ROSATI et al. 1983). The cyst belongs to the kinetosome-
resorbing type (WALKER 1976a, WALKER & MAUGEL 1976, 1980, WALKER et al. 1980).
Volume of resting cyst about 13 % of volume ofan interphase specimen; volume offused
macronucleus about 20 % of interphase value (GUTIERREZ 1985, GUTIERREZ & PEREZ-
SILVA 1981). According to WENZEL & OMILIAN-Rosso (1982) the most effective means for
activation of cysts is to offer Tetrahymena pyriformis (reactivation rate about 78 % after
4 h). Cysts which were dry preserved for 12 years did not excyst (NUSSBAUM 1897). Mor-
phogenesis during encystment and excystment was analysed by KNAIPPE (1989) and
KNAIPPE et al. (1990).
M 0 r p hog e n e sis of cell division commences with the proliferation of basal
bodies near the left transverse cirrus (Fig. 201a, Table 3; HEMBERGER 1982, P 182; TUF-
FRAU 1969, 1970, WALKER & GRIM 1973). Frontal-ventral-transverse cirral primordia
probably originate independently in the proter and the opisthe (Fig. 201b-d). Altogether
about 25 frontal-ventral-transverse cirri are fonned from streaks I-VI (Fig. 201a, e-h).
The distinctly interrupted frontoventral row (Fig. 199c, 201a) consists of the migratory
cirri of streak VI, of the anterior-most cirrus of streak IV, and all cirri of streak V, except
796 SYSTEMATIC SECTIO
N
 Gastrostyla 797

y
Fig. 199v-z Gastrostyla steinii (v,after ENGELMANN 1862; w, after SOKOWFF 1945; x, after WEYER 1930; y,
after SoKOWV 1917; Z, from FOISSNER 1987a v-y, from life; Z, differential interference contrast). v-z) Resting
cysts, v, w, Y= size not indicated, x = 46).1m, z = 51 ).1m; p 791.

the pretransverse ventral and transverse cirrus (Fig. 20lf-h; BORROR & WICKLOW 1982).
Dorsal morphogenesis is in Oxytricha pattern, that is, simple fragmentation of kinety 3
and two dorsomarginal kineties occur (Fig. 201e, t: i, j; CALVO et al. 1981, HEMBERGER
1982). WALKER & GRIM (1973) probably overlooked the dorsomarginal rows (= kineties 5
and 6). Further details on Gastrostyla morphogenesis, see G. minima. The changes of the
nuclear apparatus during morphogenesis were described by NUSSBAUM (1886), TCHANG
(1931), WALKER (1976b), WALKER & GOODE (1973, 1976), WALKER et al. (1978), and
WEYER (1930; see also RAIKov 1982). The papers by MAUPAS (1888), NIETO et al. (1981),
and TORRES et al. (1980a) deal with regeneration and cell cycle.

o c cur r e n c e and e colo g y: Common, but usually with low abundance in detri-
tus and aufwuchs of stagnant and running waters and in terrestrial habitats. Cosmopolitan.
Locus classicus is Leipzig, Germany, where ENGELMANN (1862) discovered Gastrostyla
steinii in the so-called Diebesgraben together with Cyclidium glaucoma. Locus classicus
of the synonym Gastrostyla vorax is Bonn, Germany, where NUSSBAUM (1886) discov-
ered it in a hay infusion. Locus classicus of the synonym Gastrostyla philippinesis is an
agricultural soil of the Los Banos agricultural College in the Philippines (SHIBUYA 1931).
Locus classicus of the synonym Histrio lemani is a freshwater habitat near Thonon,
France (DRAGESCO 1966b). Locus classicus of the last synonym, Gastrostyla parasteinii,
is Sedona, Arizona, USA, where GRIM (1970) discovered it in a horse pond (see also
SMALL & LYNN 1985, WALKER & GRIM 1973).

~ Fig. 199r-u Gastrostyla steinii (r, s, from GRIM 1970; t, from WALKER & GRIM 1973; U, from DRAGESCO
1966b. r-u, protargol impregnation). r-u} Ventral infraciliature, H = size not indicated, u = 135 ).1m. Arrows
in (t) are for morphometric characterisation in Figure 200a--d; p791.
 798 SYSTEMATIC SECTION

40

30 .!:
30
~

.!:
~

c
.-
~

20 ';; 10
';;

E ~
c 10

20 21 22 23 24 25 26 21 28 29 30 31 10 11 12 13

cirral number a b
cirral number

50 50

40
40

!!!
30 30
.-
~

.- c
.-
-;;
';;
20 20
E
~

10 10

4 5 1 8 9 10 11
C 10 d
c irra I number cirral number

Fig. 200a-<l Morphometric characterisation of Gastrostyla steinii (from WALKER & GRIM 1973). a) Total
number offrontal-ventral-transverse cirri (that is, all cirri except marginals and caudals). b) Number of "frontal
cirri", that is, all frontoventral cirri which are, in Figure 199t, in front of the anterior arrow. c) Number of "me-
dial cirri", that is, all ventral cirri between the 2 arrows in Figure 199t. d) Number of "anal cirri", that is, all
ventral and transverse cirri behind the posterior arrow in Figure 199t; p 791.
 Gastrostyla 799

Further records substantiated by illustrations: various soils in Tullnerfeld and Styria,

Austria (FOISSNER 1982, FOISSNER et al. 1985, 1990); freshwater in England (ScHUSTER
1899); dunghill mixed with soil in the garden of the Kaiser Wilhelm Institute of Biology
in Berlin-Dahlem, Germany (WEYER 1930); various soils near Szeged, Hungary (HOR-
vATH 1933, 1950); Asinara River near Noto, Sicily, and in soil and hay infusions from It-
aly (ROSATI et al. 1983, STELLA 1948); freshwater from Puebla del Rio, Seville, Spain
(GUTIERREZ 1985); stagnant waters and hay infusions near Geneva and littoral of Lake
Geneva, Switzerland (ANDRE 1916, Roux 1900, 1901); Mexico (BRAVO 1924); freshwater
in Iowa and Kansas, USA (EDMONDSON 1906, SMITH 1914).
Records from freshwater habitats not substantiated by illustrations: Brennersee, a
small lake in Tyrol, Austria (DALLA TORRE 1891); moderately polluted river in Lower
Austria (HAEMPEL & WITIMANN 1910); Belgium (BERVOETS 1940); in the aufwuchs of the
Moldau River, Czechoslovakia, with highest abundance in January (KALMUS 1928); near
Perth, England (OwGIE 1921); mineral and thermo-mineral waters in France (CHAOUlTE
et al. 1990); peat-bog in Germany (SCHMIDT 1916); infrequent in a moderately polluted
brook near Bonn, Germany (JlITRCZENKI 1982); Elbe River in Hamburg, Germany
(BARTSCH & HARTWIG 1984, Roy 1938); among Cladophora glomerata in the Rhine
River in Coblenz, Germany (MARSSON 1910); Italy (LoNGHI 1892, 1894, 1895); water
tanks in the Netherlands (VERSCHAFFELT 1930); mesosaprobic river in Spain (FERNANDEZ-
LEBORANS & NOVII..LO-VILLAJOS 1993, FERNANDEZ-LEBORANS et al. 1990); up to 100 ind.l·'
in the botanical garden and in a gutter in Basle, Switzerland (RIGGENBACH 1922); with low
abundance in running waters and in lake plankton in Switzerland (STEINMANN & SURBECK
1918, 1934); polluted freshwater habitats in Japan (MURAMATSU 1957); Iowa, USA
(SHAWHAN et al. 1947); hay infusion in Massachusetts, USA (WOODRUFF 1905a); Mexico
(U>PEZ-OCHOTERENA & ROURE-CANE 1970); Brazil (CUNHA 1913); New Zealand (MAsKELL
1886).
Absent in marine brackish water (BICK 1967a); however, according to HERFS (1922)
Gastrostyla steinii tolerates 2 % NaCl- (see also WEATHERBY 1964). Not recorded from
sewage treatment plants (CURDS 1975).
Records from terrestrial habitats not substantiated by illustrations: garden soil in Up-
per Austria (PETZ et al. 1988); various agricultural soils in Czechoslovakia (TIRJAKovA
1988); England (DIXON 1937; GooDEY 1911); Italy (COPPA 1921); Sweden (KOFFMAN
1934); USSR (DIXON 1937); Japan (TAKAHASHI & SUHAMA 1991).
Feeds on bacteria, algae (diatoms, Chlorogonium elongatum, Gonium pectorale),
fungi, flagellates (Chilomonas paramaecium), and ciliates, like Colpidium, Glaucoma,
Vorticella, Colpoda cucullus (BICK 1968a, DRAGESCO & DRAGESCo-KERNEIS 1986, FOISS-
NER 1982, GooDEY 1911, HORVATH 1933, 1950, NUSSBAUM 1886, SCHMERENBECK 1975,
WEYER 1930)_ In cultures Gastrostyla steinii also attacks active and encysted specimens
of its own species (NUSSBAUM 1886, WENZEL 1974, WEYER 1930). WENZEL & KORFF
(1981) obtained thriving cultures with species of the Tetrahymena pyriformis complex as
food and a suspension of yeast cells or hay-extract. Good cultures were also obtained
with Chlorogonium elongatum and Escherichia coli, Enterobacter aerogenes, or Acine-
800 SYSTEMATIC SECTION

........--- ~.""'.'.,. ......

11.'" .~
.... .....
.:
'J

"~~ ':~~"
~ "

...........-
~ .......... ~
.................
~ •••• ~'\l, .-. t)~~ ~
~.:
' ; " "• • : • • • : -
",.....
•• :....
• ••••• -
()
"~,I"~
, 0

.." .... " .... "" ...

""" .....----.ll •• I.,
...!' •••• ,
..... , _____ ........ ....
....... .... ,.... ...•

" .. " ......

.......... ....
• II • • I. ()

OJ

o 0 Q 0 11

.' ,•
 Gastrostyla 801

Fig. 201k-p Gastrostyla steinii (k, from EDMONDSON

1906; 1, from STELLA 1948; m, from SCHUSTER 1899; n,
from BRAVO 1924; 0, after KOFFMAN 1926; p, after SMCIH
1914. k-p, from life). k-p) Ventral views, k = 250 11m, I
= 70 JIm, m = 150-300 11m, n = size not indicated, 0 =
230 11m, p = 150-330 11m; p 791.

tobacter calcoaceticus as food. 0.01 % Knop's-solution (HoRvATH 1939, WEYER 1930),

Pringsheim-medium (for example, NIETO et al. 1981), or cerophyl and wheat grain extract
(DRAGESCO & DRAGESCo-KERNEIS 1986, GRIM 1970) were successfully used as culture me-
dia. Polysaccharides were stored as paraglycogen granules with 0.5 ~m diameter (VERNI
& ROSATI 1980). Generation time about 19 h at 16°C and with Chlorogonium elongatum

~ Fig.201a-j Gastrostyla steinii (from HEMBERGER 1982. a-j, protargol impregnation). a-h) Ventral infracilia-
ture of morphogenetic stages, bar = 20 11m; details, see text. (f-h) and (a) show migration of some cirri. Small
arrows in (a, h) mark cirrus IIII2 and lV/3 respectively. Large arrow denotes anterior-most cirrus of streak V
(probably homologous to cirrus V14 of the 18-cirri oxytrichids) and arrowhead marks postoral ventral cirrus
IV12. New structures black, parental white. i, j) Dorsal morphogenesis showing simple fragmentation ofkinety
3 (arrowheads) and 2 dorsomarginal kineties (arrows); parental kineties omitted. Fragmentation strongly indi-
cates that Gastrostyla is an oxytrichid. OP = oral primordium; p 791.
802 SYSTEMATIC SECTION

Table 48 Morphometric data of Gastrostyla dorsicirrata (dol, from FOISSNER 1982; do2, original data), G.
minima (mil, from HEMBERGER 1985; mi2, from FOISSNER 1997a), and G. steinii (stl, from ORAGESCO 1966b;
st2, from FOISSNER 1982). All data are based on protargol-impregnated material. All measurements in microme-
tres. ? = sample size not indicated; if only 1 value is known it is listed in the mean column, if 2 values are
available they are listed as Min and Max. CV = coefficient of variation (in %), Max = maximum value, mean
= arithmetic mean, Min = minimum value, n = sample size, SO = standard deviation

Character Species mean SO CV Min Max n

Body, length dol 106.7 9.8 9.1 90 122 8
do2 110.9 10.8 6.7 93 134 30
mil 120 145 ?
mi2 75.5 9.3 12.4 62 97 15
stl 130 160 ?
st2 78.3 10.3 13.1 67 95 10
Body, width dol 37.5 3.8 10.1 29 42 8
do2 37.6 4.0 10.5 31 45 30
mil 40 70 ?
mi2 29.4 4.1 13.8 22 37 15
st2 43.0 5.6 13.1 33 51 10
Macronuclear nodule, length dol 18.9 2.1 ILl 16 22 8
do2 2 26.1 2.7 10.3 22 31 27
mi2 13.4 2.6 19.3 9 19 15
st2 11.5 1.9 16.3 8 15 10
Macronuclear nodule, width dol 9.4 1.3 14.3 8 12 8
do2 2 10.7 Ll 10.7 9 13 27
mi2 3.2 0.2 7.3 3 4 15
stl 7.4 0.9 12.1 7 9 10
Adoral zone of membranelles, dol 39.5 3.1 7.8 37 47 8
length do2 42.6 3.2 7.6 38 52 30
mi2 26.5 2.2 8.2 23 31 15
st2 33.2 5.3 15.9 27 42 10
Adoral membranelles, number dol 35.1 1.5 4.4 33 37 8
do2 36.0 2.2 6.1 30 40 30
mil 32 35 ?
mi2 32.2 2.0 6.6 29 37 15
stl 34 38 ?
st2 34.4 3.3 9.5 31 40 10
Right marginal row, number dol 28.5 1.6 5.5 26 31 8
of cirri do2 27.5 2.8 10.3 21 32 30
mil 28 33 ?
mi2 32.2 3.9 12.0 25 39 15
stl about 30 ?
st2 27.7 2.3 8.4 23 31 10
Left marginal row, number dol 28.9 1.9 6.6 26 32 8
of cirri do2 29.1 1.8 6.1 25 32 30
mil 28 31 ?
mi2 31.7 2.1 6.6 28 34 15
st2 27.7 1.5 5.4 25 30 10
Transverse cirri, number dol 5.0 0 0 5 5 8
do2 5.1 0.4 7.9 5 7 30
 Gastrostyla 803

Table 48 Continued

Character Species mean SD CV Min Max n

Transverse cirri, number mil 5.0 ?
mi2 3 5.1 5 6 15
stl 5.0 ?
st2 4.0 0 0 4 4 10
Dorsal kineties, number doll 4.0 0 0 4 4 8
do21 4.0 0 0 4 4 22
mil 5.0 ?
mi2 6.0 6 6 15
stl 6.0 ?
st2 6.0 0 0 6 6 10

I Kinety 3 shows incomplete simple fragmentation; new kinety 4 is thus not counted separately; see Figures
202c, i and description.
2 Posterior nodule.

3 Higher value found in lout of 15 specimens investigated.

as food, 14 h at 21 DC, 11 hat 26 DC (WEYER 1930); according to TORRES et al. (1980a)

about 10 h at 20 DC and with Chlorogonium sp. as food. Cultivation failed at 6-8 DC and
at 32 DC (WEYER 1930). KJIP04 accelerated the division-rate during the early part and
retarded it toward the end of the life-cycle (WOODRUFF 1907). Biomass of 106 individuals
about 120 mg (FOISSNER 1987a).
BICK (1967a, b, 1968a, b, 1972a, 1973), BICK & DREWS (1973), BICK & KUNZE (1971),
BICK & SCHMERENBECK (1971), BICK & BERTRAM (1973), and WILBERT (1977) provided fol-
lowing autecoiogical data: 5-35 DC, pH 5.0-9.5, 0-9 mg 1.1 O2, 0-72 mg 1.1 C02fr..,
0.1-43 mg 1.1 NH/-N, 60000 to 8 X 106 bacteria mI'l (plate method). MONcH (1970)
counted about 1400 indo mI· l in experiments on the decomposition of 0.25 g mI· l peptone
at vel)' low oxygen content and at 20 mg 1.1 NH/-N. Autecoiogical date for edaphic
populations, see FOISSNER et al. (1985).
In agreement with SLADECEK et al. (1981) and WEGL (1983), ForSSNER (1988a) and
ForSSNER et al. (1991) classified Gastrostyla steinii as alphamesosaprobic indicator ofwa-
ter quality (b = 2, a = 7, P = 1, 1= 3, SI = 2.9).

Gastrostyla dorsicirrata FOissNER, 1982 (Fig. 202a-p, Tables 3, 48)

1982 Gastrostyla dorsici"ata FOISSNER, Arch. Protistenk., 126: 69 (l slide ofholotype and I slide of para-
type specimens are deposited in the Oberosterreichische Landesmuseum in Linz, Upper Austria).

Nom e n c I a t u r e: ForSSNER (1981, p 18) mentioned this taxon in a faunal list as "Am-
phisiella dorsicirrata FOISSNER"; this is a nomen nudum because no description was pro-
vided (FOISSNER & ForSSNER 1988). The following chapter includes data from the original
804 SYSTEMATIC SECTION

...':
,
,
, •
#
#
•

Fig.202a--c Gastrostyla dorsici"ata (from FOISSNER 1982. a, from life; b, c, protargol impregnation). a) Ven-
tral view, 150 ~m. b, c) Ventral and dorsal infraciliature, b = 110 ~m, c = 120 ~m. Note the postoral ventral
cirrus (arrow). For correct arrangement of dorsal kineties 3 and 4, see Figure 202i (see text for explanation);
p803.

description, unpublished data from the FOISSNER (1986a) population, and own observa-
tions (Fig. 202h-p).

M 0 r p hoi 0 g y and b i 0 log y: In life about 130-180 x 45-80 ~m (FOISSNER

1982). Population from Himalayan Mountains 110-140 x 45-70 ~m (FOISSNER 1986a,
W. FOISSNER, pers. comm.), according to own observations 150-180 x 70-80 ~m. Long
elliptical, sometimes with ± parallel margins. Not contractile, but rather flexible. Macro-
nuclear nodules in life 19-30 x 9-13 ~m, finely granulated or with large, clod-like nucle-
oli. 2-4, usually 3, in life about 3 ~m sized spherical, strongly refractile micronuclei.
Contractile vacuole with collecting canals. Cytopyge right laterally near posterior end of
cell. Cortical granules lacking. Cytoplasm often packed with yellowish, cylindrical crys-
tals, especially in posterior portion of cell so that specimens appear dark at low magnifi-
cation.
 Gastrostyla 805

Fig. 202d--g Gastrosty/a dorsi-

cirrata (from FOISSNER 1982. d, e,
from life; f, g, protargol impregna-
tion). d, e) Dorsal and lateral
view, d, e = 160 Jim. t) Ventral in-
fraciliature of posterior cell por-
tion, bar = 20 Jim. g) Dorsl!l infra-
ciliature of posterior cell portion
showing increased number of cau-
dal cirri. FB = faecal ball; p 803.

Bases of largest adoral

membranelles in life about
11-14 ~m wide. Buccal
area deep. Buccal cirrus
near anterior end of dis-
tinctly curved and intersect-
FB ing undulating membranes.
Frontal cirri 25-30 ~m,
frontoventral, postoral ven-
d e tral, and marginal cirri
15-20 ~m long in life. Left
marginal row J-shaped,
right one tenninates at level of posterior-most transverse cirrus. Transverse cirri in life
about 20-25 ~m long, protrude distinctly beyond posterior end of cell. Frontoventral row
with 2 ± distinct interruptions, tenninates slightly behind mid-body. Behind cytostome a
single cirrus left of frontoventral row (Fig. 202b, arrow), which is certainly homologous
to cirrus IV/2 of the 18-cirri oxytrichids having only 3 postoral ventral cirri. Invariably 2
pretransverse ventral cirri (n = 30).
The arrangement of the dorsal kineties was not completely clear in the original de-
scription (Fig. 202c). Slides from another population and the reinvestigation of the type-
material showed that kinety 3 is distinctly curved in the posterior third (Fig. 202i; arrow-
head). Kinety 1 distinctly shortened anteriorly, kinety 4 (= dorsomarginal row) tenninates
in about mid-body. Dorsal cilia in life about 3 ~m long. 3-6 (mean = 3.7; n = 23) caudal
cirri on kinety 1,2-4 (mean = 2.8; n = 21) on kinety 2, and 1-3 (mean = 2.1; n = 20) on
kinety 3. Caudal cirri in life about 30 ~m long, energetically motile.
Cysts (culture material) spherical, about 45 ~m in diameter. Wall 1 ~m thick, rather
smooth, surrounded by an about 10 ~m thick, inconspicuous mucous layer.

M 0 r ph 0 g e n e sis of cell division commences with the proliferation of basal bodies

in the posterior third of the cell, probably near the left transverse cirrus (Fig. 202j, Table
3). The streaks I-VI produce the following number of cirri: 1, 3, 3, 4-7 (mean = 5.6),
6-10 (mean = 7.4), and 6-9 (mean = 6.7; Fig. 202k). The frontoventral row is fonned by
806 SYSTEMATIC SECTION

Fig.202h-k Gastrostyla dorsicirrata (originals. h-k, protargol impregnation). h, i) Ventral and dorsal infra-
ciliature of an non-dividing specimen, 115 11m. The arrowhead in (i) marks the incomplete simple fragmenta-
tion in dorsal kinety 3 (see text for further explanation). j) Early morphogenetic stage in ventral view, 130 11m.
k) Late morphogenetic stage in ventral view, 135 11m. The arrowhead marks the primordiwn of the dorsomar-
ginal row (= dorsal kinety 4) of the opisthe. New structures black, parental white. 1-4 = dorsal kineties 1-4 (i;
because of incomplete fragmentation [arrowhead] of kinety 3 the posterior part [usually "kinety 4"] was not
counted separately. Thus, altogether only 4 dorsal kineties present), 1-6 = frontal-ventral-transverse cirri
streaks I-VI of the opisthe (k); p 803.

the cirral rows of the streaks VI, IV, and V (from anterior to posterior) only in the postdi-
viders (Fig. 202m, 0). Dorsal morphogenesis is practically in Oxytricha pattern, that is,
new kineties 1-3 originate within the parental kineties 1-3, kinety 3 shows simple (in-
complete; see below) fragmentation, and 1 dorsomarginal kinety occurs (Fig. 2021, k). In-
complete, simple fragmentation means that the posterior portion with the caudal cirrus
does not distinctly separate from the anterior portion (Fig. 202i, n, arrowheads) as is the
case in the typical Oxytricha pattern.

o c cur r e n c e and e colo g y: Probably confmed to terrestrial habitats (soil,

mosses). Europe and Asia. Locus classicus is an alpine, grazed and mowed meadow on
alluvial soil near the Glockner Road, Austria (site SO 1 in FOISSNER 1981). FOISSNER &
PEER (1985) provided some autecological data of an alpine population from the Gastein
 Gastrostyla 807

Fig.2021-p Gastrostyla dorsicirrata (originals. I-p, protargol impregnation). I) Late morphogenetic stage in
dorsal view, 130 J1Il1. m) Very late stage in ventral view, 135 11m. 0) Very late stage in dorsal view, 150 11m.
Arrowheads mark bend in posterior portion of new kineties 3 (= incomplete simple fragmentation). Some frag-
ments of parental kineties are still preserved. 0, p) Proter in ventral and dorsal view, 85 11m. The arrowhead
marks the posterior part of dorsal kinety 3 (in normal Oxytricha pattern this is kinety 4). New structures black,
parental white; p 803.

area, Austria. I found Gastrostyla dorsici"ata in mosses from rocks in a wood near the
village of Garsten, Upper Austria (Fig. 202h-p) and in a sub-alpine grassland near Aigen,
Styria, Austria (FOISSNER et al. 1990). FOISSNER (1986a) found it in soil from the Pisang
Peak (2200 m and 4500 m above sea level) in the Himalayan Mountains.
Feeds on cyanobacteria, fungal spores, and ciliates (FOISSNER 1982, own obser-
vations). According to FERNANDEZ-LEBORANS et al. (1987) rather insensitive to increased
concentrations of mercury. Biomass of 106 medium-sized specimens about 96 mg (FOISS-
NER 1987a).
808 SYSTEMATIC SECTION

Gastrostyla muscorum KARL, 1932 (Fig. 205a, Table 17)

1932 Gastrostyla muscorum KAHL, Tierwe1t Dtl., 25: 595 (Fig. 205a).
1972 Gastrostyla muscorum KAHL, 1932 - BORROR, J. Protozool., 19: 14.

Nom e n c I a t u rea n d t a x 0 nom y: Incorrect subsequent spelling: Gastrostyla

muscororum (BOVEE 1979, p 621). Detailed redescription necessary. It cannot be ex-
cluded that Gastrostyla muscorum is a G. steinii with an increased number of macronu-
clear nodules resulting from caryokinesis without corresponding cortical development
(WALKER & GOODE 1976).

M 0 r p hoi 0 g y: In life 130-200 ~m long. Ovoid, left margin straight, right convex.
Three anterior-most cirri of frontoventral row separated by a gap from posterior portion
of row. Five transverse cirri, protrude distinctly beyond posterior end of cell.

o c cur r e n c e and e colo g y: Rare species, in soil and freshwater. Europe and
North America. The locus classicus is near Hamburg, Germany, where KAHL (1932)
found Gastrostyla muscorum only once and with high abundance in terrestrial (?) mosses.
Records not substantiated by illustrations: Roussenski Lorn River and other rivers in
Bulgaria (DErCHEVA 1972a, 1975a, 1979b, RUSSEV et al. 1976; autecological data, see Ta-
ble 17); soil of a beech wood in Denmark (STOUT 1968); limnetic habitat in Italy (GRAN-
DORI 1934); litter layer of a pond in Japan (KUSANO 1985); soil of Great Abaco Island,
Bahamas, and soil ofa cedar glade in Tennessee, USA (CAIRNS & RUTHVEN 1972, MARTIN
& SHARP 1983); forest mosses from Lake Itasca Area, Minnesota, USA (BOVEE 1979).
Feeds on algae (Melosira sp., diatoms <65 ~m), flagellates, and ciliates (KAHL 1932, Ku-
SANO 1985). Polysaprobic (RUSSEV et ai. 1976).

Gastrostyla mystacea (STEIN, 1859) STERK!, 1878 (Fig. 203a-d)

1859 Oxytricha nrystacea STEIN, Organismus der Infusionsthiere I, p 188 (Fig. 203a-c).
1877 Oxytricha nrystacina, STEIN - WRZESNIOWSKl, Z. wiss. Zool., 29: 278 (mentions relationship to Holosti-
cha; incorrect subsequent spelling).
1878 Oxytricha nrystacea STEIN - STERK!, Z. wiss. Zoo\., 31: 57 (mentions relationship to Gastrostyla).
1882 Holosticha nrystacea STEIN sp. - KENT, Manual infusoria II, p 769.
1882 Oxytricha nrystacina (STEIN) - KOWALEWSKlEGO, Pam. ftzyogr., 2: 411 (mentions relationship to Am-
phisia; incorrect subsequent spelling).
1889 Gastrotricha nrystacea STEIN - BOTsCHLI, Protozoa, legend to Tafel LXXI, Fig. 7b (misspelling).
1932 Holosticha nrystacea STEIN, 1859 - KAHL, Tierwelt Otl., 25: 585 (Fig. 203d; redrawn after STEIN
1859b).
1972 Holosticha mystacea (STEIN, 1858) KAHL, 1932 - BORROR, 1. Protozoo\., 19: 14.
1983 Holosticha mystacea (STEIN, 1859) - BORROR & WICKLOW, Acta Protozool., 22: 115.
1991 Gastrostyla mystacea (STEIN, 1859) STERK!, 1878 - FOISSNER, BLATIERER, BERGER & KOHMANN, Infor-
mationsberichte Bayer. Landesamtes fUr Wasserwirtschaft, 1191: 270 (in part, see nomenclature and
taxonomy).
 Gastrostyla 809

Fig.203a-d Gastrostyla mystacea (a, b, from STEIN 1859b; c, after STEIN 1859b; d, after STEIN 1859b from
KAHL 1932. a-d, from life). a, b) Ventral views, sizes not indicated. Arrow in (b) marks the conspicuous pos-
toral ventral cirrus. c) Cyst. d) Ventral view, 150 ~m. The redrawing by KAHL (1932) is rather superficially
feigning a midventral row; p 808.

Nom e n c I a t u rea n d t a x 0 nom y: Incorrect subsequent spellings: Holosticha

mystacina (JiROVEC et al. 1953, p 512); Oxytricha mistacea ST. (GAYEWSKAYA 1924,
P 243); Oxytricha nuptacina STEIN (KOWALEWSKI 1883, P 243). STERKI (1878) first recog-
nised that this species could belong to Gastrostyla. The authorship of the combination
should thus be credited to him although he did not use the binomen Gastrostyla
mystacea.
The frontoventral row and the single, slightly enlarged cirrus just behind the cy-
tostome (Fig. 203b, arrow), which is very likely homologous to cirrus N12 of the 18-cirri
oxytrichids, are the most important characters indicating a classification in Gastrostyla.
This conspicuous cirrus left of the frontoventral row is lacking in Holosticha, where Gas-
trostyla mystacea was classified by KAHL (1932), BORROR (1972a), and HEMBERGER
(1982, P 91). This misclassification was supported especially by KAHL (1932) in that he
interpreted the frontoventral row as a holostichid midventral row (Fig. 203d).
Gastrostyla minima agrees rather well with the original description by G. mystacea,
especially in that the transverse cirri are displaced anteriad (cp. Fig. 203a, t). I thus syn-
onymised them in FOISSNER et al. (1991). However, FOISSNER (1997a) found a population
which matches G. minima better than G. mystacea, indicating that the synonymisation
was premature. Detailed redescription of G. mystacea necessary. The CHORIK (1968; Fig.
233e) and VUXANOVICI (1963; Fig. 205d) populations are insufficiently redescribed.

M 0 r ph 0 log y and b i 0 log y: In life about 120-170 x 40-70 J-lm. Ovoid, right
margin convex, left slightly concave, sometimes straight. Anterior third of cell often nar-
rowed (Fig. 203a). No cortical granules mentioned. Cytoplasm according to STEIN
(1859b) with a bluish shimmer. Frontoventral row according to Figures 203a, b with
810 SYSTEMATIC SECTION

about 20 cirri. Five transverse cirri, slightly displaced anteriad so that they tenninate at
about posterior cell margin. Caudal cirri inconspicuous. STEIN (1859b) described cysts,
sometimes infected by Chytridiomycetes, probably belonging to this species (Fig. 203c).

o c cur r e n c e and e colo g y: Rare, in freshwater. Europe, Asia, Australia. Locus

classicus is Tharandt near Dresden, Gennany, where STEIN (1859b) discovered Gastro-
styla mystacea with high abundance in a pool with liquid manure in community with a
mass occurrence of Euglena viridis and Chlamydomonas pulvisculus. STEIN (1859b) also
found it abundantly in similar habitats in Niemegk, a village near Berlin, Gennany, and in
a pond (also polluted with liquid manure) in Prague, Czechoslovakia; Didinium nasutum
occurred at both sites.
Records not substantiated by illustrations: Latvian rivers (LIEPA 1978); freshwater
and marine (?) habitats in the Netherlands (VERSCHAFFELT 1930); more than 100 indo 1.1at
0.8-17.8 mg 1-1 O2 in the surface layer of a basin and up to 10000 ind I-I at 2°C,
3.2 mg 1-10 2, 16.2 mg I-I COUI..., and 14 °dR carbon-hardness in the deep area of this ba-
sin in Basle, Switzerland (RIGGENBACH J 922); polluted reservoirs in Azerbaijan (ALEK-
PEROV 1981, 1982a, 1989); pond water and infusions of dried (submerged or edaphic?)
moss in Sri Lanka (BHATIA 1936; DADAY 1898, determined by G. ENTz); Lake Erie, USA
(JENNINGS 1899a); according to a table in DADAY (1898, P 9) also recorded from Aus-
tralia. GREEFF (1889) found Gastrostyla mystacea in an edaphic habitat - but where? The
marine record from the harbour of Genoa, Italy (GRUBER 1884), is uncertain.
Feeds on euglenoid flagellates and peritrichous ciliates (STEIN 1859b). Biomass of
10 specimens about 120 mg (FOISSNER et aI. 1991). Gastrostyla mystacea is a polysapro-
6

bic indicator of water quality (p-a; a = 3, P = 7, 1=4, SI = 3.7; FOIsSNER 1988a, FOIsSNER
et al. 1991, SLADECEK et al. 1981). KOLKWITZ & MARSSON (1909) and KOLKWITZ (1950)
classified it as alphamesosaprobic. Probably all classifications are based mainly on the
data by STEIN (1859b) and RIGGENBACH (1922), who found it in heavily polluted habitats
(for example, dung pools). However, these sparse data must be checked on reliably deter-
mined populations.

Gastrostyla minima HEMBERGER, 1985 (Fig. 203e-q, Table 48)

1985 Gastrostyla minima HEMBERGER, Arch. Protistenk., 130: 406 (Fig. 203f; see taxonomy; slides of the
type material are deposited in the Institut filr landwirtschaftliche Zoologie und Bienenkunde, University
Bonn, Germany).
1997 Gastrostyla minima HEMBERGER, 1985 - FOISSNER, Limnologica, 27: 225 (Fig. 203e, g-q; 8 voucher
slides with morphostatic and dividing, protargol-impregnated cells are deposited in the OberOsterrei-
chische Landesmuseum in Linz, Upper Austria).

Nom e n c I at u rea n d t a x 0 nom y: Previously, I classified Gastrostyla minima

as junior synonym of G. mystacea because of similar size and slightly anteriad displaced
transverse cirri (ForSSNER et al. 1991, p 270). However, ForSSNER (l997a) found speci-
 Gastrostyla 811

a
o
.....
,
..
..
/,
,.,-
.. ~

.,.,.,.
II

.....
~

"
~

..'
,.
.. ,.
".

..., I

,'
".

,.
".

., t
#

••
#

9 h

Fig.203e-h Gastrostyla minima (e, g, h, from FOISSNER 1997a; t: from HEMBERGER 1985. e, from life; t: h,
protargol impregnation; g, wet silver nitrate impregnation). e) Ventral view, 108 11m. C, h) Ventral infracilia-
ture and nuclear apparatus, f= 125 11m, h = 88 11m. Arrow in (f) marks postoral ventral cirrus IVI2. g) Nar-
rowly meshed silverline system and cortical granules rows (arrowheads); arrow marks cirral row; p 810.

mens in Bavarian running waters with a ciliary pattern that matches G. minima better than
G. mystacea, indicating that my synonymisation was premature. The FOISSNER popula-
tions have cortical granules, which are, however, not recognisable in life or after protar-
gol impregnation but stain deeply with methyl green-pyronin and silver nitrate. HEM-
BERGER studied only specimens impregnated with protargol and thus could not see this
special granules. FOISSNER (1997a) found further similar populations in soils of the USA
and the Galapagos Islands. These specimens lack cortical granules and thus would match
HEMBERGER'S description of Gastrostyla minima. However, they have only 3-4 cirri in the
post-peristomial portion of the frontoventral cirral row, which is dissimilar to G. minima.
 812 SYSTEMATIC SECTION

),
()

()
o ~~ ()

~1
~ ()
~
0""", ()

"':
()
, ~~ ()
"-, ()

""": ~~
.fu
()
()
()
()

~t.'
~
()
()
~ ~
~

1/ ~
q,1"
~
()
()
<' <'
()

'f <, <?

<'
-, .......... , .Y
Ii •

./ :.l' t)
<'
,..--' () t,i <' <'
.Y "OJ
t/-'o. <' '"
'" '"
Cl
()Q
<'
00 o c
0 o¢
001> k

Fig.203i-k Gastrostyla minima (i-k, from FOISSNER 1997a i-k, protargol impregnation). i) Dorsal ciliature
and nuclear apparatus, 108 11m. j) Ventral view of a very early morphogenetic stage with long, narrow oral pri-
mordium extending from left transverse cirrus to the enlarged postoral ventral cirrus. k) Early morphogenetic
stage. Arrow marks disintegrating enlarged postoral ventral cirrus; p 810.

These data indicate, that further, very similar species exist; thus, both descriptions of G.
minima are given separately.

M 0 r p hoi 0 g y and b i 0 log y: Type material after protargol impregnation

120-145 x 45-60!lm (HEMBERGER 1982, P 188; 1985). Outline oval. Two macronuclear
nodules, two micronuclei. No cortical granules mentioned (see taxonomy). Contractile
vacuole not observed, probably about in mid-body. Adoral zone of membrane lIes about
33 % of body length. 22-26 frontal, buccal, frontoventral, postoral, and pretransverse
ventral cirri. Transverse cirri (in life?) about 17 !lm long, slightly displaced anteriorly.
Marginal cirri (in life?) 15-17 !lm long. Bases of 3 posterior-most left marginal cirri re-
duced. "3 plus 2 x 112 dorsal kineties", indicating that an Oxytricha pattern (with only 1
dorsomarginal kinety) is present; possibly, HEMBERGER overlooked the very short kinety 6
(see Fig. 203i). Dorsal cilia 5-6 !lm long, last (posterior-most?) cilia slightly longer. In-
 Gastrostyla 813

m n

Fig. 2031-0 Gastrostyla minima (I-n, from FOISSNER 1997a I-n, protargol impregnation). I, m) Ventral cilia-
ture of an early and a middle morphogenetic stage. 0) Dorsal view of a middle morphogenetic stage. Parental
structures white, new structures black; p 810.

variably 3 inconspicuous, (in life?) 15-17 J.1m long caudal cirri, distinctly displaced to
right (in ventral view).
FOISSNER (1997a) population from Bavaria in life 70-120 x 25-40 J.1m. Prolate ellip-
soidal, ratio of length to width about 3: 1, left side more distinctly convex than right, both
ends evenly rounded, rather flexible. Invariably (n = 15) 2 macronuclear nodules, ellip-
soidal (1.5:1), rather close (mean = 6.7 J.1m, SD = 2.0 J.1m, CV = 30.4 %, Min = 3 J.1m,
Max = 10 J.1m, n = 15; protargol impregnation) together in middle third of body to left of
midline; nucleoli scattered, globular. 2-3 (mean = 2.1, n = 15) micronuclei, conspicuous
because rather large (after protargol impregnation 3-5 x 3-4 J.1m, mean = 3.6 x 3.2 J.1m, n
= 15) and compact, one each attached to macronuclear nodules in variable position. Con-
tractile vacuole slightly in front of mid-body. Cortex colourless, contains, as usual, nar-
rowly meshed silverline system (Fig. 203g). Cortical granules in narrowly spaced short
and long rows on ventral and dorsal side, do not stain with protargol but contrast as whit-
ish, punctate lines with dark brown pellicle in over-impregnated specimens; granules col-
814 SYSTEMATIC SECTION

ourless, about I /lm across, not recognisable in live specimens, even with interference
contrast optics, but stain dark blue with methyl green-pyronin and brown with silver ni-
trate (Fig. 203g). Cytoplasm colourless, contains some 1-4/lm long yellowish crystals
and food vacuoles 5-10 /lm across. Movement moderately rapid, scrabbling amongst de-
bris.
Adoral zone of membrane lIes about 36 % of body length. Buccal cavity rather nar-
row and flat, posterior right half covered by hyaline lip. Pharyngeal fibres distinct: In-
variably 3 frontal cirri, in life about 17 /lm long. Invariably (n = 15) 1 buccal cirrus near
at anterior end of parora!. Usually 1 cirrus slightly behind right frontal cirrus and left of
anterior end of frontoventral row (certainly homologous to cirrus III12 of I8-cirri oxytri-
chids; see Fig. 6a). Constantly 1 slightly enlarged cirrus close behind buccal vertex (cer-
tainly homologous to cirrus IV/2 of 18-cirri oxytrichids; see Fig. 6a). 9-15 (mean = 12.4,
SD = 1.9, CV = 15.2 %, n = 15), in life about 14 /lm long cirri in slightly sigmoid fron-
toventral row, which terminates usually about near mid-body in midline of cell (distance
between anterior end of cell and posterior end of frontoventral row after protargol im-
pregnation 37-55 /lm, mean = 42.1 /lm, SD = 5.1/lm, CV = 12.0 %, n = 15); frequently
some cirri strongly reduced in size and/or slightly out ofline. Invariably (n = 15) 2 pre-
transverse ventral cirri. Transverse cirri in life about 20 /lm long, close together, rather
distant from but projecting above posterior end; distance from posterior end of cell to
bases of posterior-most transverse cirrus after protargol impregnation 7-16 /lm (mean =
9.4 /lm, SD = 2.5 /lm, CV = 26 %, n = 15). However, the scanning electron micrograph
shows that transverse cirri do not protrude (Figure I7h in ForssNER 1997a). Marginal
rows open at posterior end, gap occupied by caudal cirri right of cell midline, right row
ends subterminally, left ends terminally in midline, posterior-most marginal cirri dis-
tinctly reduced in size, especially in left row. Dorsal cilia in life about 4 /lm long, kinety
6 (= outer dorsomarginal row) very short, that is, consists of2-4 dikinetids only.

M 0 r ph 0 g e n e sis: Ontogenesis is described by ForssNER (1997a, Fig. 203j-q). It

commences with the development of an oral primordium close to the frontoventral row.
Basal body proliferation occurs simultaneously in a long, narrow area between the buccal
vertex and the anterior-most transverse cirrus (Fig. 203j). Next, six cirral anlagen each
develop in the proter and opisthe (Fig. 203k, 1). The anlagen of the proter originate from
the anterior end of the paroral, the buccal cirrus, the cirrus left of the frontoventral row (=
cirrus 11112), and from the posterior cirri of the upper half of the frontoventral row. The
anterior cirri of the frontoventral row are homologous to the migratory (= frontoterminal)
cirri of the IS-cirri oxytrichids and are thus not involved in primordia formation. The an-
lagen of the opisthe originate from the oral primordium and the postoral ventral cirri, that
is, the prominent, isolated cirrus behind the buccal vertex (= cirrus IV12 of the IS-cirri
oxytrichids) and the posterior half of the frontoventral row, whose cirri are, as in the Oxy-
trichinae, completely incorporated into the oral primordium and the cirral streaks (Fig.
203k, I). In middle and late dividers cirri segregate within the anlagen. The cirri of the
three rightmost streaks arrange one behind the other: the anterior portion is made by an-
 Gastrostyla 815

.,.
o

Fig.203o-q Gastrostyla minima (from FOIssNER 1997a Protargol impregnation). 0, p) Ventral ciliature of a
late and a very late morphogenetic stage. Arrows in (P) mark movements of some cirri of streaks IV (anterior)
and V (posterior) forming the middle and posterior portion of the frontoventral row. Curved arrow denotes mi-
gration of the conspicuous postoral ventral cirrus, which is homologous with cirrus IV12 of the 18-cirri oxytri-
chids (cp. Fig. 6a). Figure 2030 does not show a typical specimen because in the proter 7 anlagen are present;
furthermore, streak IV of the opisthe has produced 3 cirri only. q) Dorsal ciliature of a late morphogenetic
stage. Dorsal morphogenesis is in Oxytricha pattern (cp. Fig. 24a). Parental structures white, new black; p 810.

lage VI, the middle portion by anlage IV, and the posterior portion of anlage V (Fig.
203m, 0, q). This pattern corresponds completely with the pattern of the I8-cirri oxytri-
chids, where the "frontoventral row" consists of 5 cirri only, namely (from anterior) VII4,
VII3, IV/3, V/4, and V/3 (Fig. 6a). The posterior-most cirrus of anlage IV (corresponding
transverse cirrus not considered) migrates behind the buccal vertex (Fig. 203p), identical
as cirrus IV/2 of the I8-cirri oxytrichids (cp. Fig. 1Oq). Thus, I am convinced that Gas-
trostyla is very closely related to the I8-cirri oxytrichids. The resemblance of Gastrostyla
with some amphisiellids, mainly caused by the frontoventral row and the "postperistomial
 816 SYSTEMATIC SECTION

cirrus" (EIGNER & ForSSNER 1994, FOISSNER 1997a), is either a convergence or the am-
phisiellids evolved from the oxytrichids. Dorsal morphogenesis is, as in other Gastrostyla
species, in Oxytricha pattern (Fig. 203i, n, q).

o c cur r e n c e and e colo g y: Locus classicus is a freshwater habitat in Peru;

however, HEMBERGER (1985) also found Gastrostyla minima in a soil sample from this re-
gion. FOISSNER (1997a) found it in two brooks in Bavaria, Germany, namely the ROslau
(periodically acidified, pH 3.9-6.6) and the Zinnbach (permanently acidified, pH 3.4 to
4.9). Both brooks are strongly shaded by spruces. Some physicochemical characteristics
from these clean running waters are given by ForssNER (1997a). Feeds on bacteria, dia-
toms, and Euglena sp. (ForSSNER 1997a).

Gastrostyia setifera (ENGELMANN, 1862) KENT, 1882 (Fig. 204a~)

1862 Pleurotricha setifera ENGELMANN, Z. wiss. Zoo!., 11: 385 (Fig.204a).

1866 Notlwpleurotricha setifera DIESING - DIESING, Sber. Akad. Wiss. Wien, 53: 118.
1882 Gastrostyla setifera, ENG. sp. - KENT, Manual infusoria II, p 784.
1889 Gasterostyla setifera ENGELM. sp. - Bo-rsclill, Protozoa, p 1248 (incorrect subsequent spelling of Gas-
trostyla).
1900 Gastrostyla setifera- WALLENGREN, Bih. K. svenska VetenskAkad. Hand!., 26: 11 (Fig. 204b).
1909 Gastrostyla setifera - FAURE-FREMIET, Arch. Protistenk., 13: 126 (Fig. 204c).
1932 Gastrostyla (pleurotricha) setifera (ENGELMANN, 1861) - KAHL, Tierwelt Dt!., 25: 595.
1972 Gastrostyla setifera (ENGELMANN, 1862) n. comb. - BORROR, J. Protozoo!., 19: 14 (see nomenclature).
1974 Pleurotricha setifera ENGELMANN - STILLER, Fauna Hung., 115: 107.

Nom e n c I a t u rea n d t a x 0 nom y: ENGELMANN (1862) assigned this species to

Pleurotricha, presumably because of the transverse cirri which are distinctly separated
into two groups, as in Pleurotricha lanceolata and P. grandis. However, KENr (1882)
recognised that the arrangement of the other cirri is as in Gastrostyla and thus transferred
it to this taxon. This was obviously overlooked by BORROR (1972a), who transferred it
once more to Gastrostyla. The illustrations by WALLENGREN (1900a; Fig. 204b) and
FAURE-FREMIET (1909; Fig. 204c) are probably redrawings from the original (Fig. 204a).
HEMBERGER (1982, p 187) found a population in a fre~:twater habitat in Peru. However,
the transverse cirri are not very distinctly separated into two groups, indicating that fur-
ther species exist. The GULATI (1925; Fig. 205e) population is insufficiently redescribed.
Detailed redescription necessary.
STOUT (1978, p 15, 17) recorded an "Oxytricha setifera" from irrigated soil in New
Zealand; since he did not mention the author of the species, it is uncertain if he observed
the present species, or ifhe made an incorrect subsequent spelling of Oxytricha setigera,
which is common in edaphic habitats.

M 0 r p hoI 0 g y and b i 0 log y: In life about 270 !lm long. Elliptical. Cell accord-
ing to ENGELMANN (1862) "formbestiindig", a term which is not identical with rigid or
 Gastrostyla 817

•
.
"VI \
'I
III ~
i III ~ :
:
i
,
• •
i ~.
'.~
"., ~i,.
\JV ii
" I
:"' ~.,~
\ \ ...:..........
, ~

; v, . .,\\
~ ~'" tIj; ~

• •
, ~ i)\. "j::
:I ,. t
I , ~::

:· f. !\l
•
:::
'
I
..
••

· ., l::
• t .,

• ,
,
I ::::

i j ~~
,: ;i ~ • ,
, : :
: .
;
'" b c
Fig.204a-(: Gastrostyla setifera (a, after ENGELMANN 1862; b, from WALLENGREN 1900a; c, from FAURE-
FREMIET 1909. 8-(:, from life). a-c) Ventral views, 260 )lm. Cirri which originate from the same streak are con-
nected by a dotted line (b). I-Vi = frontal-ventral-transverse cirri streaks I-Vi; P 816.

firm because he used it also to describe the consistency of the very flexible Urosomoida
agilis. Two postoral ventral cirri left of the frontoventral row. Transverse cirri distinctly
separate in two groups; only the 2 right transverse cirri protrude distinctly beyond poste-
rior end of cell. Dorsal cilia neither mentioned nor drawn indicating that they are short
(<5 Jlm).
According to WALLENGREN (1900a) 1,3,4,5,6,6 (in total 25) cirri originate from
frontal-ventral-transverse cirral streaks I-VI during morphogenesis (Fig. 204b). Frontov-
entral row consists of cirri from streaks VI, IV, and V (from anterior to posterior), as in
Gastrostyla dorsicirrata, G. minima, and G. steinii.

o c cur r en c e: Very rare species. Locus classicus is the town of Leipzig, Germany,
where ENGELMANN (1862) discovered Gastrostyla setifera in a lentic site covered with
Lemna sp. in the Elster River. DupLAKOFF (1933) mentioned it in a study on periphyton.
No further records known.
 818 SYSTEMATIC SECTION

Fig.20Sa Gastrostyla muscorum (from KAHL 1932). Ventral view from life, 160 /lm; p 808.
Fig.20Sb-e Species indeterminata (b) and insufficient redescriptions (c-e). b) Gastrostyla hebbalica (from
RAJASEKARASETIY & KAsTIJRI BAI 1961). Ventral view, various fixation and staining methods, 56/lm; p 824. c)
Gastrostyla hebbalica (from NAlDu 1965). Dorsal view after fixation, 65 /lm; p 824. d) Holosticha mystacea
(from VUXANOVICI 1963). Ventral view from life, 145 /lm; p 825. e) Gastrostyla setifera (from Gui.ATI 1925).
Ventral view from life, 90 /lm; p 824.

Marine species

Gastrostyla pulchra (PEREYASLAWZEWA, 1886) KAHL, 1932 (Fig. 206a-n)

1886 Stilonichia pulchra PEREYASLAWZEWA, Zap. novoross. Obshch. Estest., 10: 90 (Fig. 206a, g; incorrect
subsequent spelling of Stylonychia).
1886 Stylonychiapu/chra PEREYASLAWZEWA, Zap. novoross. Obshch. Estest., 10: 97.
1900 Gastrostyla sterkii WALLENGREN, 1900, Acta Univ. Lund., 36: 21 (Fig. 206b--d, f).
1932 Gastrostyla (Stylonychia) pulchra (PEREJASLAWZEWA, 1885) - KAHL, Tierwelt OtI., 25: 596 (Fig. 206e).
1933 Gastrostyla pulchra (PEREJASLAWZEWA 1885) - KAm., TierweIt N.- und Ostsee, 23: 112 (Fig. 206h).
1963 Gastrostyla pu/Chra (PEREJASLAWZEWA, 1885) - BORROR, Trans. Am. microsc. Soc., 82: 128 (Fig.
206i-k).
1970 Gastrostyla pu/Chra PEREJASLAWZEWA, 1885 - BURKOVSKY, Acta Protozooi., 8: 60 (Fig. 206I-n).
1972 Gastrostyla pulchra (PEREJASLAWZEWA, 1886) KAHL, 1932 - BORROR, J. Protozooi., 19: 14.
1973 Gastrostyla pulchra (PEREJASLAWZEWA, 1885) - HARTWIG, Abh. math.-naturw. K1. Akad. Wiss. Mainz,
18: 66.

Fig.206a-b Gastrostyla pulchra (a, from PEREYASLAWZEWA 1886; b-d, from WALLENGREN 1900b; e, from ~
KAHL 1932; f, after WALLENGREN 1900b from KAHL 1932; g, after PEREYASLAWZEWA from KAHL 1932; h, from
KAHL 1933. a, b, e-h, from life; c, sublimate fixation and borax-carmine staining; d, sublimate fixation and
haemalum staining). a, b, e-b) Ventral views, a, g, h = size not indicated, b = 170 /lm, e, f= 250 /lm. Arrow in
(b) marks the conspicuous single postoral ventral cirrus IV12. c, d) Nuclear apparatus.
 Gastrostyla 819

d
820 SYSTEMATIC SECTION

Nom e n c I at u rea n d t a x 0 nom y: PATIERSON et al. (1989, p 210) misdated

Gastrostyla pulchra with 1883, some other authors with 1885 (see list). However, ac-
cording to the Zoological Record and the first page of the "Zap. novoross. Obshch.
Estest., volume 10" 1886 is the correct year.
Synonymy of Gastrostyla pulchra (Fig. 206a) and G. sterkii (Fig. 206b) is very
likely, and indicated especially by the extremely anteriorly displaced transverse cirri and
the length of the frontoventral row. Synonymy was thus already supposed by KAHL
(1932), although a different number of macronuclear nodules was originally described.
However, both KAHL (1932, 1933) and BURKOVSKY (1970c) found specimens with 2 and 4
nodules, indicating intraspecific variability. Unfortunately, three important contributions
(BURKOVSKY 1970c, PEREYASLAWZEWA 1886, WAlLENGREN 1900b) are written not in Eng-
lish or Gennan, making a thorough study difficult and time consuming. I therefore used
only some morphometric data from these papers; most information presented below is
from KAHL (1932) and BORROR (1963b). Detailed redescription necessary because several
characters are only superficially (for example, dorsal infraciliature) or contradictorily (for
example, nuclear apparatus) described.

M 0 r p hoi 0 g y and b i 0 log y: In life 170 x 50 J.1m (WAlLENGREN 1900b),

170-250 J.1m long (I<AHL 1932), 140-210 x 55-67 J.1m (BORROR 1963b), 240-300 times
50-60 J.1ID and 160-200 x 50-60 J.1m (BURKOVSKY 1970c); PEREYASLAWZEWA (1886) gives
no size. Usually oblong, sometimes elliptical, margins slightly to distinctly converging
posteriorly. Right anterior margin usually slanted. Two or 4 macronuclear nodules, in life
about 32 x 15 J.1m (BORROR 1963b). Several micronuclei, in life about 10 J.1m across
(WALLENGREN 1900b; Fig. 206a, c-h, k, m, n). Contractile vacuole at about level of buc-
cal vertex (Fig. 206a, e, h). Cortical granules rod-shaped (KAHL 1932), according to BOR-
ROR (1963b) "211m long, oval, in bands between marginal cirri, clustered along median
edge of membranelles; one granule row 3-4 11m broad between marginal and ventral
cirri, and to right of right lateral cirri" (Fig. 206k, m). Food vacuoles 15-20 11m across,
remainder of cytoplasm darkly granular (BORROR 1963b). Moves hastily to and fro, some-
times torpid (K.AHL 1932).
Adoral zone of membranelles conspicuously overlapping on right margin, about 50
(BORROR 1963b) or 42-50 (BURKOVKSY 1970c) membranelles. Paroral kinety about
40 J.1m long. Frontoventral and especially transverse cirri rather prominent. Frontal cirri
in life 20 J.1m long. Frontoventral row tenninates somewhat behind level of buccal vertex,
consists of about 8 cirri (K.AHL 1932). One single postoral ventral cirrus (Fig. 206b, i, n),
very likely overlooked by PEREYASLAWZEWA (1886; Fig. 206a) and not illustrated in
KAHL's (1932, 1933; Fig. 206e, h) populations. Two pretransverse ventral cirri. Invaria-
bly 5 fimbriate, in life 25-30 11m long transverse cirri, insert at beginning of posterior
third to fifth of cell (according to BORROR 1963b 30-65 11m from posterior end), do not
protrude beyond posterior end of cell. Marginal rows overlapping posteriorly. At least 4
dorsal kineties (BORROR 1963b; Fig. 206j).
 Gastrostyla 821

-- ,
-
-- ,,
.•.'•

Fig.206i-n Gastrostyla pulchra (i-k, from BORROR 1963b; 1-0, from BURKOVSKY 1970c. i, j, I, n, Chatton-
Lwoff silver nitrate impregnation; k, m, haemalum staining). i, n) Ventral infraciliature, i = 150 ~m, n =
280 ~m. Arrows mark single postoral ventral cirrus IVf2 (cp. Fig. 206b). In (i) the transverse cirri are not very
distinctly displaced anteriorly; possibly a stain artefact. j) Dorsal infraciliature. Very likely dorsomarginal ki-
neties have been overlooked. k, m) Ventral views showing nuclear apparatus and cortical granules. I) Ventral
infraciliature of anterior cell portion; p 81S.

o c cur r e n c e and e colo g y: Marine; no record from inland salt waters. Locus
classicus is the Black Sea (PEREYASLAWZEWA 1886). Locus classicus of the synonym, Gas-
trostyla sterk;;, is the Baltic Sea at Malm6, Sweden (WALLENGREN 1900b).
Records largely substantiated by illustrations: mesosaprobic detritus from Heligo-
land, Sylt, and Kiel, Germany (KAHL 1932, 1933); mesopsammon of the Kandalaksha
Gulf, White Sea, and other marine sites in the USSR (BURKOVSKII & AzOVSKII 1985,
BURKOVSKY 1968, 1970a-c, 1971a, b, 1976, 1987, 1990, BURKOVSKY & EpsHfEIN 1982;
these papers include much ecological data, however, in Russian); Barents Sea (AzoVSKY
1996); detritus at the bottom of two sea water tanks filled with material from Alligator
Harbor, Florida, and New Hampshire tidal marshes, USA, Atlantic Ocean (BORROR
1963a, b).
822 SYSTEMA TIC SECTION

Records not substantiated by illustrations: periphyton of the Konigshafen, Sylt (Ger-

many), Northern Sea (KOSTERS 1974); Gulf of Naples, Italy, Mediterranean (NOBILl
1957); lagoon of Venice, Italy, Adriatic Sea (KrnSSELBACH 1936); mesopsammon near is-
land Jordsand, Denmark, Northern Sea (AzOVSKII 1987, HARTWIG 1973, 1974); polluted
intertidal areas of Kandalakshsky Bay, White Sea (AzoVSKY et al. 1996); mesopsammon
of the Peter the Great Bay, Japan Sea (MYSKOVA 1976); Gulf of Mexico (BORROR 1962).
Feeds on algae, mainly on diatoms (BORROR 1963b, FENCHEL 1968, KARL 1932); ac-
cording to BURKOVSKY et al. (1980) food vacuoles contained 30.8 % Protoceratium sp.,
30.5 % Peridinales, 0.7 % Navicula salinarium, and 11.3 % N. ammophila.

Gastrostyla stenocephala (BORROR, 1963) BORROR, 1972 (Fig; 207a-t)

1963 Oxytricha (Urosoma) stenocephala BORROR, 1963, Arch. Protistenk., 106: 512 (Fig. 207a-d; the
Chatton-Lwoff silver nitrate-impregnated holotype is deposited in the United States National Museum,
Smithsonian Institution, Washington, D. C.; CoRLISS 1972).
1968 Oxytricha stenocephala BORROR, 1963 - BORROR, Trans. Am. microsc. Soc., 87: 236 (Fig. 207a).
1972 Gastrostyla stenocephala (BORROR, 1963) - BORROR, J. Protowol., 19: 14.
1985 Gastrotyla stenocephala (BORROR, 1963) - ALAnRO LUBEL, An. Inst. BioI. Univ. Mex., 55: 28 (Fig.
207f; incorrect subsequent spelling of Gastrostyla).
1990 Gastrostyla stenocephala (BORROR, 1963) - ALOORO LUBEL, MARTINEZ-MURIllO & MAvEN ESTRADA,
Manual de ciliados, p 137 (Fig. 207e).

Nom e n cia t u rea n d t a x 0 nom y: Oxytricha stephanocephala BORROR, 1963

in HARTWIG (1980, P 128) is an incorrect subsequent spelling. The ventral infraciliature of
this species is in fact reminiscent of Gastrostyla; however, a detailed redescription is
needed to clarify the systematic position finally. Recently transferred to Hemigastrostyla
by SONG & WILBERT (l997a; see Addendum 5).

M 0 r p hoi 0 g y: In life 160-220 x 37-44 Ilm (BORROR 1963a), 194 x 42 Ilm (ALAoRO
LUBEL 1985, ALAoRO LUBEL et al. 1990). Cephalised region about 20 x 20 Ilm. Margins
almost in parallel, anterior end abruptly narrowed (that is, more or less cephalised), pos-
terior narrowly rounded and usually distinctly curved to the right, which is often not visi-
ble in specimens under coverslip as body is extremely fragile, often losing posterior end
prior to examination. Gastrostyla stenocephala may fragment within minutes following
removal from its natural habitat, making observation rather difficult. Ventral side flat,
dorsal one rounded. Macronuclear nodules in life about 20 x 10 Ilm, each with an adja-
cent, in life about 6 x 4 Ilm sized micronucleus. No contractile vacuole. Ectoplasm 1 Ilm
thick, clear. Rosettes of cortical granules at bases of dorsal cilia and some cirri. Cyto-
plasm filled with remains of diatoms and small, oval, 1 Ilm sized bodies. Extremely thig-
motactic, adhering with the aid of caudal cirri and posterior dorsal cilia. Bends ventrally
but not laterally, when swimming freely.
Adoral zone of membrane lies only about 25 % of body length, about 40 mem-
braneIles. Anterior membrane lIes in life 25 /lm long, covered basally by clear ectoplas-
 Gastrostyla 823

Fig. 207a-f Gastrostyla stenocephala (a-d, from BORROR 1963a; e, from ALADRO LUSEL et aI. 1990; f, from
ALADRO LusEL 1985. a, d, shape from life and ciliature after Chatton-Lwoffsilver nitrate impregnation; b, e, f,
from life; c, Chatton-Lwoff silver nitrate impregnation). a, e, 1) Ventral view, a = 200 ~m, e = 215 ~m, f =
195 ~m. Note the abruptly narrowed (cephalised) anterior end. Arrow in (a) marks postoral ventral cirrus. b)
Ventral view showing nuclear apparatus, pharyngeal fibres, and ingested food, 200 ~m. C, d) Ventral and dor-
sal infraciliature, c = 146 ~m, d = 200 ~m. Tiny arrow in (c) = postoral ventral cirrus, large arrow = cirri right
of right transverse cirri, arrowhead = caudal cirri. See text for details; p 822.

mic plate. Paroral (kinety?) about 32 11m long, conspicuous. Endoral on bottom of buccal
cavity. Cytophatynx in life at least 20 11m long. Arrangement of cirri as in Figure 207a, c.
Frontoventral row consists of about 8 cirri, usually 2 distinctly separated cirri behind
frontal row. According to original description a single cirrus immediately behind buccal
vertex (Fig. 207a, c, tiny arrow) which is very likely homologous to the prominent pos-
toral ventral cirrus of the other Gastrostyla species; however, it is lacking in populations
described by ALADRO LUBEL (1985; Fig. 207f) and ALADRO LUBEL et al. (1990; Fig. 207e).
Five distinctly enlarged, in life about 25 11m long transverse cirri, overlapping posterior
end by almost half their length. Marginal cirri in life about 12 11m long, a fibre extends
12 11m anteriorly from median edge of right marginal cirri. 4-6 dorsal kineties; however,
such a high variability is very unlikely, indicating misobservation by BORROR (1963a).
824 SYSTEMATIC SECTION

Dorsal cilia in life 16 flm long. Two extremely fimbriate caudal cirri to right of posterior
end (in ventral view; Fig. 207c, arrowhead). Figure 207c (large arrow) shows, as in Oxy-
tricha enigmatica, a pair of cirri right of right transverse cirri (details, see Addendum 5).

o c cur r e n c e and e colo g y: Marine. Locus classicus is Alligator Harbor, Flor-

ida, USA, Atlantic Ocean (BORROR 1963a, 1968). Gastrostyla stenocephala occurred
regularly and was vel)' common in sand, especially in the uppermost layer where pore
spaces may have been large enough to permit its entry, and where the diatom population
was highest; did not occur in diatom detritus, although feeding exclusively on diatoms
(BORROR 1963a, FENCHEL 1968). Highest abundance (13 indo ml-1) at late January. Re-
cords substantiated by illustrations: in sediment (0-2 cm) of the Thalassia testudinum
zone at Enmedio Island (Gulf of Mexico near Veracruz) and other sites in Mexico (ALA-
DRO LUBEL 1985, ALAnRO LUBEL et al. 1990).

Species indeterminata
Gastrostyla hebbalica RAlASEKARASETIY & KAsTURI BAl, 1961, J. zool. Soc. India, 13:
238 (Fig. 205b).Remarks: In life 60-80 x 30-35 11m. Ovoid. Three (?) spherical macro-
nuclear nodules, 2 micronuclei. Contractile vacuole in right posterior portion of cell.
Three enlarged frontal cirri. Ventral cirri irregularly arranged. Five transverse cirri. Some
observations (for example, position of contractile vacuole) indicate that it is not a valid
species. Freshwater in Hebbal, Bangalore, India.

Insufficient redescriptions
Gastrostyla hebbalica RAlASEKHARASETIY & KAsTURlBAl, 1961 - NAlDu, 1965, Hydrobio-
logia, 25: 560 (Fig. 20Sc). Remarks: Freshwater in Vijayawada, India.
Gastrostyla (Stylonychia) pulchra (PEREJASLAWZEWA, 1886) WAHLGREN, 1890 - JONES,
1974, Univ. South Alabama Monogr., 1: 41 (Fig. 242j). Remarks: Transverse cirri not
displaced anteriorly! Mobile Bay, USA.
Gastrostyla seti/era ENGELMANN - GULATI, 1925, J. Bombay nat. Hist. Soc., 30: 752 (Fig.
205e). Remarks: The transverse cirri are not separated into two distinct groups. In life
90 x 30 flm. Small pond near Data Ganj Bakhash, India.
Gastrostyla sp. - LEPSI, 1927, Studii Cerc. Acad. RPR, 12: 122 (Fig. 226k). Remarks: In
life 76 x 30 flm. Among algae_ Romania.
Gastrostyla steini? - SCHNEIDER, 1930, Arch. Protistenk., 72: 523 (Fig. 233h). Remarks:
Incorrect subsequent spelling. Rather invariably 6 macronuclear nodules. After addition
of iodine ink some specimens excreted a homogeneous mucous layer. Infusions of grass.
Germany?
Holosticha mystacea STEIN, 1859 - CHORIK, 1968, Free-living ciliates, p 130 (Fig. 233e).
Remarks: Possible a redrawing of KARL's (1932) redrawing (cp. Fig. 203d). Freshwater
in Moldova.
 Kerona 825

Holosticha mystacea STEIN, 1859 - VUXANOVICI, 1963, Studii Cerc. BioI., 15: 208 (Fig.
205d). Remarks: Certainly not identical with Gastrostyla mystacea since it obviously has
more than 2 macronuclear nodules. Stagnant water body in Bucharest, Romania.

Kerona MULLER, 1786

1786 Kerona MOLLER, Animalcula Infusoria, p 233 (see nomenclature) - Type (subsequent designation by
EHRENBERG 1838; see nomenclature): Cyclidium pediculus MOLLER, 1773.
1838 Kerona - EHRENBERG, Infusionsthierchen, p 368.
1852 .Alastor PERlY, Mitt. naturf. Ges. Bern, year 1852: 64 - Type (by monotypy): Kerona polyporum
EHRENBERG, 1835.
1852 Alastor PERlY, Zur Kenntniss kldnster Lebensfonnen, piSS.
1859 Kerona. EHRBG. - STEIN, Organismus der Infusionsthiere I, p 171.
1882 Kerona, EHRENBERG- KENT, Manual infusoria II, p 763.
1932 Kerona EHRENBERG, 1838 - KAHL, Tierwelt Otl., 25: 569.
1972 Kerona EHRENBERG, 1835 - BORROR, J. Protozool., 19: 9.
1974 Kerona EHRENBERG - STILLER, Fauna Hung., lIS: 44.
1982 Kerona EHRENBERG, 1825 - HEMBERGER & WILBERT, Arch. Protistenk., 125: 269.

C h a r act e r i sat ion: Undulating membranes almost straight and in parallel. Numer-
ous frontal and frontoventral cirri arranged in 6 slightly to distinctly curved rows. Usually
5 transverse cirri. One left and 1 right row of marginal cirri almost confluent posteriorly.
Numerous dorsal kineties covering dorsal side almost completely. Caudal cirri present.
Six frontal-ventral-transverse cirral streaks segregating numerous frontal and frontoven-
tral cirri. Three dorsal primordia, each showing mUltiple fragmentation and 2 dorsomargi-
nal kineties. Ectocommensal on hydras and bryozoans.

Nom e n c I a t u r e: Alaster polyporum PERTY, 1852 in HEMBERGER & WILBERT (1982,

P 262) is an incorrect subsequent spelling. There is great confusion about Kerona, which
was established by MOLLER (1786) comprising 14 species ofhypotrichous ciliates. How-
ever, Kerona pediculus was not included. In EHRENBERG'S early days one of these 14 spe-
cies was in Kerona, namely K. pustulata (see EHRENBERG 1838 for details of the time be-
tween MOLLER and EHRENBERG). EHRENBERG (1835) transferred this last species to Sty/ony-
chia. Simultaneously, he introduced the name Kerona polyporum for the hypotrichous
ciliate living on hydras. Thus, Kerona sensu EHRENBERG was differently defined than
Kerona sensu MOLLER (which was almost not defmed). This caused later workers (for ex-
ample, STEIN 1859b, KENT 1882, KAHL 1932) to treat EHRENBERG as the author and thus
the founder of Kerona. However, this is incorrect because EHRENBERG never establish a
genus Kerona. Considering EHRENBERG as the author of Kerona is of course unacceptable
because it would be the junior homonym of Kerona MOLLER. There are two ways to solve
this nomenclatural problem: (i) MOLLER (1786) is accepted as the author of Kerona with
Cyclidium pediculus MOLLER, 1773 as type species, ignoring the problem that none of
MOLLER'S species is now in Kerona. (ii) The synonym, Alastor PERTY, 1852 is used which
826 SYSTEMATIC SECTION

was expressly established for Kerona polyporum EHRENBERG and Cyc/idium pediculus
MOLLER. This solution was proposed by JANKOWSKI (1979, P 49; see also CURDS et al.
1983, p 426). I prefer the first proposal, mainly to preserve the well established name
Kerona. Furthermore, Alastor PERTY, 1852 is a nomen oblitum (CORLISS 1979, P 208).
The taxonomy of Kerona is discussed in the relevant chapter ofthe single species.

Single species

Kerona pediculus (MOLLER, 1773) BLOCHMANN, 1886 (Fig. 208a-y, 209a, b)

1773 Cyclidium pediculus MOLLER, Vennium Terrestrium et Fluviatilium, p 54 (no illustration).

1786 Cyclidium pediculus - MOLLER, Animalcula Infusoria, p 84, Tab. XI, Fig. 15-17 (see taxonomy).
1835 Kerona polyporum EHRENBERG, Abh. preuss. Akad. Wiss., year 1835: 164 (no illustration).
1838 Kerona polyporum EHRENBERG, 1835 - EHRENBERG, Infusionsthierchen, p 368 (Fig. 208c, d).
1850 Kerona polyporum EHRENBERG - DIESING, Systema Helminthum I, p 161.
1852 Alastor polyporum - PERTY, Zur Kenntniss Ideinster Lebensfonnen, p 155.
1859 Kerona polyporum. EHRBG. - STEIN, Organismus der Infusionsthiere I, p 173 (Fig. 208a).
1860 Kerona polyporum - BALBIANI, Journal de la physiologie de l'homme et des animaux, 3: Planche IV,
Fig. 16-18.
1882 Kerona polyporum, ERR. - KENT, Manual infusoria 11, p 763.
1886 Kerona pediculus O. F. MOLL. - BLOCHMANN, Mikroskopische Thierwelt, p 76.
1930 Kerona polyporum EHRENB. - CAY ALLINI, Archo zool. ital., 14: 1 (Fig. 208j-1).
1932 Kerona polyporum EHRENBERG - KMu., Tierwe1t Dtl., 25: 569, Fig. 8614 (redrawing from STEIN 1859b).
1933 Kerona pediculus (0. F. MOLLER) EHRENBERG - WANG & NIE, Contr. bioI. Lab. Sci. Soc. China, 10: 53
(Fig. 208b).
1965 Kerona - HiLL, Rep. Oundle Sch. nat. Hist. Soc., 1965: 95, Fig. 2 (low quality sketch).
1973 Kerona polyporum - SCHLICK, Mikrokosmos, 62: 201, Bild 4 (low quality illustration).
1974 Kerona polyporum EHRENBERG - PATSCH, Arb. Inst. landw. Zool. Bienenkd., 1: 54 (Fig. 208h).
1974 Kerona polyporum EHRENBERG - STILLER, Fauna Hung., 115: 44.
1975 Kerona polyporum - WILBERT, Mikrokosmos, 64: 175, Bilder 7, 8 (micrographs of ventral and dorsal
infraciliature after protargol impregnation).
1982 Kerona polyporum EHRENBERG, 1835 - HEMBERGER & WILBERT, Arch. Protistenk., 125: 261 (Fig.
208e--g, n-v, X, y).
1982 Kerona polyporum EHRENBERG, 1835 - HEMBERGER, Dissertation, p 62 (Fig. 208e--g, n-y)
1983 Peritromus hydrarum CHARDEZ, Protistologica, 19: 231 (Fig. 209a, b; new synonym).
1985 Kerona polyporum EHRENBERG, 1838 - FLEURY, !FrODE, DEROUX & FRYD-VERSAVEL, Protistologica, 21:
508 (Fig. 208i).
1985 Kerona polyporum - SCHNEIDER, Mikrokosmos, 74: 313, Bilder 1-5 (micrographs ofliving specimens).
1987 Kerona pediculus (0. F. MOLLER, 1773) nov. comb. - FOISSNER, Arch. Protistenk., 133: 225 (see no-
menclature).
1991 Kerona pediculus (MUELLER, 1773) BLOCHMANN, 1886 - FOISSNER, BLATTERER, BERGER & KOHMANN, In-
fonnationsberichte des Bayer. Landesamtes fur Wasserwirtschaft, 1191: 265.

Nom e n c I a t u rea n d t a x 0 nom y: Incorrect subsequent spellings: Kerona

polymorphum (FOISSNER 1979b, P 3); Keronapolypora (ESTEVE 1982, P 519).
 Kerona 827

MOLLER (1786) provided three small illustrations which rather distinctly show a hy-
potrichous ciliate living on hydras. EHRENBERG (1838) described Kerona polyporum as-
suming that MOLLER'S (1773, 1786) Cyclidium pediculus is the peritrichous ciliate also
living on hydras. EHRENBERG'S error was recognised already by STEIN (1859b), however,
without suppressing the junior synonym, Kerona po/yporum EHRENBERG. Thus, the name
K. polyporum was used until recently, when FOISSNER (1987d) reintroduced the older
name, K pediculus, obviously overlooking that Cyclidium pediculus had already been
transferred to Kerona by BLOCHMANN (1886; see list of synonyms).
Kerona pediculus is usually classified in the Keronidae DUJARDIN, 1841 (for example,
CoRLISS 1979, HEMBERGER & WILBERT 1982, SMALL & LYNN 1985, TUFFRAu 1987, TUF-
FRAU & FLEURY 1994). However, the morphogenesis shows the main apomorphy of the
oxytrichids, namely fragmentation of dorsal kineties (Fig. 208w). I thus include it in the
oxytrichids, assuming that parallel evolution of this conspicuoUs character is rather un-
likely. Furthermore, the frontal-ventral-transverse cirri originate from six longitudinal
streaks (Fig. 208s, t) like in the 18-cirri oxytrichids. The inclusion of Kerona in the oxy-
trichids requires the assumption of parallel or convergent evolution of a corona of frontal
cirri because there is no evidence that the second taxon of the Keronidae, Keronopsis PE-
NARD, 1922, is also an oxytrichid. Keronopsis (and its sister-group or synonym, Para-
holosticha KARL, 1932) divides in cysts and lacks fragmentation of dorsal kineties
(DIECKMANN 1989), indicating that it branches outside the oxytrichids.
As in some other taxa (for example, Gastrostyla, Paraurostyla), I was unable to find
the sister-group of this presumably highly specialised species. It thus is classified as taxon
of unknown systematic position in the oxytrichids. The multiple fragmentation in all 3
dorsal kinety anlagen (Fig. 208x, y) reminds of Onychodromus quadricornutus (FOIssNER
et al. 1987b). This species also has several frontal cirri, which, however, originate from
several streaks, whereas in Kerona they are (largely) the descendants of the leftmost
streak, that is, the left frontal cirrus is split (see below). Furthermore, lack of cortical
granules, the (rather) inflexible body, the long (50 % of body length) adoral zone of
membrane lIes, and some details of the ventral morphogenesis indicate that Kerona possi-
bly belongs to the Stylonychinae, implying that the number of frontal-ventral-transverse
cirri increased secondarily. The confluent marginal rows are reminiscent of Histriculus,
the undulating membranes are almost in Stylonychia pattern. EIGNER (1997) classified
"Kerona polyporum" in the Parakahliellidae.
The synonymy of Peritromus hydrarum CHARDEZ, 1983 and Kerona pediculus is
rather evident, especially concerning the body shape, the number of adoral membraneUes,
and the habitat, namely freshwater hydras. The semischematic illustration after silver im-
pregnation probably shows the dorsal kinety pattern (Fig. 209a, b).
Kerona pediculus is very easily identified by the kidney-shaped body, the unique in-
fraciliature, and especially the extraordinary habitat. Kerona polyporum sensu DUMAS
(1929) and FROMENTEL (1876) are insufficiently redescribed.
 828 SYSTEMATIC SECTION

... '

.... , .... ."

....
" ~
' .. "

: ; ,.' ~ ~ .

- e
Fig.208a-g Keronapediculus (a, from STEIN 1859b; b, from WANG & NIE 1933; c, d, from EHRENBERG 1838;
9

e-g, from HEMBERGER & WILBERT 1982. a-d, from life; e-g, protargol impregnation). a, b, d) Ventral views, a
= 130-205 Jlm, b = 150 Jlm, d = up to 170 Jlm. c) Several specimens (arrows) creeping on a hydra. e-g) Dor-
sal and ventral infraciliature and nuclear apparatus, e, f = 190 Jlm. Arrow in (e) marks caudal cirri, small ar-
rows in (f) mark interruption in frontoventral rows 3 and 4. Large arrow marks buccal cirri. 1-6 = frontoventral
rows 1-6 (very likely homologous to the circal streaks I-VI of 18-cirri oxytrichids, cp Fig. 6a); p 826.
 Kerona 829

Fig. 208h, i Kerona pediculus (h, from PATSCH 1974; i, from FLEURY et a1. 1985. h, i, protargol impregnation).
Ventral infraciliature, h = ISO J!m, i = 140 J!m. The specimen in (h) has only 4 transverse and 2 caudal cirri.
BC = buccal cirri, RMR = right marginal row, TC = transverse cirri, uM = undulating membranes, l...{i = fron-
toventral cirral rows l...{i; p 826.

M 0 r p hoi g y and b i 0 log y: In life 120-205 x 70-110 ~m, usually about

0
180 x 95 ~m; 130-150x 70-80 ~m (CAVALLINI 1930); 150-180 ~m long (PATSCH 1974),
150 x 80 ~m on average (WANG & NIE 1933), 120 ~m long (CHARDEZ 1983). Kidney or
bean-shaped, that is, right margin convex, left deeply indented centrally, anterior end
evenly, posterior narrowly rounded to slightly pointed; plano-convex, strongly flattened
dorso-ventrally. Margins rather thin and flexible, attach to body of hydras, remaining
body of Kerona, however, rather firm than flexible, but not as rigid as, for example, in
Stylonychia. Two macronuclear nodules (16-17 ~m long; CHARDEZ 1983), about in me-
dian of cell; each nodule with a micronucleus (about 5 ~m across; CHARDEZ 1983). Con-
tractile vacuole about at level of buccal vertex. Cortical granules lacking. Rapid move-
ment, about 500 !lm S·l (BULLINGTON 1925).
Adoral zone of membrane lIes about 50 % of body length, arranged at body margin,
except proximal portion which points to mid-body. 30-74, usually around 60 adoral
membranelles; 30-35 (PATSCH 1974),35-60 (WARREN & ROBSON 1986),60-64 (HEMBER-
GER & WILBERT 1982), 68-74 (CHARDEZ 1983). Undulating membranes slightly curved.
Usually 4 (2-6) slender buccal cirri immediately in front of undulating membranes;
 830 SYSTEMATIC SECTION

Fig. 208j, k Kerona pediculus (from CAVAlLINI 1930. j, k, from life). j) Hydra sp. with many specimens of K.
pediculus. k) Ventral view showing arrangement of cirri and nuclear apparatus, about 130 11m; p 826.
 Kerona 831

Fig.2081 Kerona pediculus (from CAY ALLIN' 1930). Hydra sp. with many specimens of K. pediculus; p 826.
 832 SYSTEMATIC SECTION

Fig. 208m Kerona pediculus (original scanning electron micrograph kindly supplied by C. BAROELE, Univer-
sity of Tuebingen, Germany). Ventral view showing the arrangement of cirri, about 75 J.lm. Arrow marks the
almost straight paroral. AZM = adoral zone of membranelles, TC = transverse cirri; p 826.

sometimes they are almost not separated, giving the impression of a large buccal cirrus.
Six slightly to distinctly curved rows of frontal and frontoventral cirri which very likely
correspond (are homologous) with the cirri originating from the 6 primordia of the 18-
cirri oxytrichids (for arrangement and numbering, see Fig. 208t). Row 1 with about 18
 Kerona 833

(17-19) cirri, homologous with frontal cirrus III of the 18-cirri oxytrichids; row 2 about
15-16 cirri, homologous with middle frontal cirrus; row 3 with 17-18 cirri, usually inter-
rupted and thus possibly homologous to right frontal cirrus (11113) and frontoventral cirrus
11112 of the 18-cirri oxytrichids; row 4 with 20-21 cirri, usually slightly interrupted at
level of buccal vertex; row 5 usually with 27 (25-29) cirri, terminates near left transverse
cirrus; row 6 usually with 25 cirri (25-27), begins about at level of buccal vertex, termi-
nates at right transverse cirrus. Five prominent transverse cirri, becoming larger from left
to right (Fig. 208m); according to PATSCH (1974) only 4 transverse cirri. Left marginal
row with about 40 (38-45), right one with about 62 (60-67) cirri, almost confluent poste-
riorly; marginal cirri distinctly longer than frontoventral cirri. Dorsal cilia 2-4 Jlm (WAR-
REN & ROBSON 1986), according to HEMBERGER & WILBERT (1982) 5-6 Jlm long, arranged
in numerous (about 16-20) indistinct rows, that is, dorsal side completely ciliated (Fig.
208e). Three (HEMBERGER & WILBERT 1982) or2 (PATSCH 1974) caudal cirri very closely
arranged at posterior end of cell (Fig. 208e, f). Some ultrastructural details are shown in
FLEURY et al. (1985).
M 0 r p hog e n e sis of cell division is described by WICKLOW (1979), HEMBERGER
(1982; Fig. 208n-y), and HEMBERGER & WILBERT (1982). It commences with the forma-
tion of an distinctly invaginated oral primordium immediately behind the buccal vertex
(Fig. 2080, 0). Simultaneously 2 primordia are formed right and left of the anterior end of
the undulating membranes (Fig. 2080). Somewhat later one primordium each occurs
within the parental rows 3 and 4 (Fig. 208p). The primordium in row 4 splits up into 3
streaks to form the frontoventral transverse primordia IV-VI of the proter, which is remi-
niscent of Stylonychia pustulata (Fig. 161g). In the opisthe the primordia V and VI origi-
nate from the middle portion of the parental row 5 (Fig. 208r). The exact origin of the
other frontal-ventral-transverse cirri streaks is difficult to ascertain (Fig. 208r, s) but very
likely it proceeds rather similarly to in 18-cirri oxytrichids, for example, Stylonychia pus-
tulata. A late stage shows 6 streaks in both the proter and the opisthe, each forming a
rather high number of cirri. Row 6 does not, as in the l8-cirri oxytrichids, participate in
primordia formation. The cirral rows originating from streaks II-IV are distinctly sepa-
rated; the posterior part of streak II forms the buccal cirri (Fig. 208g). New marginal rows
originate, as in Onychodromus quadricornutus, distinctly right of the parental rows,
which are completely retained in late stages (Fig. 208s, t).
Dorsal morphogenesis commences, as is usual, with the formation of 3 longitudinal
anlagen in both the anterior and posterior half of the cell. Because of the complex inter-
phasic pattern it is impossible to ascertain if a correlation with certain parental kineties
exists (Fig. 208v). Later, all anlagen begin with multiple fragmentation, forming large
fields of usually parallel kinety fragments. Simultaneously, 2 dorsomarginal kineties oc-
cur (Fig. 208w-y). In interphasic specimens, the kinety fragments cover the dorsal side
almost completely (Fig. 208u). One caudal cirrus originates at the posterior end of each
of the 3 dorsal anlagen.
Division of nuclear apparatus is described by BALBIANI (1860a) and conjugation is
figured and described by BALBIANI (1861, 1862).
 834 SYSTEMATIC SECTION

p -- ." ,,
....
...

... ,
..... .-.. ,, ,

)0
\ . ,,. '", ..

Q
... ,, .-
..-.- " '" "
,, "~
,
," I, <0SS&
'<:::::;

..,
.,
,
•
,,
,
",
\ \
,
" ""
"

\ ..
\ t?~"

00

Fig.208n-q Kerona pediculus (from HEMBERGER 1982. n-q, morphogenetic stages in ventral view after pro-
targol impregnation). n) Very early stage, about 90 J.lm. Arrow marks oral primordium. 0) Early stage. Arrows
mark 2 primordia left and right of the buccal row. They become the primordia I and II of the proter. p) Early
stage showing one primordium each in row 3 and 4. q) Middle stage showing 2 primordia right of the oral pri-
mordium; p 826.
 Kerona 835

.'. _.- u

Fig.20Sr-u Kerona pediculus (from HEMBERGER 1982. r-t, morphogenetic stages in ventral view after protar-
got impregnation; u, protargol impregnation). r) Middle stage showing 6 primordia both in the proter and the
opisthe. Arrows mark rightmost anlagen (3 in the proter, 2 in the opisthe) originating from frontoventral row 4
(proter) and 5 (opisthe). This pattern is reminiscent of the 18-cirri oxytrichids like Sty/onychia pustulata (cp
Fig. 161g). s, t) Late and very late stages showing 6 primordia and segregation of cirri. Transverse cirri became
larger from left to right. Note that new marginal cirral rows originate right of parental ones which are almost
completely retained even in late stages. u) lnterphasic pattern of dorsal infraciliature; p 835.
 836 SYSTEMATIC SECTION

\I
/ }

j
I I

w
v

~
"
1;':-
."".
.\

p\
-7
!
/ I:
l ,1/ t

...;
\
"
,
\ fl
\,
:1,
,:~ ;:1.
i fJ
\ \I

x y

Fig.208v-y Kerona pediculus (from HEMBERGER 1982. v-y, morphogenetic stages in dorsal view after protar-
gol impregnation). v) Early stage with 3 longitudinal anlagen for both the proter and the opisthe. Parental basal
body pairs omitted. w) Anlagen 2 and 3 begin with fragmentation. x) All anlagen show multiple fragmentation.
Arrows mark dorsomarginal kineties. y) Each primordium produces several kineties which later cover the
whole dorsal surface (cp Fig. 208u); p 826.
 Kerona 837

Occurrence and eco-

log y: Ectocommensal on fresh-
water hydras (Hydra attenuata,
H. circumcincta, H. fusca, H.
vulgaris, H. viridissima; Fig.
208c, j, 1) and thus often in com-
pany with the peritrich, Tricho-
dina pediculus (for example,
BRECKENFELD 1886, ForssNER et
al. 1991, 1992c, HILL 1965,
KEISER 1921, MEuCHE 1939,
PATSCH 1974, SCHNEIDER 1994);
but also on the bl)'ozoans Plu-
matella and Cristatella mucedo
(WARREN & ROBSON 1986, 1987,
SVEC 1897); single specimens
sometimes swim freely or creep
near the host. Obviously prefers
the body to the arms of the hy-
dras (SCHLICK 1973, SCHNEIDER
1985). Fig. 209a, b Peritromus Irydrarum (from CHARDEl 1983a. a, b,
Locus classicus is vel)' likely silver impregnation, schematic). This is very likely a synonym
Denmark, where MOLLER (1773, of Kerona pediculus. The dotted lines are probably the dorsal
infraciliature; p 826.
1786) discovered Kerona pedicu-
lus on Hydra pallida. Locus clas-
sicus of the synonym Kerona
polyporum is Berlin, Germany, where EHRENBERG (1838) discovered it with high abun-
dance on Hydra vulgaris and H. oligactis. CHARDEZ (1983) found the synonym Peritro-
mus hydrarum on the same hydras in a pond in the village ofWaremme, Prov. de Liege,
Belgium, in January at 4--6 °C and pH 7.6.
Records substantiated by illustrations: Mare Nozay, a pond in France? (FLEURY et al.
1985); vel)' abundant on Hydra viridissima in a trout pond in Germany (?) and also vel)'
abundant on H. fusca and H. grisea in Prague, Czechoslovakia (STEIN 1859b); meso-
saprobic brook in Germany (PATSCH 1974); numerous on Hydra attenuata and H. viridis-
sima in a clay-pit in Germany (SCHNEIDER 1985); Italy (CAVALLINI 1930); rarely in fresh-
water in Nanking, China (WANG & NIE 1933).
Records not substantiated by illustrations: oligotrophic and eutrophic ponds in Aus-
tria (KlwNER 1988, own observations); on Hydra attenuata in a side branch of the Da-
nube River in Vienna, Austria (VORNATSCHER 1938; further records from Austria: AOOLR
1995c, BLATTERER 1994, DALLA TORRE 1891, MITIS 1941); rather abundant on bl)'ozoans
and hydras in a pond in Czechoslovakia during autumn (SVEC 1897); saprobic brook near
Prague and Mo1dau River, Czechoslovakia (BuCHAR 1957, KALMus 1928); England
838 SYSTEMATIC SECTION

(CRAIGIE 1921); mesosaprobic running waters in Germany (HEuss 1976, SCHMITZ 1986);
Hamburg Harbour, Germany (BARTSCH & HARTWIG 1984); lake (ploner See) in Germany
(ZACHARIAS 1893, 1894); unpolluted foothill stream (Breitenbach) near the village of
Schlitz, Germany (PACKROFF & ZWICK 1996); numerous and in masses in oligosaprobic
and oligosaprobic to betamesosaprobic reservoirs in Germany (NUSCH 1970; further re-
cords from Germany: HENDERSON 1905, SONG & WILBERT 1989, WETZEL 1928b, WILBERT
1969); on Hydrajusca in Lake Balaton and in Budapest, Hungary (FRANcE 1897, KRE-
PUSKA 1917); beta- to alphamesosaprobic running water in Italy (MAnoN! & GIIETri 1980;
further records from Italy: CANELLA 1954, LaNGill 1895); Switzerland (ANDRE 1912);
Volga Basin and reservoirs in the USSR (KRAvCHENKO 1969, MAMAEVA 1979b, MYLNIK-
OVA 1981, ZYKOFF 1903); at 13°C abundantly on Hydra in Lake Baika~ USSR (GAJEVS-
KNA 1927, GAJEWSKNA 1933); rare in a pond in the botanical gardens of the Pennsylvania
University, USA, in March and October 0NANG 1928); New Zealand (MAsKELL 1887b).
It was originally presumed that K pediculus preyed upon the living tissues of the
polyp which serves it as host, the characteristic nematocysts of the hydras being fre-
quently found inside it. These cells, however, are only incepted with other waste matter
thrown off from the surface of the polyp's integument, and in the removal of which
Kerona undoubtedly plays the role of a useful scavenger. Other food, especially algae
(Euastrum, Euglena, Phacus, Trachelomonas, desmids, diatoms like Amphora, Fragi-
laria, Navicula), are devoured with equal avidity, and form an important addendum to its
customary bill of fare (CHARDEZ 1983, KAHL 1932, KENT 1882, SCHLICK 1973, STEIN
1859b). STEIN (1859b) supposed that the hydras are only a temporary dwelling for the
keronas, the available food being insufficient there.
The nematocysts of the hydras are not released by Kerona (for example, HiLL 1965,
SCHNEIDER 1985, ZICK 1932), whereas they are released by other ciliates, for example, the
rather small Coleps hirtus. Possibly, the special infraciliature (and the flexible body mar-
gins) of Kerona are adaptations for undisturbed living (and thus protected against preda-
tors) on the hydras. Biomass of 106 individuals about 230 mg (FOISSNER et al. 1991).
One Hydra is usually colonised by 8-18 keronas; according to CAVALLINI (1930) and
COLEMAN (1965), however, up to 100 or 200 individuals per Hydra are possible (Fig.
208j, I); but in this case the growth rates of the keronas are rather low and many of them
leave the Hydra. HILL (1965) stated that Kerona harmed hydras when dominant, and that
an infested Hydra would lose its ciliates if it became very strong, and keronas transferred
to a healthy Hydra had little chance of remaining there. Conversely, SCHLICK (1973)
stressed that heavily colonised hydras also reproduce. According to COLEMAN (1965),
Kerona did not survive on intact, starved hydras.
Kerona pediculus is used as indicator of water quality (FOISSNER 1988a, SLADECEK et
al. 1981, WEGL 1983). The above mentioned hydras and bryozoans live in oligo- to be-
tamesosaprobic standing and running waters (HEITKAMP 1986, WEGL 1983). FOissNER et
al. (1991) thus classified Kerona pediculus as follows: b-o; 0 = 4, b = 5, a = I, I = 2, SI =
1.7.
 Kerona 839

Species transferred to genera not treated in this book

Kerona ciliata GOURRET & ROESER, 1888, Archs BioI., 8: 178, Planche XIV, Fig. 11, 12.
Remarks: According to KAHL (1932, P 482), a supposed synonym of Peritromus faurei
KAHL,1932.

Kerona rotunda FROMENTEL, 1876, Microzoaires, p 271, Planche XIII, fig. 11. Remarks:
Probably a synonym of Aspidisca lynceus.

Kerona vannus MOLLER, 1786, Animalcula Infusoria, p 240, Tab. XXXIII, Fig. 21-23.
Remarks: Now Euplotes vannus.

Species indeterminata

DUMAS (1929, 1930, 1937) and FROMENTEL (1876) described many "Kerona" species. If
not otherwise indicated, it is either a fragment, a deformed hypotrich, or the description
and the illustration are so insufficient that identification is impossible.

Kerona absidata DUMAS, 1930, Microzoaires, p 59 (Fig. 243h).

Kerona anfracta DUMAS, 1929, Microzoaires, p 78 (Fig. 227j, 2431). Remarks: "Rede-
scribed" by DUMAS (1937, P 24).

Kerona aper FROMENTEL, 1876, Microzoaires, p 271, Planche XIII, fig. 15.

Kerona arcuata DUMAS, 1929, Microzoaires, p 79 (Fig. 227p).

Kerona becciformis DUMAS, 1929, Microzoaires, p 77 (Fig. 227f, g).

Kerona bicaudata DUMAS, 1937, Microzoaires, p 21 (Fig. 243k).

Kerona bipes DUMAS, 1937, Microzoaires, p 21 (Fig. 243j).

Kerona calvaeformis DUMAS, 1929, Microzoaires, p 80 (Fig. 243b).

Kerona caudata DUMAS, 1929, Microzoaires, p 76 (Fig. 227b).

Kerona congyrans DUMAS, 1929, Microzoaires, p 80 (Fig. 2270). Remarks: Possibly a

Slylonychia because, in the legend to Planche XXXVII, DUMAS designated it as Kerona
silurus (congyrans).
840 SYSTEMA TIC SECTION

Kerona cuneata DUMAS, 1930, Microzoaires, p 57 (Fig. 243c). Remarks: Possibly a Sty-
lonychia.

Kerona curvata DUMAS, 1929, Microzoaires, p 76 (Fig. 227e). Remarks: Possibly a

Gonostomum. Very likely not identical with the next "species".

Kerona curvata DUMAS, 1930, Microzoaires, p 58 (Fig. 243e). Remarks: Very likely not
identical with the previous species.

Kerona cylindrica DUMAS, 1929, Microzoaires, p 75 (Fig. 243a).

Kerona diademata DUMAS, 1929, Microzoaires, p 77 (Fig. 227c).

Kerona elongata FROMENTEL, 1876, Microzoaires, p 270, Planche XIII, fig. 7.

Kerona epalxiformis DUMAS, 1937, Microzoaires, p 22 (Fig. 243m).

Kerona extuberans DUMAS, 1930, Microzoaires, p 58 (Fig. 243f).

Kerona lacerata FRoMENTEL, 1876, Microzoaires, p 269, Planche XII, fig. 11.

Kerona mamillata DUMAS, 1929, Microzoaires, p 78 (Fig. 227n). Remarks: The illustra-
tion is shown on Planche XXVIII, and not on Planche X as indicated. In 1937, DUMAS de-
scribed a further species with the same name (see next entry).

Kerona mamillata DUMAS, 1937, Microzoaires, p 23 (Fig. 2430, p). Remarks: DUMAS
probably did not realise that he had already described a species having the same name
(see previous entry).

Kerona oblonga DUMAS, 1929, Microzoaires, Legend to Planche XXVIII (Fig. 227h).
Remarks: According to the text (p 78) it is Kerona elongata FROMENTEL.

Kerona ondulata DUMAS, 1930, Microzoaires, p 58 (Fig. 243g).

Kerona ovata DUMAS, 1929, Microzoaires, p 79 (Fig. 227k).

Kerona rastellum MOLLER, 1786, Animalcula Infusoria, p 233, Tab. XXXIII, Fig. 1,2.
Remarks: I did not find this name in the comprehensive index by EHRENBERG (1838). Per-
haps an Aspidisca?

Kerona rostrata DUMAS, 1937, Microzoaires, p 24 (Fig. 243i).

Kerona subrotondata DUMAS, 1930, Microzoaires, p 58 (Fig. 243d). Remarks: Kerona

subrotundata in the legend to Planche XVI is another original spelling. The same name
 Paraurostyla 841

occurred already in DUMAS (1929, legend to Planche XXVIII), which is, according to the
text (DUMAS 1929, P 79), K. rotundata FROMENTEL (Fig. 227i; incorrect subsequent spell-
ing).

Kerona tremula DUMAS, 1929, Microzoaires, p 76 (Fig. 227a).

Kerona triangularis FROMENtEL, 1876, Microzoaires, p 272, Planche XII, fig. 5.

Kerona truncata DUMAS, 1929, Microzoaires, p 77 (Fig. 227d).

Kerona unguicularis DUMAS, 1937, Microzoaires, p 22 (Fig. 243n).

Insufficient redescriptions

Kerona polyporum EHRENBERG - DUMAS, 1929, Microzoaires, p 76, Planche XXVI, fig. 6.

Kerona polyporum EHRENBERG - FROMENTEL, 1876, Microzoaires, p 270, Planche XIII,

fig. 10.

Paraurostyla BORROR, 1972

1972 Paraurostyla BORROR, J. Protozool., 19: 4, 9 - Type (original designation): Urostyla weissei STEIN,
1859.
1986 Paraurostyla BORROR, 1972 - DRAGESCO & DRAGESCO-KERNEIS, Faune tropical, 26: 435 (revision of Af-
rican species).

C h a r act e r i sat ion: Body flexible. Adoral zone of membranelles formed like a
question mark. Undulating membranes in Oxytricha pattern. Numerous frontoventral
cirri arranged in 3-7 longitudinal rows. Usually 6-9 transverse cirri. One left and 1 right
row of marginal cirri. 5-7 dorsal kineties. Caudal cirri present. More than six frontal-
ventral-transverse cirri primordia, and number of cirri each originating from the fourth
to the rightmost primordium usually distinctly >4. Dorsal morphogenesis in Oxytricha
pattern.

Nom e n c I a t u rea n d t a x 0 nom y: Incorrect subsequent spellings: Parauros-

tula weissei (STEIN) and Paraurostula sp. (YANG 1989, P 157); Parurostyla weissei STEIN
(DETCHEVA 1981, P 24; 1986, p 63).
For a detailed description of the patterns mentioned in the characterisation, see chap-
ter 2 in the general section. BORROR (1972a) established Paraurostyla within the Urostyli-
dae to include species previously thought to be closely related to Urostyla grandis, but
with decidedly different appearance of the cirri of the frontal area, as well as entirely dif-
ferent morphogenesis. Somewhat later, he transferred Paraurostyla to the Oxytrichidae
842 SYSTEMATIC SECTION

because the morphogenetic pattern reminds of that of the 18-cirri oxytrichids (BORROR
1979). Furthermore, fragmentation of dorsal kinety 3 is a very strong evidence that
Paraurostyla belongs to the oxytrichids. However, the exact position within this group is
uncertain. WIRNSBERGER et ai. (1986) considered it as sister-group of all other oxytrichids,
but without giving an autapomorphy for Paraurostyla. This position is supported by
some characters of the resting cyst (see there). Another possibility is that Paraurostyla
evolved from flexible 18-cirri oxytrichids (tat is, the Oxytrichinae) by a secondary in-
crease of the number of frontal-ventral-transverse cirri streaks and an increase of the
number of cirri originating from the right streaks. A similar origin is proposed for Ony-
chodromus within the rigid oxytrichids. Conversely, SMALL & LYNN (1985, P 457) and
TUFFRAU & FLEURY (1994, p 141) considered Paraurostyla as member of the amphisiel-
lids. However, this group lacks fragmentation of dorsal kinety 3. EIGNER (1997) classified
it in the Parakahliellidae.
Originally, ten species were transferred to Paraurostyla (BORROR 1972a). However,
some of these deViate distinctly from the type species, and in several species some impor-
tant characters are unknown (for example, fragmentation of dorsal kinety 3) or not sub-
stantiated by silver impregnation (for example, lack of midventral cirri). These species,
briefly discussed in the following paragraphs, are omitted in the present revision.

Paraurostyla dispar (KAHL, 1932) BORROR, 1972 (basionym: Urostyla dispar KAlIL,
1932, Tierwelt DtI., 25: 565, Fig. 98). Remarks: Many frontal cirri, very likely arranged
in a bicorona, indicating that a midventral row is present. Marine.

Paraurostyla gibba (MOLLER, 1786) BORROR, 1972 (basionym: Trichoda gibba MOLLER,
1786, Animalcula Infusoria, p 179, Tab. 16-20). Remarks: According to the first detailed
redescription (STEIN 1859b, P 184) a marine urostylid.

Paraurostyla latissima (DRAGESCO, 1970) BORROR, 1972 (basionym: Urostyla latissima

DRAGESCO, 1970, AnnIs Fac. Sci. Univ. fed. Cameroun (Numero Hors-serie): 100, Fig.
73). Remarks: Very likely a synonym of "Paraurostyia viridis".

Paraurostyla marina (KAHL, 1932) BORROR, 1972 (basionym: Urostyla marina KAlIL,
1932, Tierwelt DtI., 25: 567, Fig. 100). Remarks: Both the original description and the
redescription by BORROR (1979) show that midventral cirri are present, that is, it is an
urostylid. Marine.

Paraurostyla naumanni (LEPSI, 1935) BORROR, 1972 (basionym: Urostyla naumanni

LEPSI, 1935, Bui. Muz. natn. 1st. nat. Chisinau, 6: 16, 1 fig). Remarks: Exact arrangement
of cirri unknown, thus transfer to Paraurostyla unfounded. No redescription available.
About 6 left marginal rows. Marine.

Paraurostyla rubra (ANDRUSSOWA, 1886) BORROR, 1972 (basionym: Urostyla rubra AN-
DRUSSOWA, 1886, Trudy imp. S-peterb. Obshch. Estest., 17: 246, plate II, fig. 10). Re-
 Paraurostyla 843

marks: Superficially described. No redescription available. Obviously no transverse cirri

and numerous macronuclear nodules. Marine.

Paraurostyla viridis (STEIN, 1859) BORROR, 1972 (basionym: Urostyla viridis STEIN,
1859, Organismus der Infusionsthiere I, p 206, Taf. XIII, Fig. 13, 14). Remarks: See
Paraurostyla enigmatica.

The following 5 more recent species were originally classified in Paraurostyla. However,
they differ significantly from the type species and were thus transferred to other taxa.

Paraurostyla buitkampi ForSSNER, 1982, Arch. Protistenk., 126: 40, Abb. 4a-f. Remarks:
BERGER & FOISSNER (1987a, P 197) transferred it to Pseudouroleptus HEMBERGER, 1985
because of the deviating ventral infraciliature. Furthermore, P. buitkampi has only 3 dor-
sal kineties, that is, fragmentation ofkinety 3 and dorsomarginal rows are lacking.

Paraurostyla enigmatica DRAGESCO & DRAGESCO-K.ERNEIS, 1986, Faune tropicale, 26:

437, Planche 125C. Remarks: This conspicuous species from saline marshes in Benin has
about 9 cirral rows left of the median. DRAGESCO & DRAGESCo-K.ERNEIS (1986) thus pro-
posed a new genus, namely Mixotricha (M enigmatica). However, this name is preoccu-
pied by Mixotricha SUTHERLAND, 1933, a termite inhabiting flagellate. JANKOWSKI (1989)
thus introduced the replacement name Ponturostyla. Ponturostyla enigmatica is not
treated in this book because nothing is known about the dorsal morphogenesis (that is, if
fragmentation of kinety 3 is present or absent). Very likely related to Urostyla viridis
STEIN, 1859.

Paraurostyla macrostoma ForssNER, 1982, Arch. Protistenk., 126: 43, Abb. 5a-e and
Paraurostyla terricola BllTKAMP, 1977, Decheniana, 130: 118, Abb. 2. Remarks: Now in
the kahliellid taxon Parakahliella BERGER, ForssNER & ADAM, 1985, which differs from
Paraurostyla by the lack of transverse cirri and in that dorsal kinety 3 does not fragment.
The number of left marginal cirral rows is increased and some parental dorsal kineties are
retained after division.

Paraurostyla pulchra BllTKAMP, 1977, Decheniana, 130: 119, Abb. 3. Remarks: Lacks
caudal cirri and has only 3 dorsal kineties, that is, fragmentation of kinety 3 and dorso-
marginal kineties are very likely absent. Thus, a classification in Paraurostyla and in the
Oxytrichidae is unfounded. JANKOWSKI (1979, p 83) transferred it to Bakuella, which is
also unfounded because midventral cirri are lacking.

Consequently, Paraurostyla comprises four species. Two form the Paraurostyla weissei
complex, namely P. weissei and P. polymicronucleata. ForSSNER (1997b) established
Apoamphisiella for Onychodromopsis tihanyiensis (Fig. 198.1a-g). When I made the
layout of the book, I recognised that Paraurostyla hymenophora (STOKES) BORROR
844 SYSTEMATIC SECTION

(Fig. 21Sa-d) is very similar (possibly synonymous) to that species; thus, it is also trans-
ferred to Apoamphisiella (however, it was too late to change the figure numbers).
Apoamphisiel/a differs from Paraurostyla mainly in that it has only two frontoventral
rows against three or more in Paraurostyla.

Key to species

1 About 9 macronuclear nodules (Fig. 218a) .............. P. polynucieata (p 877)

Usually 2 macronuclear nodules ........................................ 2
2 Two frontoventral rows (for example, Fig. 21Sa-d) ....... Apoamphisie//a (p 781)
3-7 frontoventral rows (for example, Fig. 210g, 211a) ...................... 3
3 Cortical granules yellow-green, spherical, about 1 Jlm across (Fig. 212i; Parauro-
styla weissei complex; p 844) .......................................... 4
Cortical granules colourless; rod-shaped, about 2 x I Jlm (Fig. 216c, d) ......... .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . " P. granulifera (p 874)
4 Usually 3 distinctly enlarged anterior frontal cirri and 3 enlarged posterior frontal
cirri; usually 6 transverse cirri; third frontoventral row begins near distal end of ado-
ral zone of membranelles; rightmost frontoventral row (usually the fourth from left)
not interrupted in the middle region (Fig. 211d, e) ... P. po/ymicronucieata (p 844)
Usually 4 distinctly enlarged anterior frontal cirri and 2 enlarged posterior frontal
cirri; usually 8 transverse cirri; third frontoventral row begins at about level of buccal
vertex; rightmost frontoventral row often interrupted in the middle region (Fig.212r)
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. P. weisse; (p 844)

Paraurostyla weisse; complex (Fig. 8, 9a, b, 21Oa-x, 211a-k, 212a-x,

213a-l, 214, Table 49)

1859 Urosty/a weissei STEIN, Organismus der Infusionsthiere I, p 192 (Fig. 21Oa-c).
1866 Urostyla weissei STEIN - DIESING, Sber. Akad. Wiss. Wien, 53: 120.
1866 Urostyla sp. nov? - WRZESNlOWSKlEGO, Wykaz Szkoty Glownej Warszawskiej, 5: 19 (Latin description,
no illustration).
1867 Urostylajlavicans WRZESNIOWSKIEGO, Rozprawa, p 65 (Fig. 212a, b).
1867 Urostylajlavicans WRZESNlOWSKlEGO, Rocznik ces. kr61. Towarzystwa Naukowego Krakowskiego, 12:
293 (Fig. 212a, b).
1870 Urostylajlavicans WRZESNlOWSKI, Z. wiss. Zool., 20: 480 (Fig. 212c).
1877 Urostyla weissei STEIN, nova varietas - MERESCHKOWSKY, Trudy imp. S.-peterb. Obshch. Estest., 8: 239
(variety not named; no illustration).
1879 Urostyla weissei STEIN, nova varietas - MERESCHKOWSKY, Arch. mikrosk. Anat. EntwMech., 16: 166
(variety not named; no illustration; translation ofMERESCHKOWSKY 1877).
1882 Urosty/a weissii, STEIN - KENr, Manual Infusoria II, p 764 (incorrect subsequent spelling).
1882 Urostylajlavicans, WRZ. - KENr, Manual Infusoria II, p 766.
1894 Urostyla vernalis STOKES, Proc. Am. phil. Soc., 33: 342 (Fig. 21Ik).
1930 Urostylaparagrandis WANG, Contr. bioI. Lab. Sci. Soc. China, 6: 9 (Fig. 212d).
 Paraurostyla 845

1932 Urostyla vernalis STOKES, 1894 - KARL, TieIWelt Otl., 25: 568.
1932 Urostyla weissei STEIN, 1859 - KARL, TieIWeIt Otl., 25: 568 (Fig. 21Og).
1932 Urostylaflavicans WRZESNIOWSKI, 1870 - KARL, TieIWeIt Otl., 25: 568.
1936 Urostyla weissei STEIN - BHATIA, Protozoa: Ciliophora, p 366.
1937 Urostyla polymicronucieata MERRIMAN, Arch. Protistenk., 88: 427 (Fig. 212e).
1939 Urostyla coei TURNER, Trans. Am. microsc. Soc., 58: 395 (Fig. 212g).
1939 Urostyla Iynchi HORVAlH, Arch. Protistenk., 92: 544 (Fig. 212f; new synonym).
1945 Urostyla weissei, STEIN - FAuRE-FREMIET, Bull. bioI. Fr.. Belg., 79: 112 (Fig. 210h).
1961 Urostyla weissei STEIN - BucK, Jh. Ver. vaterl. Naturk. WUrtt., 116: 211 (Fig. 210e).
1962 Urostyla weissei STEIN - LIEBMANN, Handbuch der Frischwasser- und Abwasser-Biologie I, p 360 (Fig.
210d, t).
1963 Urostyla weissei STEIN, 1859 - VUXANOVICI, Studii Cerc. BioI., 15: 203 (Fig. 21Oi).
1965 Urostyla hologama HECKMANN, Arch. Protistenk., 108: 55 (Fig. 212h).
1965 Urostyla weissei STEIN - JERKA-DzlADOsz, Acta Protozool., 3: 345 (Fig. 210k).
1966 Urostyla weissei STEIN - ORAGESCO, Protistologica, 2: 81 (Fig. 2101, n-s).
1968 Urostyla weissei STEIN, 1859 - CHoRIK, Free-living ciliates, p 128 (Fig. 211h).
1969 Urosryla weissei - JERKA-DzlADOSZ & FRANKEL, J. Protozool., 16: 612 (Fig. 211d).
1970 Urostyla weissei STEIN, 1859 - ORAGESCO, Annis Fac. Sci. Univ. fed. CameroUn (Numero Hors«rie):
102 (Fig. 21Ow, x).
1972 Urostyla weissei STEIN - BlcK, Ciliated Protozoa, p 178.
1972 Paraurostyla weissei (STEIN, 1859) - BORROR, J. Protozool., 19: 4, 9.
1974 Urostyla weissei STEIN - STILLER, Fauna Hung., 115: 42.
1974 Parurostyla weissei (STEIN) BORROR - PATSCH, Arb. lnst. landw. Zool. Bienenkd., 1: 55 (Fig. 211g; in-
correct subsequent spelling of Paraurostyla).
1983 Paraurostyla weissei (STEIN, 1859) - SHEN, Protozoa of the Tibetan Plateau, p 203 (Fig. 211 i, j).
1985 Paraurostyla weissei (STEIN, 1859) - WIRNSBERGER, FOlSSNER & ADAM, Zool. Scr., 14: 1 (Fig. 2120-x,
213a-1; detailed redescription of P. weissel).
1986 Paraurostyla weissei (STEIN, 1859) BORROR, 1972 - ORAGESCO & ORAGESCo-KERNEIs, Faune tropicale,
26: 435 (Fig. 210m, 211t).
1988 Paraurostyla weissei (STEIN, 1859) - SHIN & KIM, Korean J. syst. Zool., 2: 70 (Fig. 21Ot-v).
1988 Paraurostyla weissei - KRAMER, Oiplomarbeit, p 19 (Fig. 211e).
1991 Paraurostyla weissei (STEIN, 1859) BORROR, 1972 - FOlSSNER, BLATTERER, BERGER & KOllMANN, lnfor-
mationsberichte des Bayer. Landesamtes fUr WasseIWirtschaft, 1191: 260 (Fig. 212i-n; see also Abb.
16, 17 on p 264, showing 2 large scarming electron micrographs of the ventral and dorsal side).
1994 Paraurostyla weissei (STEIN, 1859) - SHIN, Oissertation, P 53 (Fig. 211a-c).

Nom e n cia t u rea n d t a x 0 nom y: Incorrect subsequent spellings: Parauro-

styla eissei (STEIN) (DETCHEVA 1993, p 34); Paraurostyla weisei (STEIN) (BEER 1958,
P 196; MATIS 1975, P 586); Urostila weissei ST. (AWERINTZEV 1899, P 244); Urostyla
hologamma (JERKA-DZIADOSZ & JANUS 1975, p 311); Urostyla wessei STEIN (SHADIN et al.
1931, p 72); Urostyla weissi STEINII (COPPA 1921, P 184; GULATI 1925, P 753); Urostyla
weissii STEIN (DADAY 1891, pIlI; DADAY 1903, P 324; GULATI 1925, P 752; LEPSI 1929b,
P 269; LEVANDER 1892, P 117; SCHEFFELT 1922, P 116; ZACHARIAS 1888, P 222).
Paraurostyla weissei is very likely a complex of at least two species (WIRNSBERGER et
al. 1984, 1985b). Significant differences occur mainly between European and American
populations and concern especially ventral infraciliature and number of dorsomarginal ki-
neties (see following paragraphs for details). Furthermore, JERKA-DzIADOSZ (pers. comm.
to WIRNSBERGER et al. 1985b) and KRAMER & SCHLEGEL (1989) observed conjugation only
846 SYSTEMATIC SECTION

Fig. 210a-( Paraurostyla weissei complex (It-C, from STEIN 1859b; d, f, from LIEBMANN 1962; e, from BUCK
1961. a-t; from life). a, b, d-I) Ventral views, a, b = up to 300 ~m, d = 170 ~m, e = size not indicated, f =
140 ~m. Specimen in (b) with an ingested Paramecium aurelia. c) Late morphogenetic stage in dorsal view.
The dotted longitudinal rows are the cortical granules; p 844.

among European and among American strains, but not between them. Furthennore, KRA-
MER (1988) found a genetic identity which is typical for morphologically separable spe-
cies. European populations agree rather well with the original description by STEIN
(1859b) and should thus be designated as Paraurostyla weissei (STEIN, 1859) BORROR,
1972. American populations should be detennined as Paraurostyla polymicronucleata
(MERRIMAN, 1937) comb. nov. because this is the first (oldest) description of this deviat-
ing type (Fig. 212e; Urostyla vernalis STOKES, 1894 from America is older, however, it is
more like the European type than on the deviating American type). Urostyla coei TURNER,
1939 (Fig. 212g), Paraurostyla weissei sensu JERKA-DzIADOSZ & FRANKEL, 1969 (Fig.

Fig.210g-m Paraurostyla weissei complex (g, from KAHL 1932; h, from FAURE-FREMIET 1945b; i, from Vux- ~
ANOVICI 1963;j, after BORROR 1979 from DRAGESCO & DRAGESCo-KERNEIS 1986; k, from JERKA-DzIADOSZ 1965;
I, from DRAGESCO 1966b; m, from DRAGESCO & DRAGESCO-KERNEIS 1986. g-i, from life; j, I, m, protargol im-
pregnation; k, Parducz fixation). g-m) Ventral views, g = 280 ~m, h = 220 ~m, i = 200-240 ~m, j = 185 ~m,
k = size not indicated, I = 165 ~m, m = 175 ~m; p 844.
Paraurostyla 847

, .
•••••
,

" ,
-'. " j

. . ....
.... ...."..
.. .."
... .." .. ..
,,
......
....
.
....
...
..
. ..
.
m
848 SYSTEMATIC SECTION

211 d), and P. weissei syngen 2 of KRAMER (1988; Fig. 211 e) are surely conspecific with
P. polymicronucleata.
WIRNSBERGER et al. (1 985b) proposed the name Paraurostyla hologama (HECKMANN,
1965) for populations with cortical granules, 4 anterior frontal cirri, generally 4 fronto-
ventral rows, and 8 transverse cirri arranged like a J. The name Paraurostyla weissei was
proposed for populations probably without cortical granules (for example, "u. weissei
sensu STEIN 1859"), but with 6 frontal cirri arranged in 2 rows of 3 cirri each, 6 or 7
transverse cirri in a straight row, and a number offrontoventral rows varying from 4 to 7.
However, this proposal is somewhat confusing because STEIN (1859b, p 193) unequivo-
cally described and illustrated cortical granules (Fig. 21 Oc).
Synonymy of Urostyla coei, U. jlavicans, U. hologama, U. paragrandis, U. polymi-
cronucleata, and U. vernalis with the Paraurostyla weissei complex is evident and was
already proposed by BORROR (1972a) and WIRNSBERGER et al. (1985b). Urostylalynchi
HORVATH, 1939 is surely a further synonym of Paraurostyla weissei (Fig. 212t). HORVATH
(1939a) assumed that the number of macronuclear nodules and micronuclei is reduced
during cultivation in KNop's solution, that is, in nature he found this species always hav-
ing numerous macronuclear nodules and many micronuclei. Very likely, he confused
Urostyla grandis and Paraurostyla weissei. According to HEMBERGER (1982, P 30) Tri-
chotaxis fossicola (having only 3 frontoventral rows), is a further synonym of Parauro-
styla weissei. I consider it as supposed synonym of Paraurostyla granulifera (see there
for details). Oxytricha multipes CLAPAREDE & LACHMANN, 1858 is considered as supposed
synonym of Paraurostyla weissei to retain stability in nomenclature (see below).
MERESCHKOWSKY'S (1877, 1879) unnamed variety differs from STEIN'S (1859b) de-
scription in that 6 (against 5) cirral rows are present. However, this is within the range of
clonal populations (Table 49). The redescriptions by GULAn (1925; Fig. 218c) and ROSA
(l957a; Fig. 218b) are insufficient.
At superficial live observation species of the Paraurostyla weissei complex can be
easily confused with Allotricha mollis because it is very similar in size, shape, cortical
granulation, the nuclear apparatus, and, most importantly, the 2 or 3 marginal rows of A.
mollis feign frontoventral rows. However, after careful checking of the ventral ciliature
they can be easily distinguished because Allotricha has the typical 18 frontal-ventral-
transverse cirri pattern of, for example, Oxytricha (Fig. 91a-c).
The following chapter is largely based on the detailed redescription of an Austrian
population by WIRNSBERGER et al. (1985b) unless otherwise indicated.

M 0 r p hoI 0 g y and b i 0 log y: In life 150-300 J.1m long, usually 200 to

250 x 50-80 J.1m; up to 220 J.1m long (DRAGESCO 1966b), 180-205 J.1m (DRAGESCO 1970;
fIXed specimens), 120-180 x 35-55 J.1m (HECKMANN 1965), 200-300 x 90-100 J.1m
(JERKA-DzIADOSZ 1965), 225 x 65 J.1m (MERRIMAN 1937), 200-250 J.1m (PATSCH 1974),
150-250 x 50-90 J.1m (SHIN & KIM 1988), up to 300 J.1m (STEIN 1859b), 254-340 J.1m
(STOKES 1894), 200 x 50 J.1m (TURNER 1939), 100-260 J.1m (VUXANOVICI 1963), 158 to
 Paraurostyla 849

···
: : II" .' :

::.,~::
···: '.'.".. , . '. ":
. .."" . .. .......... .
...
.
. I, .. · . . . .. I"

···
ill

........ "
'.
.... .." ~....
:.. "..
t '.. ! ~ ~ ..
~ ~

'... ..... .' .". . ···

" ..... .... . ..:.. ...
"

.: .: .:...: .::
• • II '.
~
\ ~ ~! ....
", II. ..

..

. .... . .
; : .. .. .. ....
......~!" .
\ ~ ~ : .. : ", I. ".' :

... .......'. ..... ":

'"
\.......... ~.

.' .
.
" :
: .. : .. ;
.,..
\",\ .... I' . :
i
.....
n .' p

Fig. 2100-v Paraurostyla weissei complex (o-s, from DRAGESCO 1966b; t-v, from SHIN & KIM 1988. 0-5, v,
protargol impregnation; t, u, from life?). o-s) Variability of ventral infraciliature. t-v) Nuclear apparatus, ven-
tral view (225 J.lm), and ventral infraciliature of Korean population; p 844.
 850 SYSTEMA TIC SECTION

Fig. 210w, x Paraurostyla weissei complex (from DRAGESCO 1970. Protargol impregnation). Ventral infra-
ciliature of specimens from Cameroon, W = 210 ~m, x = 140 ~m. The majority of the popUlation had 2 macro-
nuclear nodules and not 3 as illustrated. I doubt that the arrangement of the cirri is correctly drawn; p 844.
 Paraurostyla 851

.
• ,
, f>

: , ~IJ '
• f>

., .. .,
#
,,', #/j #

, , f '

, ~

,
,
..• •

.. '

a b c
Fig. 211a-c Paraurostyla weissei complex (from SHIN 1994. a, from life; b, c, protargol impregnation). Ven-
tral view and ventral and dorsal infraciliature, a = 245 Ilm, b, c = 240 Ilm. (b, c) agree almost completely with
Figures 212r, s, indicating that SHIN'S illustrations are redrawings; p 844.

200 Ilm (WANG 1930),220 Ilm (WRZESNIOWSKI 1870), poorly nourished specimens only
80 Ilm long (TURNER 1939).
Body slender elliptical, narrowing and sometimes tapering towards posterior end.
Right margin straight to slightly concave, left slightly convex. Flattened about 2: 1 dorso-
ventrally. Body rather flexible, very likely because cortex with only one layer of (longitu-
dinal) microtubules (JERKA-DZIADOSZ 1982). Usually 2 macronuclear nodules with numer-
ous little nucleoli, nodules in life about 20-30 x 10-15 Ilm, both lying left of median;
however, sometimes only 1 nodule or 3 nodules are present (Fig. 210w, x). DNA content
of macronucleus 348 ± 12.3 pg (mean ± SD), GC content of macronucleus DNA about
32 % (STEINBRUCK et al. 1981). Usually 3-6, in life about 6.5 x 5 Ilm sized micronuclei
(WIRNSBERGER et al. 1985b), according to AMMERMANN & MUENZ (1982) 5.6 micronuclei
per cell on average, Paraurostyla polymicronucleata with 3-11 micronuclei (MERRIMAN
1937). Contractile vacuole above mid-body, during diastole with 2 canals, which almost
reach the ends of the cell (Fig. 2120). Cortical granules about 1 Ilm in diameter, yellow-
 852 SYSTEMATIC SECTION

•
,I
.
,, ",,
.
I

,, •,
I

I
, ,
, ,
, , • •
f •
•• • ,
• ", •, , ,,
I

,
f
"I
I

•
,
, ,, •
I

"
I
OJ
, ,
f I
•
• , •
,• , ,"
,
" I
.: "'1 ;
, , ,
J
I
I
I
,• l.,......
,
, ,•
I
I
I

",I
I ,
I,
.' ,,
...... ,.
.. ,.~ ,..;, d
•

Fig. 2l1d, e Paraurostyla weissei complex (d, after Fig. lA from JERKA-DzlADOSZ & FRANKEL 1969; e, from
KRAMER 1988. d, e, protargol impregnation). Ventral infraciliature of American populations (that is, P. polymi-
cronucleata), d = 220 JUIl, e = 192 11m. In (d) the anterior part of the right marginal row is out offocus. Cirri
between arrowheads are caudal cirri. Anterior arrows mark the 3 enlarged "anterior" frontal cirri, posterior
ones mark "posterior" frontal cirri comprising the buccal cirrus, the cirrus behind the right frontal cirrus (ho-
mologous to cirrus IIII2 of the 18-cirri oxytrichids) and a further cirrus right of it Further differences to P.
weissei: third frontoventral row (arrow in e) begins near distal end of adoral zone of membranelies and right-
most frontoventral cirrus shows no interruption, as is often the case in P. weissei (see Fig. 212r, arrow); p 844.

greenish, giving specimens a greenish-brown shimmer at low magnification, grouped

mainly along cirral rows and dorsal kineties, turn pale when the cover glass exerts pres-
sure for a while and water enters the cell (Fig. 210c, 212i); granules are extrusomes of the
mucocyst type. HECKMANN (1965) misinterpreted them as symbiotic algae ("Zoochlorel-
len"). According to WIRNSBERGER et al. (l985b, p 9) several populations (for example,
STEIN 1859b, WRZESNIOWSKI 1870, WANG 1930) probably without cortical granules. How-
ever, this is incorrect because STEIN (1859b, P 193) expressly mentioned them and others
Paraurostyla 853

Fig. 211f-k Paraurostyla weisse; complex (f,

from DRAGESCO & DRAGESco-KERNEIs 1986; g,
from PATSCH 1974; h, from CHORIK 1968; i, j,
from SHEN 1983; k, from STOKES 1894. f, g, pro-
targol impregnation; h-k, from life?). f. g) Ven-
tral infraciliature, f = 138 11m, g = 2 I311m. b-k)
Ventral views, h =310 11m, i,j = 176--200 11m, k
= 254-340 11m; p 844.
854 SYSTEMATIC SECTION

Fig. 212a-d Paraurostyla weissei complex (a, after WRZESNIOWSKlEGO 1867a, b; c, from WRZESNiOWSKI 1870;
d, from WANG 1930. 3-d, from life). Ventral views, a-c =220 J.lm, d =200 J.lm; p 844.

described a yellow-brownish colour of the body. MERRIMAN (1937) wrote in the original
description of Paraurostyia poiymicronuc/eata that "living specimens invariably appear
to be a dark green or brown and are very nearly opaque", strongly indicating that the
American species of the P. weissei complex also has greenish cortical granules. Cyto-
plasm full of numerous intensely yellow shining inclusions of different shape, often ag-
gregated in posterior portion of cell (Fig. 212q). Food vacuoles in life 8-26 ~m in diame-
ter. Movement moderately rapid, nestling close to particles of mud.
Adoral zone of membranelles about 35 % of body length, 45-50 adoral mem-
branelles on average (Table 49), according to JERKA-DZIADOSZ & FRANKEL (1969) 50-80
membranelles. Undulating membranes curved and intersecting, that is, in Oxytricha pat-
tern. Ventral cirral pattern, especially number of ventral rows variable within a popula-
tion. In addition, some significant differences between European and American popUla-
tions exist, that is, between Paraurostyla weissei and P. poiymicronuc/eata. The follow-
ing description is based mainly on WIRNSBERGER et al. (1985b) and thus concerns the
European populations (that is, P. weissei) unless otherwise indicated. Usually with 4 "an-
 Paraurostyla 855

Fig. 212e-h Paraurostyla weissei complex (e, from MERRIMAN 1937; (from HORvArn 1939a; g, from TURNER
1939; h, from HECKMANN 1965. e, g, h, from life; ( aniline blue-staining). e, g) Paraurostyla polymicronu-
c1eata (e; 225 ~m) and its synonym, Urostyla coei (g; 200 lIm). C, h) Two synonyms of P. weissei, namely
Urostyla Iynchi (f; 165 ~m) and Urostyla hologama (h; 120--180 ~m). Arrow in (h) marks the interruption in
the rightmost frontoventral row which is caused by incomplete morphogenesis; p 844.

terior" frontal cirri (that is, the anterior-most cirri of streaks I-IV distinctly enlarged) and
2 "posterior" frontal cirri comprising the buccal cirrus and the cirrus behind the third
frontal cirrus. Behind the fourth (rightmost) "anterior" frontal cirrus is the first frontoven-
tral row (1. VR in Fig. 212r), which is separated into usually 3 slightly enlarged cirri in
the frontal area and a short arched portion beginning near the buccal vertex (Fig. 212r;
originating from same streak! See below). Second frontoventral row extends from frontal
area to the left posterior quarter of the cell. Third row from pharynx to leftmost trans-
verse cirri. Sometimes a very short ventral row follows (4. VR in Fig. 212r). Usually, 4th
frontoventral row (5. VR in Fig. 212r) extends from distal end of adoral zone of mem-
branelles to transverse cirri. Sometimes this rows is interrupted or overlapping in mid-
body, which is the result of incomplete morphogenesis (see below). Transverse cirri in
856 SYSTEMATIC SECTION

Fig.212i-k Parauroslyla weissei (from FOISSNER et al. 1991. i, Normarski differential interference contrast;j,
bright-field; k, scanning electron micrograph). i) Cortical granules, yellow-green, about I Ilm in diameter. j)
Resting cyst with spines, 70-85 Ilm in diameter. k) Ventral view of anterior cell portion. AZM = adoral zone
of membrane lies, BC = buccal cirrus, p = paroral; p844.

life about 20 Jlm long, hook-shaped arranged, rarely reaching posterior end of cell. Left
marginal row J-shaped, approaches the dorsal side more or less. Dorsal cilia in life 3 Jlm
long; number of kineties varying, Austrian population usually with 3 dorsomarginal kine-
ties (Table 49, Fig. 2I2s). Caudal cirri in life about 11 Jlm long, often hardly distinguish-
able from cirri of left marginal row. Caudal cirri according to Oxytricha pattern (Fig.
24a), that is, on posterior end of dorsal kineties 1,2, and 4; usually 3-5 cirri per kinety.
American populations (that is, Paraurostyla polymicronucleata) usually with only 3
enlarged "anterior" frontal cirri and 3 slightly enlarged "posterior" frontal cirri (Fig.
211 d, e). Usually 4 frontoventral rows; third row begins near distal end of adoral zone of
membrane lies (Fig. 21Ie, arrow). Rightmost frontoventral row not composed of cirri of2
streaks (WIRNSBERGER et al. I985b). Transverse cirri not hook-shaped arranged. Usually 6
dorsal kineties, that is, only 2 dorsomarginal kineties.
JERKA-DzIADOSZ & BANACZYK (1983) described a naturally occurring variant with
more than one left marginal row and having several phenotypic defects including modi-
fied shape, increased frequency of cortical reorganisations, slower growth rate (>24 h
against 12-18 h; 20 QC), and formation of monsters.
 Paraurostyla 857

;
J

1·
CI:
..
I
'. " ':1
0
~ 0
a:

Fig. 2121-0 Paraurostyla weissei (from FOlSSNERet aI. 1991. ~ m, scanoingelectron micrographs; n, protargol
impregnation). 1-0) Ventral infraciliature. Large arrow in (n) marks the discontinuity in the rightmost fronto-
ventral row due to incomplete fusion of the cirri of the two rightmost cirral streaks. The small arrow marks fi-
brils originating from a cirrus. CC = caudal cirri, LMR = left marginal row, Ma = anterior macronuclear nod-
ule, p = paroral, RMR = right marginal row, TC = transverse cirri; p 844.
 00
Vl
0
00
'&\8 8
°° a 008°0~
00 0 oo~o,,0o a AFC
0 '-;-';:--7
8J'~o 'b
0 00
0 g
0 0
o ° 'if' 0

08
°aU OJ '~\-6 o
0 o
00 o
Jf' o{oo
0 0 •k
°o&' 0 og
00 o
00
°00
000
,-J-5 u
00 o~o •
00 0
I. )t. C/)
II~MR-
P ;'-~2.VR /"0 :, :' -<
C/)
OP 0 ' ',J
•
•.
-- • 0,
• @
" Qt
.• . •. - ~
• . • ~
~?O 4)JR-I-'--"~"
: #.
•• ...-. , n
C/)
. • m
o (J
5,VR-\~~ ~ ~ . ~
1 \ ~. • o
• •• # ..
• 00
--•.-
~

•• •••... - o·
.
0,
~
0 0,
r C-\-'.-.•• .-/ " 0 ,
• • "
~ ~q .. -..." r s ~ ...~{ CC ~oo~t
Fig. 212o--t Paraurostyla weissei (from WIRNSBERGER et al. 1985b. o--q, from life; r-t, protargol impregnation). 0) Ventral view, 200 11m. p, q) Cortical granules and cy-
toplasmic, yellow shining inclusions (crystals), bar = 50 11m. r, s) Ventral and dorsal infraciliature, 155 11m. Arrow in (r) marks discontinuity in rightmost frontoventral
row. t) Ventral infraciliature ofa very early morphogenetic stage, 180 11m. Explanation of original labels: AFC = anterior frontal cirri (comprising the anterior-most cirri
of streaks I-IV), AMZ = adoral zone of membranelles, CC = caudal cirri, CV = contractile vacuole, FV = food vacuole, IPM = endoral, LMR = left marginal row, OP =
oral primordium, OPM = paroral, PF = pharyngeal fibres, PFC = posterior (enlarged) frontal cirri, RMR = right marginal row, TC = transverse cirri, l.VR-5.VR = fron-
toventral rows 1-5, 1-7 = dorsal kineties; p 844.
 Paraurostyla 859

Resting cyst spherical, in life about 70-85 Ilm across, with many spine-shaped,
~ Ilm long protuberances (Fig. 212j). Cyst wall in life about 4 Ilm thick, consists of 4
morphologically distinct layers. Ultrastructure and cytochemistry similar to that of other
kinetosome-resorbing cysts. However, the cytoplasm shows characters of''urostylid-type''
cysts, indicating that P. weissei is a "transition ciliate" between the oxytrichids and uro-
stylids (DELGADO et al. 1987). Specimens starving at 20°C are unable to encyst (HECK-
MANN 1965).

M 0 r ph 0 g e n e sis: This part of life cycle is described by JERKA-DzIADOSZ (1965),

JERKA-DZIADOSZ & FRANKEL (1969), and WIRNSBERGER et al. (1985b). Morphogenesis of
ventral ciliature is rather complicated, especially during early stages of streak formation.
The general course is shown in Figures 212t-x, 213a-l. The following paragraphs are
largely from WIRNSBERGER et al. (l985b), describing only important stages and differing
observations to the paper by JERKA-DzIADOSZ & FRANKEL (1969).
Stomatogenesis: In the earliest stage of stomatogenesis small groups of basal bodies
appear vel)' close to the posterior-most cirri of the first frontoventral row (Fig. 212t).
Somewhat later this anarchic field migrates slightly to the left (Fig. 212u). In the Austrian
population the primordium of the undulating membranes of the opisthe is not formed "de
novo", as JERKA-DzIADOSZ & FRANKEL (1969) stated, but derives from disaggregation of
1-2 postoral cirri of the first frontoventral row.
Development of cirral primordia: The first 3 frontal-ventral-transverse streaks (with-
out the primordium of the undulating membranes) in the proter originate either from the
posterior 3 frontal cirri (JERKA-DzIADOSZ & FRANKEL 1969) or from the posterior 2 frontal
cirri and the anterior-most cirri ofthe first frontoventral row (WIRNSBERGER et al. 1985b)
proving that the right "posterior" frontal cirrus of the American populations (P. polymi-
cronucleata) is homologous to the anterior part of the first frontoventra1 row of the Euro-
pean populations. Furthermore, this pattern agrees largely with the pattern of the 18-cirri
oxytrichids (Table 4). The equivalent 3 frontal-ventral-transverse streaks in the opisthe
separate from the posterior undulating membrane field, which develops from dispersion
of basal bodies of 1-2 postoral cirri of the first frontoventral row (Fig. 212v, w, 213a).
Later, these streaks later elongate (Fig. 213b, c). JERKA-DzIADOSZ & FRANKEL (1969) sup-
posed that these streaks were apparently formed de novo, although involvement of some

Fig.212u-x Paraurostyla weissei (from WIRNSBERGER et al. 1985b. u-x, morphogenetic stages after protargol ~
impregnation). u-w) Ventral infraciliature of very early and early stages, u = 170 /lm, v = 230 /lm, w =
200 /lm. x) Dorsal infraciliature of an early stage, 205 /lm. Explanation of original labels: DP = dorsal primor-
dia within kineties 1-3, FVTP = frontal-ventral-transverse cirri primordia, UMP = primordium of the undulat-
ing membranes; p 844. Figures on p 860.

Fig. 213a-c Paraurostyla weissei (from WIRNSBERGER et aI. 1985b. a-c, ventral infraciliature of morphoge- ~
netic stages after protargol impregnation). Middle stages showing formation of frontal-ventral-transverse cirri
primordia and marginal primordia formation, a = 205 /lm, b = 160 lim, c = 200/lm. 1-9 = frontal-ventral-
transverse cirri primordia !-IX; p 844. Figures on p 861.
860 SYSTEMATIC SECTION

x
" . "

, .. , , ' .
\ .....
", ,
.,
.... ', " .... ', . " .. "
"#\'''''' t ....... of. 0".:'
I
c..
C

o b
o o
o
'" 0., o
o

>
•
.
~

:::I
,...
C\I
C\I
 Paraurostyla 861

. . ';,~?~~~~.~.:. :.:. . .
.
Q

.
"
.' ."
'

Q ~v
"~"'~·"-"". n",,,,,,,.~,,.,..~,,~.,~ 1) <) <) <) ~ <) <) <) ~ <) ~ ,!.;:.",
..

<> co
<>
G

~::··:~'~:~:;"i."o. 0 ~
o .:,:. . {::.:,: .•Li 0 0.

co
,..
C")

N
 00
0\
N

l r.: 8 0 0 "

,~,;::\. co c • #
..
~.. .... 0 •
0
::~:•• 0 0 : ••
/._ 0 l) 0· • •
;>'.::;>"'
, l
~

~ ---
~
"'" C/)

•
t \~jtt/
" . co a - .. .""",,.
,\ •
,
~
• •,,;::
.l~
q
IJ
-<
C/)

;l
o· f~ Q

"'"I
\% . # ·l~ ~
~
DP-\-t:~ .:. g ~
j \ ,••
:~. ~ I
\.1 :=lo ~ n
~. '.~ \ C/)
\' ~, f tTl
\, (1
i dllPI , ::l
• . : ::::.
• ! ;0.
,
4 ••• 0
o 0
~
I: l
\ ~
;;
4 0 ~
i: J
• Q o 0
a
o 0
• Q Q
. """ . a
• 0 a
" . • a Q
.Q~ 0
.... "
.::.oY" f 9
Fig. 213d~ Paraurostyla weissei (from WIRNSBERGER et aI. 1985b. d-g, morphogenetic stages after protargo! impregnation). d, 1) Ventral infraciliature of middle
stages showing segregation of new cirri and formation of dorsomarginal kineties, d = 180 ~m, f = 160 ~m. Parental structures white, new ones black. e, g) Dorsal infra-
ciliature and nuclear apparatus of middle stages, e = 180 ~m, g = 208 ~m. Arrowheads mark fragmentation of dorsal kinety 3. DP = new dorsomarginal kineties ofopis-
the; p 844.
 Paraurostyla 863

parental cirri cannot be completely ruled out. The remaining streaks develop from por-
tions of the parental cirral rows (Fig. 213a, b). Typically, one streak originates within
each row, except the 2 overlapping streaks associated with the farthest right ventral row
in the anterior region (JERKA-DzIADOSZ & FRANKEL 1969). In the Austrian population the
rule is that 2 streaks are built at a time in the 2 right ventral rows, in both proter and opis-
the (Fig. 213c, d).
In the Austrian population usually 8, seldom 9, frontal-ventral-transverse cirri streaks
are formed (WIRNSBERGER et al. 1985b), whereas JERKA-DzIADOSZ & FRANKEL (1969) re-
port a greater variation of 5-9, mostly 6 or 7.
Differentiation of frontal-ventral-transverse cirri: European populations (P. weisse!)
usually have a higher number of transverse cirri (usually 8) than American populations
(P. polymicronucleata; usually 6; Table 49). According to WIRNSBERGER et al. (1985b)
the streaks segregate the following cirri: streak I (= streak 1 in Fig. 213c), left "anterior"
frontal cirrus (homologous to cirrus III of 18-cirri oxytrichids), undulating membranes;
streak II, "anterior" frontal cirrus 2 (homologous to cirrus II13 of 18-cirri oxytrichids),
left "posterior" frontal cirrus (homologous to buccal cirrus [II12] of 18-cirri oxytrichids),
leftmost transverse cirrus; streak III, "anterior" frontal cirrus 3 (homologous to cirrus
IIII3) of 18-cirri oxytrichids), "right posterior" frontal cirrus (homologous to cirrus IIII2
of 18-cirri oxytrichids), next transverse cirrus; streak IV, rightmost "anterior" frontal cir-
rus (often not vel)' prominent), anterior (right of undulating membranes) and posterior
(behind buccal vertex) part offrrst frontoventral row (1. VR in Fig. 212r), next transverse
cirrus; streak V, second frontoventral row (2. VR in Fig. 212r), next transverse cirrus;
streak VI, third frontoventral row (3. VR in Fig. 212r), next transverse cirrus; streak VII,
posterior half offourth frontoventral row (5. VR in Fig. 212r because this Figure shows a
specimen with a "surplus" row [4. VR]), next transverse cirrus; streak VIII, anterior half
offourth frontoventral row, rightmost 2 transverse cirri (according to my terminology the
anterior of these 2 transverse cirri is in fact a pre transverse ventral cirrus which is vel)'
likely homologous to cirrus VI/2 of the 18-cirri oxytrichids; furthermore, the conspicuous
anteriad migration of the main portion of this streak implies that it is homologous to the
migratol)' cirri [VII3 and VI/4] of the 18-cirri oxytrichids).
The American populations differ from this pattern in the following details (JERKA-
DZIADOSZ & FRANKEL 1969): the anterior part of the first frontoventral row consists of
only 1 cirrus (instead of usually 3) and is thus considered as third (rightmost) posterior
frontal cirrus (posterior-most arrow in Fig. 21Id); the third frontoventral row begins more
anteriorly, namely near the distal end of the adoral zone of membranelies (Fig. 21le, ar-
row) and not near the buccal vertex; the fourth (= rightmost) frontoventral row is the
product of only 1 streak (vel)' likely streak VII), that is, streak VIII, which produces 2
transverse cirri in the European strains, is lacking, resulting in usually 6 instead of 8
transverse cirri. The last difference especially (lack of streak VIII) is vel)' conspicuous
and should be checked again. The micrographs in JERKA-DzIADOSZ & FRANKEL (1969) do
not allow an unequivocal interpretation because the stages are too early (Fig. 9) or too
late (Fig. 10).
 864 SYSTEMATIC SECTION

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Fig. 213h-j Paraurostyla weissei (from WIRNSBERGER et a11985b. h-J, morphogenetic stages after protargol
impregnation). h,j) Ventral infraciliature ofa late and a very late stage, h = 160 Ilm,j = 187 Jim. Arrows mark
the anterior portion of the rightmost streak which migrates anteriorly to fuse with streak on its left. In the opis-
the of(j) this process is obviously already completed. Very likely, the anterior portion of the rightmost streak is
homologous to the migratory cirri (VU3, VU4) of the 18-cirri oxytrichids. Parental structures white, new black.
i) Dorsal infraciliature and nuclear apparatus ofa late stage, 192 11m; p 844.
 Paraurostyla 865

,,
:

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..
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Fig. 2131<, I Paraurostyla we issei (from WIRNSBERGER et aI. 1985b. k, I, protargo\ impregnation). Ventral and
dorsal infraciliature and nuclear apparatus of an opisthe, 100 ~m. Parental structures white, new black; p 844.

Dorsal morphogenesis proceeds in Oxytricha pattern (JERKA-DZlADOSZ 1982), that is,

kinety 3 shows simple fragmentation and 2 (American strains) or 3 (Austrian strains) dor-
somarginal kineties occur. Division takes about 180 min (JERKA-DZlADOSZ 1965).
Conjugation is total in Paraurostyla weissei, that is, the left conjugant, the donor be-
comes absorbed into its mate, the recipient (Fig. 214; FRONTCZAK-BANIEWICZ & JERKA-
DZIADOSZ 1992, HEcKMANN 1965, ILoWAISKY 1916, MOLDENHAUER 1965). It undergoes 3
rounds of cortical reorganisation. Remarkably, the first reorganisation is abortive: on the
ventral surface only a minute adoral zone of membranelles primordium arises, which
does not, however, give rise to membranelles; somatic cirral primordia are also absent.
That the fITSt reorganisation does occur is testified by the differentiation of bristle primor-
dia on the dorsal surface. As in Pseudourostyla cristata, after the first cortical reorganisa-
tion the cell reabsorbs almost all ciliature, and rounds off to become a "zygocysf' before
initiating the second reorganisation. The ciliature generated during the second reorganisa-
tion is incomplete, although the cell can feed. Normality is restored by the third reorgani-
sation, which resembles a physiological reorganisation (NG 1990, P 40). Changes of cor-
tical structures during conjugation and formation of single exconjugant were studied by
HEUMANN (1975) and JERKA-DzIADOSZ & JANUS (1975).
 866 SYSTEMATIC SECTION

o
o o
o
o
o o
o
o

o
o
o
o

Fig.214a Total conjugation of Paraurostyla weissei (from JERKA-DZIADOSZ & JANUS 1975; continued in Fig.
214b). The open circles represent the frontal cirri, the striped fields the adoral zone of membranelles, the solid
lines represent cirraI rows. Primordial structures indicated as fields of fine dots. On left side of each drawing
main features of nuclear apparatus. In 2-5 the drawing of the micronucleus applies to both mates, in 6-9 only
to the right conjugant. On the right side of some drawings the time (in hours) since mating is indicated. 2) For-
mation of one common adoral zone of membranelles. 3) Beginning of resorption of the adoral zone of mem-
branelles, micronuclei are swollen. 4) Resorption of ventral infraciliature in left mate. Micronuclei in early pro-
phase. 5) Resorption of ventral and transverse cirri in the right mate. Tapering of both posterior ends. Micronu-
cleus in parachute stage. 6) Proliferation of basal bodies on the ventral surface. Migration of pronuclei. 7) Pro-
liferation of basal bodies of the dorsal primordia (small sketch at right side). Division of synkaryon; p 844.
 Paraurostyla 867

72h

()
o o
o
o o
o o

o o 10
00
o
11

o 0
o 0

0 0
0
0
0
o
0
0

12 13
14
Fig. 214b Schematic drawing of total conjugation of Paraurostyla weissei (from JERKA-DzIAOOSZ & JANUS
1975; continued from Fig. 214a; see there for general remarks). 8) Resorption of ventral primordia and old
ventral ciliature. Second division of synkaryon. 9) Formation of zygocyst. Resorption of ventral ciliature. Dif-
ferentiation of new nuclei. 10) The zygocyst. The macronuclear anlagen and 2 micronuclei. 11) The primordia
of ventral ciliature in the zygocyst. The anlagen of macronucleus and micronucleus in prophase. 12) Small ex-
conjugant after the cortical reorganisation. Note the incomplete cortical pattern. Single macronucleus and 4
micronuclei. 13) The third cortical reorganisation. Formation of normal pattern. Micronucleus in telophase,
macronucleus elongated. 14) The exconjugant after third cortical reorganisation showing normal pattern;
p 844.
868 SYSTEMATIC SECTION

Table 49 Morphometric data of the·two species of the Paraurostyla },I'eissei complex (1, P. weissei from
WIRNSBERGER et aI. 1985b; 2, three P. weissei populations from Germany-and one from Poland from KRAMER
1988; 3, two Polish P. weissei populations from KRAMER 1988; 4, three North American P.po/ymicronuc/eala
populations from KRAMER 1988). All data are based on protargol-impregnated specimens. All measurements in
micrometres. CV = coefficient of variation (in %), Max = maximum value, mean = arithmetic mean, Min =
minimum value, n = sample size, SD = standard deviation

Character Species mean SD CV Min Max n

Body, length 1 126.8 34.3 27.1 73 198 28
2 162.5 130 212 135
3 160.7 122 195 51
4 172.6 141 206 73
Body, width 47.8 13.4 28.0 26 73 28
2 50.3 22 72 133
3 48.8 32 70 51
4 50.8 35 76 72
Adoral membranelles, number 1 45.4 9.3 20.5 28 63 28
2 48.2 35 62 59
3 44.3 39 50 39
4 52.5 49 58 28
Adoral zone of membranelles, length 47.1 10.2 21.7 30 66 28
Macronuclear nodules, number 2.0 0.2 9.3 2 3 28
2 2.0 1 3 160
3 1.8 3 51
4 2.1 2 4 123
Macronuclear nodule, length 17.2 2.3 13.6 13 22 28
Macronuclear nodule, width 8.5 1.1 12.9 6 11 28
Micronuclei, number 4.0 0.9 21.8 3 6 28
2 5.0 3 8 50
3 4.2 0 11 25
4 5.7 2 12 50
Micronucleus, length 5.0 0 0 5 5 28
Micronucleus, width 4.0 0 0 4 4 28
Left marginal row, number of cirri 1 34.8 7.2 20.6 20 59 28
2 39.4 29 49 S9
3 39.4 28 47 37
4 43.5 33 52 28
Right marginal row, number of cirri 1 35.0 6.1 17.4 24 46 28
2 37.2 27 47 58
3 37.3 29 44 36
4 43.5 34 52 28
Frontoventral rows between 3.9 0.54 13.7 3 5 28
marginal rows, number 2 4.5 3 7 135
3 4.1 3 5 51
4 4.3 4 7 73
Frontoventral row 1, number of cirri 9.0 2.1 23.8 5 12 28
2 9.2 2 14 59
3 7.5 2 13 39
4 7.4 4 11 27
Frontoventral row 2, number of cirri 1 15.3 2.4 15.9 10 19 28
2 14.4 8 19 60
 Paraurostyla 869

Table 49 Continued

Character Species mean SD CV Min Max n

Frontoventral row 2, number of cirri 3 13.7 5 26 39
4 14.7 8 18 27
Frontoventral row 3, number of cirri 1 14.4 4.4 30.1 6 27 28
2 14.9 3 23 60
3 13.7 5 26 39
4 25.4 16 30 27
Frontoventral row 4, number of cirri 25.2 7.2 28.7 8 35 23
2 21.2 2 34 58
3 23.4 3 33 35
4 32.5 22 39 27
Frontoventral row 5, number of cirri 24.3 7.5 30.8 20 33 3
2 25.6 7 34 26
3 27.3 25 30 6
4 30.7 27 34 3
Frontoventral row 6, number of cirri 2 19.3 8 35 3
Frontoventral row 7, number of cirri 2 21.0 I
Ventral cirri, total number 61.9 14.3 23.1 34 86 28
Anterior frontal cirri, number 3.9 0.3 6.7 3 4 28
2 3.9 3 5 135
3 3.1 3 4 50
4 3.0 3 3 73
Posterior frontal cirri, number I 2.0 0 0 2 2 28
2 2.0 2 2 135
3 1.8 I 2 50
4 3.0 3 3 73
Transverse cirri, number 1 7.8 0.7 9.0 6 9 28
2 8.4 7 10 135
3 7.3 5 9 50
4 6.4 6 8 71
Dorsal kineties, number 1 7.2 0.9 12.0 6 8 28
2 6.5 6 9 62
3 6.1 6 7 40
4 5.3 5 6 22
Caudal cirri, number 13.5 1.8 13.5 10 18 23
2 11.4 9 16 69
3 10.7 7 13 44
4 13.4 11 16 32

Species of the Paraurostyla weissei complex are the object of numerous studies con-
cerning mainly morphogenetic processes, ultrastructure, genetics, and biochemistry:
BAKOWSKA (1980, 1981), BAKOWSKA & JERKA-DZIADOSZ (1977, 1978, 1980), BALBIANI
(1862), Bllli et al. (1994), BRUGEROLLE & AooUTIE (1988), DELGADO & FLEURY (1993),
DUBIELECKA & JERKA-DZIADOSZ (1989), ENGELMANN (1875), FAUllli-FREMIET (1945a,
1948), FLEURY & GUYADER (1990), FLEURY & LAURENT (1993, 1994), FLEURY et al.
870 SYSTEMATIC SECTION

(1993), FRANKEL (1973, 1974, 1975), JERKA-DzIADOSZ (1967, 1972, 1974, 1976, 1977,
1980, 1981a-c, 1983, 1985, 1987, 1989, 1990), JERKA-DZIADOSZ & BEISSON (1990),
JERKA-DzIADOSZ & CZUPRYN (1997), JERKA-DzIADOSZ & DUBIELECKA (1985), JERKA-
DZIADOSZ & FRANKEL (1970), JERKA-DzIADOSZ & GOLINSKA (1977), JERKA-DZIADOSZ &
WIERNICKA (1992), JERKA-DzIADOSZ et al. (1989a, b), KRAUT et al. (1986), LAURENT &
FLEURY (1994), LAURENT et al. (1992), PUYTORAC et al. (1994), RAIKov (1989), SAPRA et
al. (1985), SCHLEGEL & STEINBROCK (1986), STEINBROCK (1990), TUFFRAU (1965a).

o c cur r e n c e and e colo g y: Very common all over the year in moderately to
heavily polluted freshwater habitats; benthic. Locus classicus is Niemegk, a small town
near Berlin, Germany, where STEIN (1859b) discovered Paraurostyla weissei in slowly
running waters covered with Lemna sp. He also found it in Tharandt, a village near Dres-
den, Germany, and Prague, Czechoslovakia. According to STEIN (1859b) it was very
abundant in shaded ditches and ponds, especially when they were almost desiccated, that
is, when Lemna almost rested upon the decaying leaf litter. It also occurred very abun-
dantly - together with Stylonychia mytilus and Urostyla grandis - in collecting jars when
Lemna began to putrefY. Locus classicus of the synonym Urostyla flavicans is Warsaw,
Poland, where WRZESNIOWSKIEGO (1867a, b; see also WRZESNIOWSKI 1870) discovered it in
a small pond among Lemna minor. STOKES (1894) discovered the synonym Urostyla ver-
nalis near Trenton, New Jersey, USA, in shallow wayside pools in the early spring. Locus
classicus of the synonym Urostyla paragrandis is Pe Ji Koh, Nanking, China, where
WANG (1930) discovered it in great abundance in a shallow Sphagnum pond in May and
June. Locus classicus of Paraurostyla polymicronuc/eata, the second species of the com-
plex, is the Edgewood Park near the outskirts of New Haven, Connecticut, USA, where
MERRIMAN (1937) discovered it in a small pool close to the edge of the West River in
autumn. New Haven is also the locus classicus of the synonym Urostyla coei, where
TURNER (1939) discovered it in the Mill River. Locus classicus of the synonym Urostyla
Iynchi is Lengyelkapolna near Szeged, Hungary, where HORVATH (1939a) discovered it in
moss; unfortunately, he did not mention if the moss was from a freshwater or from a ter-
restrial habitat. Locus classicus of the synonym Urostyla hologama is probably a lake
(Federsee) at the village of Bad Buchau, Germany (HECKMANN 1965).
Records of the Paraurostyla weissei complex substantiated by illustrations: on sedi-
ment on the shore of an oligotrophic lake (Mondsee) in Austria (WIRNSBERGER et al.
1985b); pond in Thonon-les-Bains, France (DRAGESCO 1966b); mesosaprobic running wa-
ters in Germany (FOISSNER et al. 1991, 1992a, b); common, but not very abundant in vari-
ous running waters of Nordwilrttemberg, Germany (BUCK 1959, 1961); ponds and small
running waters near Bonn, Germany (PATSCH 1974); bog in Germany (MOLDENHAUER
1965); Moldova (CHORIK 1968); pond and rivers in Poland (JERKA-DzIADOSZ 1965); Bu-
charest, Romania (VUXANOVICI 1963); Swan Lake, Johnson County, Iowa, USA (JERKA-
DZIADOSZ & FRANKEL 1969); Tibetan plateau (SHEN 1983).
Records not substantiated by illustrations: among algae and mosses in springs near
Vienna, Austria (KOHN 1940); among Sphaerotilus and detritus in heavily polluted run-
 Paraurostyla 871

ning water in Austria (STRUHAL 1969); Tyrol, Austria (DALLA TORRE 1891); with
0.7-9.7 % frequency in Bulgarian rivers (DETCHEVA 1979a--<:, 1981, 1986, 1991); sapro-
bic brook and Moldavia River near Prague, Czechoslovakia (BuCHAR 1957, KALMus
1928, SRAMEK-HuSEK 1953); Cymbella aufwuchs on stalks of Phragmites in a lake (Ober-
see) near Reval and in ponds in Estonia (JACOBSON 1928, SCHNEIDER 1909); with very
high abundance (10 800 indo ml- I) in the Kahmhaut of a 17 d old waste water, Leipzig,
Germany (LIEBMANN 1936); alphamesosaprobic running waters in Germany (BEER 1958,
CASPERS & SCHULZ 1962, KLAPPER 1963, REICHENBACH-KLINKE 1959, ZIEMANN 1985); ben-
thaI and aufwuchs of Hamburg Harbour and Elbe River upstream from Hamburg, Ger-
many (BARTSCH & HARTWIG 1984, GRIMM 1968, TENT 1981); cooling plant of a power sta-
tion in Germany (BERNERTH 1982); mesotrophic lake in Germany (MOCKE 1979); bogs in
Germany (SCHEFFELT 1922); rarely in mine-water and in caves in Germany (GITTLESON &
HOOVER 1969, GRIEPENBURG 1934, 1935, WETZEL 1929; further records from Germany:
LEVANDER 1892, WETZEL 1928a, b); Hungary (DADAY 1891); Latvian rivers (LIEPA 1978);
the Netherlands (VERSCHAFFELT 1930, WIERSMA-VERSCHAFFELT 1936); Henares River,
Spain (SOLA et al. 1996); Sweden (QUENNERSTEDT 1865); littoral of Lake Geneva, Swit-
zerland (ANDRE 1912, 1916; further records from Switzerland: MERMon 1914, RIGGEN-
BACH 1922); reservoir in Azerbaijan (ALEKPEROV 1984b, c); cooling plant in Moldova
(CHORIK & VIKOL 1973); moss-puddle near Helsinki, Finland (LEVANDER 1894a); Danube
River in Romania (ENACEANU & BREZEANU 1970); Palestine (BODENHEIMER 1937, LEPsr
1929a); Tibet (WANG 1977); Lake Onega, USSR (MERESCHKOWSKY 1877, 1879); rivers
(for example, Oka, Volga) in the USSR (MAMAEVA 1979a, b, NEISWESTNOWA-SHADINA
1935, SASSUCHIN 1924, 1930, SASSUCHIN et al. 1927, SHADIN 1940, SHADIN et al. 1931);
amongst putrefying algae and other sites in Lake Baikal, USSR (GAJEVSKAJA 1927, GA-
JEWSKAJA 1933, ROSSOLIMO 1923); Turkestan, USSR (DADAY 1903, 1904; further record
from the USSR: MlNKEWITSCH 1898); Yuelushan Area, China (YANG 1989); pond water in
Punjab, Lahore, India (BHATIA 1936); paddy fields in Japan (TAKAHASHI & SUHAMA 1991);
Argentina (SECKT 1924); Brazil (CuNHA 1913, PROWAZEK 1910); pond in Cameroon
(NJINE 1977); paddy field near Tamatave, Madagascar (SONDHEIM 1929).
MuwwwrrsCH (1989) found Paraurostyla weissei predominantly at low salinity
(0-200 mg 1-1 Cn, KLAPPER (1963) at up to 6.6 g 1-1 CI-. BOUTCHINSKY (1895), BUTSCHIN-
SKY (1897), WILBERT (1995), and ZACHARIAS (1888) recorded it from saline lakes in the
USSR, Canada, and Germany.
Also in trickling filters (CURDS 1975, LIEBMANN 1962) and activated sludge (KLI-
MOWICZ 1973, LIEBMANN 1962); however, Paraurostyla weisse; avoids water with liquid
manure (LIEBMANN 1962). KLIMOWICZ (1972) counted 383000 ind.l- I in an activated
sludge plant under the following conditions: mean loading of aeration tank
0.l44 kg BODs m-3 d- I; mean loading of activated sludge 0.0385 kg BODs kg-I d- I; mean
concentration of activated sludge 3.9 kg m-3; time of sewage aeration about 20 h d- I; de-
struction of organic matter 90.4 %; mean oxygen content in aeration tank 3.6 mg 1-1; mean
temperature of activated sludge 4-9 °C; mean pH of effluent and affluent 7.8 and 7.3 re-
spectively.
872 SYSTEMATIC SECTION

Records from terrestrial habitats (COPPA 1921, GREEFF 1889, ROSA 1957a, Fig. 218b;
VARGA 1961) unreliable, although resting cysts are formed (Fig. 212j).
Feeds on bacteria, algae (green algae, diatoms, desmidiaceans), flagellates (Chilomo-
nas), testaceans (Arcella), ciliates (Tetrahymena pyriformis, Colpidium), and detritus
(BICK 1972a, BICK & KUNZE 1971, FAURE-FREMIET 1961a, JERKA-DzIADOSZ 1965, KAm..
1932, KALMus 1928, MERruMAN 1937, MOLDENHAUER 1965, W~G 1930, WIRNSBERGER et
al. 1985b). Also ingests starch, which is digested within 3 days (STRUHAL 1969). TURNER
(1939) observed a specimen ingesting a small nematode. The worm survived in an active
condition within the ciliate until the latter died 3 days later without dividing. Also canni-
balistic (MOLDENHAUER 1965).
WIRNSBERGER et al. (1985b) used tap water as culture medium, and yeast and a spe-
cies of the Tetrahymena pyriformis complex, grown on city egg yolk, was added as food.
JERKA-DZIADOSZ & FRANKEL (1969) maintained long-term stocks of P. polymicronuc/eata
as steady-state cultures containing Pringsheim solution and a food-chain made up of a
few wheat and rice grains, bacteria, Tetrahymena, and Paraurostyla. Under these condi-
tions the Tetrahymena popUlation is kept at low density, and overfeeding of the Parauro-
styla is avoided - overfed P. weissei cannot reproduce and eventually die. Length of cell
cycle 24 h at about 23°C in Pringsheim medium and fed daily with Chlorogonium elon-

Fig. 215a-d ApoamphisieZIa hymenophora (a, from STOKES 1886; b, from LUNDIN & WEST 1963; c, d, from
GRIMES & L'HERNAULT 1978. a, b, from life; c, d, after protargol impregnation and scanning electron micros-
copy). a, b) Ventral views, a = 170-205 !lm, b = size not indicated. c) Ventral infraciliature, size not indicated.
Arrow marks the single postoral ventral cirrus. This population differs from (a, b) in that (i) the undulating
membranes are distinctly curved, (ii) the left frontoventral row begins more anteriorly, and (iii) more (6-7
against 5) transverse cirri are present. d) Ventral infraciliature of an early morphogenetic stage; p 786.
 Paraurostyla 873

gatum (AMMERMANN & MUENZ 1982). According to JERKA-DZIADOSZ (1965) well fed P.
weissei divides 1-3 times per day at 18-20 °C.
BICK & KUNZE (1971) provided the following autecological data (summarised from
literature): 0.3-7.8 mg I" O2, 13.5-24.5 mg 1-' BODs. Avoids less than 3 mg 1-' O2 and
pH<6.5 (LIEBMANN 1962). Conversely, MATIS (1975) recorded P. weissei in a drainage at
1.3-2.2 mg 1-' O2 (15-20 °C, pH 6.9). DETCHEVA (1972a, 1975a, b, 1976b, 1979c, 1982b,
1983b, c, 1993) found it in Bulgarian running waters at following conditions: 0.9-3.3 %
frequency, 9-24 °C, pH 7.4-7.8,5.8-11 mg 1-' O2 (54-100 % saturation), 1.5-15.7 mg 1-'
BODs, 54-195 mg 1-' Ca2+, 0-80 mg 1-' Mg2+, 155-283 mg 1-' HC0 3-, 58-941 mg 1-' SOlo,
13-257 mg 1"' CI-, 0.02-1.53 mg 1-' NH/-N, 0.3-11.2 mg 1-' N03--N, 0-0.12 mg 1-' NOi-
N, 0-1.5 mg 1-' Fe2+. MnwLoWITSCH (1989) provided following autecoiogical data (n =
4-5) from populations found in salt-polluted running waters in Germany: 4.2-20.5 °C,
pH 7.6-7.9, 18.6-89.4 mg 1-' CO2, 8.0-9.3 mg 1"' O2, 0.09-D.5 mg 1-' NH/-N, 0.04 to
0.1 mg 1-' NOi-N, 2.3-3.0 mg 1"' N03--N.
Species of the Paraurostyla weissei complex are indicators of polluted waters (for
example, KOLKWITZ 1950, KOLKWITz & MARsSON 1909, MARsSON 1903, MAUCH 1976).
SLADECEK et al. (1983), WEGL (1983), and ForSSNER (1988a) classified it as strict aI-
phamesosaprobic species (a = 10). However, P. weissei occurs in a rather wide BOD·
range, proving that this classification is unrealistic. ForSSNER et aI. (1991) thus changed it
to a; b = 2, a = 7, P = 1, I = 3, SI = 2.9.

Supposed synonym of the Paraurostyla weissei complex

Oxytricha multipes CLAPARimE & LACHMANN, 1858

1858 Oxytricha multipes CuPARE1>E & LACHMANN, Mem. Inst. natn. genev., 5: 143, Planche 5, Fig. 1.
1876 Kerona multipes- FROMENTEL, Microzoaires, p 271, Planche XIII, fig. 19 (insufficient redescription).
1932 Urostyla (Oxytricha) multipes CLAP. u. L., 1858 - KAHL, TielWelt Ott, 25: 567, Fig. 9711 (redrawing
from original description).
1972 Urostyla multipes (CLAPAREDE & LACHMANN, 1858) KAm., 1932 - BoRROR, J. Protozool., 19: 9.
1974 Urostyla trichogaster STOKES, 1885 var. multipes (CLAPAREDE & LACHMANN, 1858) comb. n. - STIlLER,
Annis hist.-nat. Mus. natn. hung., 66: 132.

Rem ark s : Synonymy of Oxytricha multipes and Paraurostyla weissei is evident and
was already proposed by I<:ENT (1882, P 765). Differences are very likely due to some mi-
nor misobservations both by CLAPAREoE & !.ACHMANN (1858) and STEIN (1859b). To
avoid suppression of the well established name Paraurostyla weissei I consider Oxytri-
cha muitipes as supposed synonym. Only 2 faunistic records (HAsSDENTEUFELOvA-
MORAvcovA 1955, LIEPA 1978).
Previously, I also considered Oxytricha urostyla CLAPAREDE & LACHMANN, 1858
(p 141, Planche 5, fig. 2) as supposed synonym of Paraurostyla weissei (ForSSNER et al.
874 SYSTEMA TIC SECTION

1991, P 260). However, this species has about 10 frontal cirri arranged in a bicorona,
strongly suggesting that it is not identical with P. weissei (see Fig. 2271, m).

Paraurostyla granulifera BERGER & FOISSNER, 1989 (Fig. 216a-f, Table 50)

1989 Paraurostyla granulifera BERGER & FOISSNER, Bull. Br. Mus. nat. Hist. (Zoo!.), 55: 29 (type slides
[holotype: reference number 1988:2:1:20; paratype: 1988:2:1:21] are deposited in the British Museum
ofNaturaJ History in London).

T a x 0 nom y: The ventral infraciliature is almost identical to that of Trichotaxis fossi-

cola KARL (Fig. 217; note that Trichotaxis is an incorrect subsequent spelling of Tricho-
totaxis). However, we avoided an identification since KARL (1932) did not mention extru-
somes. But this argumentation is weak because KARL (1932) did not even described the
conspicuous mucocysts of Paraurostyla weissei. I thus propose T. fossicola as supposed
synonym of P. granulifera (see below). Redescription of a European population should
be awaited before definitive synonymisation.

M 0 r p hoI 0 g y and b i 0 log y: In life 170-230 x 70-100 /lm. Right margin

straight, left convex and in the anterior portion markedly indented. Anterior and posterior
end rounded. Flattened about 2: 1 dorso-ventrally. Flexible. Invariably (n = 7) 2 macronu-
clear nodules, distinctly left of median. Micronuclei in life about 7 x 5 /lm. Contractile
vacuole in about mid-body, during diastole with canals. Perpendicular to pellicle very
many, about 2 x I /lm sized, colourless cortical granules (extrusomes). Individual extru-
somes difficult to discern because very closely spaced, forming a distinct lucent seam
giving the organism a brownish colour at low magnification. Extrusomes ejected after ad-
dition of methyl green-pyronin, stain red, U- or horseshoe-shaped; among these extru-
somes many thin structures, probably fully exploded extrusomes. Cytoplasm colourless
with many crystals and food vacuoles. Rapid movement
Adoral zone of membranelies about 40 % of body length, bases of largest mem-
branelles in life about 8 /lm wide. Undulating membranes slightly curved, almost of same
length, paroral consists of short oblique kineties, at least in middle part. Invariably 3 (n =
7) distinctly enlarged frontal cirri. Invariably I (n = 7) buccal cirrus near anterior end of
paroral. Bases of buccal cirrus, the cirrus behind right frontal cirrus (invariably 1; n = 7),
and the in life about 25 /lm long frontal cirri distinctly enJarged. Left and right frontoven-
tral row begin about at level of the cirrus behind right frontal cirrus, left one terminates
about in mid-body, right one at the transverse cirri like the middle row, which begins at
about level of buccal vertex. One or 2 slightly enlarged postoral ventral cirri (Fig. 216f,
arrow). Transverse cirri in life about 30 /lm long, terminate at posterior edge of cell. Mar-
ginal rows almost confluent posteriorly, cirri in life about 20 /lm long. Invariably (n = 7)
3 caudal cirri. Dorsal cilia short. Dorsal infraciliature unknown because hidden by the ex-
trusomes, which often stain with protargol.
 Paraurostyla 875

,,
,
,
,
,

,
,,
,
I
,,
.,
• I •

....','.
,," ...---
•• :~,............. f

Fig. 216a-f Paraurostyla granulifera (from BERGER & FOISSNER 1989a 1Hl, from life; e, methyl green-
pyronin stain; t: protargol impregnation). a) Ventral view, 195 j.lm. b) Right lateral view. c) Dorsal view show-
ing the arrangement of the extrusomes (right half of illustration) and the contractile vacuole. d, e) Extrusomes
in lateral view and after ejection. t) Ventral infraciliature, 225 j.lm. Arrow marks single postoml ventral cirrus,
arrowhead marks caudal cirri; p 874.

o c cur r e n c e and e colo g y: Locus classicus is Ryu-Ga-

Take Mountain, Amakusa, Kumamoto Prefecture, Japan, where we
(BERGER & FOISSNER 1989a) discovered Paraurostyla granulifera in
the soil of deciduous forest at 470 m above sea level. This is a brown
soil with many fragments of leaves and roots and pH 4.8. Not found
since. Feeds on fungal spores, flagellates, cysts of amoebas, and testa-
ceans (Trinema enchelys).

Sup p 0 sed s y non y m: Trichotaxis fossicola KAHL, 1932, Tier-

welt Dtl., 25: 589 (Fig. 217). Remarks: KAHL (1932) classified Tricho-

Fig. 217 Trichotaxis fossicola from life (from KAHL 1932). Supposed synonym of
Paraurostyla granulifera, 200 j.lm. Note the single postoral ventral cirrus between buc-
cal vertex and anterior macronuclear nodule.
876 SYSTEMATIC SECTION

Table 50 Morphometric data of Paraurostyla granulifera (from BERGER & FO/sSNER I 989a). All data are
based on protargol-impregnated specimens. All measurements in micrometres. CV = coefficient of variation
(in %), Max = maximum value, mean = arithmetic mean, Min = minimum value, n = sample size, SD = stan-
dard deviation

Character mean SD CV Min Max n

Body, length 211.9 17.4 8.2 195 240 7
Body, width 92.7 12.7 13.7 70 Il2 7
Adoral membranelles, number 59.9 3.2 5.4 55 65 7
Adoral zone of membranelles, length 84.0 5.4 6.4 73 90 7
Macronuclear nodules, distance between 35.0 4.4 12.5 27 39 7
Posterior macronuclear nodule, length 26.1 4.0 15.4 22 34 7
Posterior macronuclear nodule, width 13.9 0.9 6.5 13 15 7
Micronuclei, number 4.1 1.1 25.8 3 6 7
Posterior micronucleus, length 5.8 0.9 15.5 4 7 7
Posterior micronucleus, width 3.4 0.5 15.6 3 4 7
Left marginal row, number of cirri 48.0 5.3 11.0 40 56 7
Right marginal row, number of cirri 40.0 1.8 4.6 38 43 7
Frontoventral rows, number 3.0 0 0 3 3 7
Left frontoventral row, number of cirri 18.3 2.4 13.3 15 22 7
Middle frontoventral row, number of cirri 14.0 2.0 14.3 12 18 7
Right frontoventral row, number of cirri 31.1 1.7 5.4 30 34 7
Postoral ventral cirri, number 1.4 0.5 37.4 I 2 7
Ventral cirri near transverse cirri, number 2.0 0 0 2 2 7
Transverse cirri, number 7.0 0.6 8.2 6 8 7
Distance II 30.0 6.7 22.3 21 39 7

I Distance between base of posterior-most transverse cirrus and posterior end of cell.

taxis (an incorrect subsequent spelling of Trichototaxis) as subgenus of Holosticha; the

correct name in his revision is thus Holosticha (Frichotaxis) fossicola. BORROR (l972a,
p 10) transferred it to Paraurostyla, P. fossicola. HEMBERGER (1982, p 30) proposed syn-
onymy with P. weissei, which sometimes also has only 3 ventral rows. However, the sin-
gle postoral ventral cirrus is reminiscent of Paraurostyla granulifera. I preliminarily
avoid a final synonymisation with our species because KAHL (1932) did not mention ex-
trusomes and it is known that almost identical species with and without cortical granules
exist. Furthermore, his population is from freshwater. However, further studies are
needed to solve this problem.
In life 200-220 Ilm long, slenderly elliptical. Two macronuclear nodules, each with a
single micronucleus. 4-5 enlarged frontal cirri, that is, 3 frontal cirri, 1 buccal cirrus, and
1 cirrus behind right frontal cirrus. 7-8 transverse cirri, protrude only slightly beyond
posterior end of cell (at fITSt KAHL found 4 or 5 specimens without transverse cirri). Three
frontoventral rows, middle one beginning at level of buccal vertex. Marginal cirri almost
confluent posteriorly, distinctly protruding beyond posterior end of cell. Locus classicus
unknown, but very likely near Hamburg, Germany, where KAHL (1932) lived and worked.
He found it only once with high abundance in freshwater on dead leaves covered with al-
 Paraurostyla 877

Fig. 218a Paraurostyla polynucleata (from ALEKPEROV 1993). Ventral infraciliature after Chatton-Lwoff sil-
ver impregnation and nuclear apparatus after Feulgen staining, 95 J.lm. Arrowhead marks curved cirral row be-
hind frontal cirri. Large arrow marks pore of contractile vacuole. Tiny arrows mark anterior end of left and
posterior end of middle frontoventral row respectively.
Fig. 218b, c Insufficient redescriptions of Paraurostyla weisse; (b, from ROSA 1957a; c, from GULAn 1925; b,
c, from life). Ventral views, b = size not indicated, c = 130 x 42 J.lm; p 878.

gae during spring. Records not documented by illustrations: benthal of Hamburg harbour,
Germany (BARTSCH & HARTWIG 1984); in a reservoir of the Seversky Donets Basin,
Ukraine (KRAvCHENKO 1969). Feeds on diatoms (KARL 1932).

Paraurostyla polynucleata ALEKPEROV, 1993 (Fig. 218a)

1993 Paraurostyla poiynucleata ALEKPEROV, Zoosystematica Rossica, 2: 26 (bolotype slide L(S-R) No.3 is
deposited in the Institute of Zoology, Academy of Sciences of Azerbaijan, Baku).

T a x 0 nom y: Detailed redescription and morphogenesis are needed to confirm the

classification in Paraurostyla.

M 0 r p hoi 0 g y and b i 0 log y: In life 150-170 ~m long, fixed cells up to

155 ~m. Body elliptical to oval, slightly flattened dorso-ventrally. Nine ellipsoidal to
reniform macronuclear nodules, 5-7 micronuclei. Contractile vacuole about at level of
S7S SYSTEMATIC SECTION

buccal vertex. Cytoplasm (endoplasm) light brown, without inclusions. Adoral zone of
membrane lIes about 37 % of body length, 55-60 adoral membranelles. Three distinctly
enlarged frontal cirri. Behind them 6 slightly enlarged cirri form a curved row beginning
near left frontal cirrus and terminating slightly in front of posterior end of buccal row.
Four buccal cirri. Left cirral row beginning at level of buccal vertex, terminating close to
right transverse cirrus. Middle frontoventral row beginning near distal end of adoral zone
of membranelIes, ending slightly behind mid-body. Right frontoventral row also begin-
ning near distal end of adoral zone, slightly shortened posteriorly; according to original
description this is the inner right marginal row (morphogenesis is needed to ascertain the
correct terminus). Nine obliquely arranged transverse cirri. Four "complete" dorsal kine-
ties and one shortened, that is, dorsal kineties very likely in Oxytricha pattern with 1 dor-
somarginal row. Caudal cirri not mentioned, that is, either absent or almost indistinguish-
able from marginal cirri and thus overlooked.

o c cur r e n c e: Locus classicus is Repino near st. Petersburg, Russia, where ALEK-
PEROV (1993) discovered Paraurostyla polynucleata in soil.

Insufficient redescriptions

Urostyla weissei - RosA, 1957, Pi'irodov. Sb. ostrav. Kraje, IS: 45 (Fig. 21Sb). Remarks:
In life 250 11m long. Soil in Czechoslovakia.

Urostyla weissii STEIN - GULATI, 1925, 1. Bombay nat. Hist. Soc., 30: 752 (Fig. 21Sc).
Remarks: Incorrect subsequent spelling. In life 130 x 42 Ilm. Long elliptical, slightly
converging anteriorly. Very flexible. Many macronuclear nodules. Contractile vacuole
slightly behind mid-body. Yellowish. Adoral zone of membrane lIes slightly more than
one third of body length. 3-5 frontal cirri, 5 ventral rows, 7-S transverse cirri. Marginal
rows confluent posteriorly. Among filamentous algae in freshwater from Lahore, India.
Especially the nuclear apparatus indicates that the identification is wrong.

Parentocirrus Voss, 1997

1997 Parentocirrus Voss, Europ. J. Protistol., 33: 31 - Type (original designation): Parentocirrus hortualis
Voss, 1997.

C h a r act e r i sat ion: Body little flexible. Adoral zone of membrane lIes formed like
a question mark. Undulating membranes in Oxytricha pattern. Numerous frontoventral
cirri arranged in at least two longitudinal rows. One left and 1 right row of marginal cirri.
Six dorsal kineties. Caudal cirri present. Six frontal-ventral-transverse cirri primordia,
number of cirri originating from primordia II-VI distinctly increased (compared with IS-
 Parentocirrus 879

cirri oxytrichids). Rightmost frontoventral row not involved in primordia formation. Dor-
sal morphogenesis in Oxytricha pattern.

T a x 0 nom y: See relevant chapter of single species.

Single species

Parentocirrus hortualis Voss, 1997 (Fig. 218d-k, Tables 50.1, 50.2)

1997 Parentocirrus hortualis Voss, Europ. J. Protistol., 33: 31 (Fig. 218d-h, j, k; two slides of protargol-
impregnated specimens with the holotype and several paratypes and one slide with specimens in divi-
sional morphogenesis are deposited in the OberOsterreichische Landesmuseum in Linz, Upper Austria.
Accession numbers: 27-29/1996).

T a x 0 nom y: In Parentocirrus hortualis dorsal morphogenesis proceeds according to

the Oxytricha pattern (Fig. 21Sk) strongly indicating that it is a member of the Oxytrichi-
dae. Conversely, Voss (1997) assigned it to the Kahliellidae because three long ventral
rows on the right side of the body and V-shaped primordia are present; furthermore, fron-
toventral row 6 (Fig. 21Sh) is not involved in primordia formation. However, all three
characters are unspecific and do not prevent a classification in the Oxytrichidae. Unfortu-
nately, as in some other taxa which do not have the typical IS-cirri pattern (for example,
Paraurostyla, Gastrostyla), I am unable to decide whether Parentocirrus is a ''primitive''
member of the oxytrichids or a "IS-cirri oxytrichid" with a secondarily increased number
of cirri. EIGNER (1997) classified it in the Parakahliellidae.

M 0 r ph 0 log y and b i 0 log y: In life 160-225 x 75-105 !lm. Outline elliptical,

both ends broadly rounded, flattened about 2: 1 dorso-ventrally. Body little flexible. Nu-
clear apparatus left of median. Contractile vacuole about in mid-body, with two incon-
spicuous collecting canals during diastole. Cytoplasm colourless, with many cytoplasmic
crystals and yellowish fat globules; hence, cells darkish in bright field at low magnifica-
tion. Cytoplasm of well-fed specimens with inconspicuous spherical inclusions, 1-2!lm
across, do not stain with protargol or methyl green-pyron in, "no special cortical granules"
(important note: according to the legend of Figure 3 in the Voss paper [= Fig. 21Sfin the
present book] "Conspicuous cortical granules which do not stain with methyl green-
pyronin"); consequently, this character has to be checked in further populations! Newly
excysted specimens appear slim, anterior portion of body bent to the left, cytoplasm clear
and colourless.
Adoral zone of membranelles about 50 % of body length (according to Table 50.1,
about 46 %). Buccal area large and deep, opens and enlarges by shifting undulating mem-
branes to right body margin; thus, cell is able to ingest very large food particles. Undulat-
ing membranes in Oxytricha pattern, that is, curved and crossing optically; endoral termi-
880 SYSTEMATIC SECTION

Fig. 21Sd-g Parentocirrus hortualis (from Voss 1997. d--g, from life). d) Ventral view, 200 Jim. e) Resting
cyst, 85 Jim. 1) Detail of surface showing "conspicuous cortical granules which do not stain with methyl green-
pyronin" (original figure legend); however, according to text "no special cortical granules" present (bar scale
division = 5 Jim). g) Body shape ofa recently encysted specimen; p 879.

nates at proximal end of adoral zone of membranelles, paroral slightly in front of it (Fig.
218h). Cirri in life about 25 ~m long. Bases of frontal cirri distinctly enlarged. Arrange-
ment of frontoventral rows, see Figure 218h. I have no doubt that the frontoventral cirral
rows 1-6 (Fig. 218h) are homologous to the primordia I-VI of the 18-cirri oxytrichids
(Fig. 6a). 3-4 only slightly enlarged transverse cirri (thus, the mean of 4.1 in Table 1 of
Voss 1997 is probably incorrect), do not protrude beyond posterior end of cell (Fig.
218d). Marginal rows slightly separated posteriorly. Dorsal kineties 1-4 about as long as
body, kinety 5 about half of body length with 10-13 basal body pairs, kinety 6 usually
with 5-7 pairs; rarely a third dorsomarginal kinety (kinety 7) present. Dorsal cilia in life
about 6 ~m long. Usually 1 caudal cirrus each at posterior end of dorsal kineties 1,2, and
4, that is, as in most oxytrichids.
 Parentocirrus 881

-
-
II

""
/II

I
•
•
.. ••
fI'· h

Fig. 218h, i Parentocirrus hortualis (h, from Voss 1997; i, original kindly supplied by H. BLAlTERER, Linz; h,
protargol impregnation; i, from life). h) Ventral infraciliature (168 11m) and nuclear apparatus. Cirri originating
from same cirra1 primordium are connected by a dashed line. Arrows mark filamentous tie between macronu-
clear nodules. i) Ventral view, 165 )lm. 1-6 = frontal-ventral-transverse cirri anlagen I-VI; p 879.

Rarely specimens with a supernumerary cirral row: 1.1 % (17 of 1534 specimens)
with a row between frontoventral row 6 and right marginal row, 0.6 % with a cirral row
between frontoventral rows 5 and 6,0.06 % with a cirral row between frontoventral rows
4 and 5, and 0.3 % with a supernumerary left marginal row. Slow movement, creeping
and gliding on substratum, sometimes resting for a moment.
Resting cysts about 42-74 /lm in diameter (Table 50.1; according to Fig. 218e, how-
ever, about 80--85 /lm!), outer cyst wall covered with scaly, hyaline, mucous layer, inner
wall smooth.

M 0 r p hog e n e sis: Detailed description and documentation with illustrations and mi-
crographs of protargol-impregnated specimens, see Voss (1997). Some important data
are summarised in Table 50.2. The right frontoventral row is not involved in primordia
 882 SYSTEMATIC SECTION

J l
J )
3J
J
I [

I f
J
J ,f
t
f
J C'
.!
, ! J r
(
'/ .( r
1 .
:{ ,I
'i
" , I
r
" .'
';
) f
'/ / ,r
'f \
'/ " :-
"J
I r ,r
I~, 5J : r /~

CC~
'I
,\
\ to, \ t
- '.......
"
"
./'
.1 ,r
.
.!
j
\ 0<)0
k

Fig. 218j, k Parentocirrus hortualis (from Voss 1997. j, k, protargo1 impregnation). j) Dorsal infraciliature,
170 J.lm. k) Middle stage of dorsal morphogenesis, 190 J.lm. Arrows mark fragmentation of new dorsal kineties
3. CC = caudal cirri (on dorsal kineties 1,2, and 4), 1-{) = dorsal kineties 1-{) (kineties 5 and 6 are dorsomar-
ginal rows); p 879.

fonnation; this is a very important difference to Apoamphisiella, where both frontoven-

tral rows are involved. Dorsal morphogenesis of Parentocirrus proceeds in Oxytricha
pattern indicating that this species is closely related to the other taxa revised in the pre-
sent book. Fragmentation of a dorsal kinety was never described in (typical) kahlielids, to
which Parentocirrus was originally assigned.

o c cur r e n c e and e colo g y: Locus classicus is the village of Feldhausen in

Bottrop, Gennany, where Voss (1997) discovered P. hortualis in a bird bath in August.
H. BLATIERER (Fig. 218i) found it in the activated sludge of a paper-mill in Salzburg,
Austria. Voss (1997) isolated several specimens and put them into Petri dishes together
with water and some leaf mud from the sampling site. Non-carbonated mineral water and
squeezed wheat grains were added to establish raw cultures. Clone cultures were set up
from these raw cultures in non-carbonated mineral water together with a few drops of
 Parentocirrus 883

Chlorogonium elongatum concentrated by centrifugation and mixed with a suspension of

baker's yeast as food. Feeds also on green algae, diatoms, mineral particles, cysts of
amoebas and of conspecifics, and eggs of rotifers, for example, Rotaria rotatoria.

Table 50.1 Morphometric data of Parentocirrus hortualis (from Voss 1997). All data are based on protargol-
impregnated specimens, except otherwise indicated. All measurements in micrometres. CV = coefficient of
variation (in %), Max = maximum value, mean = arithmetic mean, Min = minimum value, n = sample size, SD
= standard deviation

Character mean SD CV Min Max n

Body, length (in life) 186.6 22.2 ll.8 163 223 10
Body, length 183.1 37.2 20.6 128 301 25
Body, width (in life) 86.9 ILl 12.8 74 104 10
Body, width 94.7 22.5 23.8 62 166 25
Macronuclear nodules, number 7.8 1.5 19.2 6 12 25
Macronuclear nodules, diameter 13.4 3.6 26.1 9 26 25
Micronuclei, number 3.5 1.0 28.6 2 6 25
Micronuclei, diameter 3.9 0.4 10.0 3 5 25
Adoral zone of membranelles, length 85.0 22.3 26.2 55 158 25
Adoral membranelles, number 45.0 4.6 10.2 38 57 25
Frontal cirri, number 3.0 0 0 3 3 25
Cirri behind right frontal cirrus, number 2.0 0 0 2 2 25
Buccal cirri, number 2.8 0.5 17.9 2 4 25
Frontoventral row 4, number of cirri 1 4.2 1.3 31.0 3 9 25
Frontoventral row 5, number of cirri 1 19.1 2.4 12.6 16 24 25
Frontoventral row 6, number of cirri I 24.3 2.2 9.1 19 30 25
Transverse cirri, number 4.1 0.3 7.3 3 4 25
Right marginal row, number of cirri 32.2 4.0 12.4 23 42 25
Left marginal row, number of cirri 32.0 4.4 13.8 23 43 25
Dorsal kineties, number 6.1 0.3 4.9 6 7 25
Caudal cirri, number 3.1 0.3 9.7 3 4 25
Cyst, diameter (in life) 49.6 10.2 20.6 42 74 10

I Designation, see Figure 218h. Without transverse cirrus (the posterior-most cirrus of the row).
2 Uncertain data because mean (4.1) higher than maximum value (4).

Table 50.2 Origin of frontal-ventral-transverse cirri primordia and number of cirri produced in the primordia
of Parentocirrus hortualis (from Voss 1997)

Primordium1 Proter Opisthe Number of cirri

undulating membranes oral primordium 1
2 buccal cirri oral primordium 3-5
3 posterior frontal cirri oral primordium 4-5
4 frontoventral row 4 frontoventral row 4 5-7
5 frontoventral row 52 frontoventral row 5 17-25
6 frontoventral row 5 frontoventral row 5 20--31

I Primordia 1-6 correspond to primordia I-VI of the 18-cirri oxytrichids (see Fig. 6a). Designation offron-
toventral rows, see Figure 218h.
2 Primordium 5 ofproter is also generated by frontoventral row 4.
884 SYSTEMATIC SECTION

Territricha BERGER & FOISSNER, 1988

1988 Territricha BERGER & FOISSNER, Zoo!. Anz., 220: 127 - Type (original designation): Territricha stra-
menticola BERGER & FOISSNER, 1988.

C h a r act e r i sat ion: Body flexible. Adoral zone of membranelies formed like a
question mark. Undulating membranes in Oxytricha pattern. Frontoventral cirri form V-
shaped pattern. More than 3 postoral ventral cirri behind buccal vertex. Two pretrans-
verse ventral cirri and more than 5 transverse cirri. One right and I left row of marginal
cirri. More than six dorsal kineties. Caudal cirri present. Number of frontal-ventral-
transverse cirri primordia >6. Dorsal morphogenesis in Oxytricha pattern, however, ki-
nety 3 not with simple, but with multiple fragmentation.

T a x 0 n 0 my: For a description of the patterns mentioned in the characterisation, see

chapter 2 in the general section. Further details, see relevant chapter of single species.

Single species

Territricha stramenticola BERGER & FOISSNER, 1988 (Fig. 219a-j, Table 51)

1988 Territricha stramentico/a BERGER & FOISSNER, Zoo!. Anz., 220: 128 (I slide of ho[otype and 1 slide of
paratype specimens are deposited in the OberOsterreichische Landesmuseum in Linz, Upper Austria).

T a x 0 nom y: Territricha is monotypic and was originally classified in the Urostylidae

following BORROR & WICKLOW'S (1983) definition in that in this group "frontal and mid-
ventral cirri differentiate during pre-fission morphogenesis from the longitudinal field of
more than 5 (without streak I) oblique ciliary streaks". We assumed that Territricha stra-
menticola links typical urostylids,~uch as Holosticha, with typical oxytrichids, like, for
example, Oxytricha (BERGER & FOI15~R 1988a). A classification in the amphisiellids, as
proposed by TUFFRAU & FLEURY (1994, p 141), is very likely artificial because a fragmen-
tation of dorsal kineties and an arrangement of cirri in couples is unknown in this group.
I include Territricha in the Oxytrichidae because it shows, like all other oxytrichids,
fragmentation of dorsal kinety 3. This character is so conspicuous that parallel evolution
is very unlikely. However, I have no idea if the increased (that is, >6) number of cirral
primordia is plesiomorphic or if it originated by secondary polymerisation (Fig. 219j), as,
for example, in Onychodromus quadricornutus. A relationship to the Stylonychinae,
however, can be excluded because the body is flexible and extrusomes are present.
Territricha stramenticola should not be confused with Holosticha macrostoma
(DRAGESCO, 1972) BORROR, 1972 (needs redescription; very broad buccal area, 5 dorsal
kineties) or Pattersoniella vitiphila (larger, rigid cortex, 13-15 macronuclear nodules,
about 5 contractile vacuoles, Fig. 197a).
 Territricha 885

'I·:
::.:.':>
":'.:)

-::':::.
.
~:::,
":.:.7!

2: d

Fig. 219a-f Territricha stramenticola (from BERGER & FOISSNER 1988a a--e, from life; f, methyl green-
pyronin stain). a) Ventral view, 135 ~m. The dorsal cilia of the anterior portion of kinety 1 (arrowhead) are
distinctly longer (about 7 ~m) than the remaining dorsal cilia (about 3 ~m). Note distinctly anteriad displaced
transverse cirri. b) Right lateral view. c) Dorsal view showing the contractile vacuole and the fringe of extru-
somes. d, e) Side and surface view of extrusomes, 3-5 x 0.8-1 ~m. f) Extrusomes after ejection; p 884.

FOISSNER (1993, P 7, 501; 1994e, p 225) confused Territricha with another similarly
named hypotrich, namely Terricirra. The latter has fusifonn food vacuoles like the colpo-
dids Parabryophrya and Notoxoma.

M 0 r p hoI 0 g y and b i 0 log y: In life 100-175 x 50-75 Jim. Right margin

straight, left convex, in anterior portion markedly indented. Anterior and posterior end
rounded. Flattened about 2: 1 dorso-ventrally. Body flexible. Two macronuc1ear nodules,
in life 25-35 x 10-17 ~m, fmely granulated, lying distinctly left of median of cell. Micro-
nuclei in life about 6-8 x 3-5 ~m. Contractile vacuole at about mid-body, during diastole
with canals. Cells more or less brownish at low magnification. Very many fusiform, in
life about 3-5 x 0.8-1 ~m sized, colourless extrusomes arranged perpendicular to pellicle
and making a bright fringe at middle magnification; extrusomes ejected after addition of
methyl green-pyronin, stain blue, comifonn (hom-shaped, hollow), about 5-8 ~m long;
 00
00
0\
23 Y
\ / ~
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\\11\,(1/ ~
:' ,f ~ "-'/'
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---;: ,r \ ~ Ii' 0 ....
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r \ ~\~\mS=
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:::t. ...••:.: ..
\,\ ~:(~
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••• \ :!~;~
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'-l.
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I, .P, ~r' .' \ ,jl ", ::l
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,,
-~ .......' ~. ". ~
,, l/f'' ( J" " ~
'~~~ilJ rn
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,, ,"'-
',-I.
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-t', (I
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" 0
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:(J
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if
c:J
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>-l
, • t ~~ 0 c:J
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, 'I I, (
,, "'~\',' . .I 00°
',0 CJ ~
, ~ '\, f ,/1 o
i\' 'I (',,~i. c§) CJ
D
,,
CJ
I·.
,, x.:x" • f ( / 1 " ('
'I " - I, a
\x' ,/ ' '/ c:J
c:J
~I,
I, v' (' --, J I' oG
If, ; ,(' f'-" /'
; - .' I
~i J
Fig. 219g-j Territricha stramenticola (from BERGER & FOISSNER 1988a Protargol impregnation). g, b) Ventral infraciliature of specimens with 7 and 9 postoral ventral
cirri, g = 110 11m, h = 115 11m. Arrows mark the 2 pretransverse ventral cirri. I) Dorsal infraciliature, 115 11m. Dorsal kinety 3 shows multiple fragmentation and the
posterior-most fragment with the caudal cirrus is denoted as kinety 4. Arrowheads mark the dorsomarginal kinety arranged next to kinety 4 and the right marginal row
(see also next Figure). j) Late reorganisation stage, 90 11m. Arrowheads mark the right- and leftmost dorsomarginal kinety primordium (in i the same dorsomarginal ki-
neties are marked with arrowheads). Omitting 2 of the frontal-ventral-transverse cirri streaks marked with an asterisk results in the well-known I8-cirri pattern of typical
oxytrichids (cp. Fig. 6a). Old structures white, new ones black. Cirri originating from same primordium are connected by a broken line. 1-4 = dorsal kineties 1-4; p 884.
 Territricha 887

Table 51 Morphometric data of Territricha stramenticola (from BERGER & FOISSNER 1988a). All data are
based on protargol-impregnated specimens. All measurements in micrometres. CV = coefficient of variation
(in %), Max = maximum value, mean = arithmetic mean, Min = minimum value, n = sample size, SD = stan-
dard deviation

Character mean SD CV Min Max n

Body, length 111.5 6.7 6.0 99 126 22
Body, width 42.7 4.1 9.5 36 50 21
Adoral membranelles, number 42.2 2.5 5.8 39 47 15
Adoral zone of membranelles, length 44.8 3.0 6.6 39 50 21
Posterior macronuclear nodule, length 18.5 2.0 10.9 15 22 21
Posterior macronuclear nodule, width 10.0 1.1 10.9 8 12 21
Macronuclear nodules, distance between 5.4 2.2 41.7 2 10 21
Micronuclei, number 2.5 0.7 29.2 2 4 10
Posterior micronucleus, length 5.3 0.6 10.8 4 6 20
Posterior micronucleus, width 4.5 0.5 1l.S 4 5 20
Left marginal row, number of cirri 32.8 2.4 7.2 29 38 15
Right marginal row, number of cirri 33.4 3.0 9.1 27 39 18
Frontal cirri, number 3.0 0 0 3 3 21
Buccal cirri, number 1.0 0 0 1 21
Transverse cirri, number 6.8 0.8 12.2 4 8 19
Distance 11 17.2 3.6 21.1 8 23 21

I Distance between posterior-most transverse cirrus and posterior end of cell.

do not impregnate with protargol (Fig. 219c-f). Posterior portion of cell with many
1-5 ~m sized, yellowish cytoplasmic crystals and numerous homogenous 2-5 ~m sized
globules. Food vacuoles 10--20 ~m across. Rapid movement.
Adoral zone of membranelles about 40 % of body length, bases of largest mem-
branelles in life about I 0 ~m wide. Buccal area rather narrow, roofed by a prominent
peristomial lip. Undulating membranes slightly curved, almost the same length. Frontal
cirri in life about 25 ~m long, bases distinctly enlarged. Buccal cirrus almost at anterior
end of undulating membranes. Cirri right of adoral zone arranged as in typical oxytrichids
(for example, Oxytricha). Number of postoral ventral cirri >3 (7-9) due to higher (usu-
ally 2 or 3) number of primordia and not due to increased number of cirri originating
within a primordium. Arrangement of cirri rather invariably as shown in Figures 219g, h;
postoral cirri in life about 20 ~m long. Invariably (n = 14) 2 pretransverse ventral cirri.
Transverse cirri in life about 30 ~m long, bases distinctly enlarged and displaced anteriad
so that only posterior transverse cirri sometimes protrude slightly beyond posterior end of
cell. Marginal cirri in life about 20 ~m long. Right marginal row begins at about level of
buccal cirrus, posteriorly only slightly separate from left one. Dorsal infraciliature rather
complicated (Fig. 219i). Basal body pairs, especially in kinety I, very closely arranged.
Dorsal cilia in anterior portion ofkinety 1 about 7 ~m, in posterior portion and remaining
kineties about 3 ~m long (Fig. 219a, arrowhead). In posterior portion of cell, between ki-
neties 3 and 4, about 3 short kineties; very likely they are the remainder (except kinety 3
888 SYSTEMA TIC SECTION

and 4) of the multiple fragmentation of kinety 3. Constantly (n = 9) 3, in life 15-20 Jim

long caudal cirri, one each on posterior end of kineties 1,2, and 4. About 4-5 dorsomar-
ginal kineties (Fig. 2I9j).
Figure 2I9j shows a reorganisation stage, indicated by the presence of both the pa-
rental migratory cirri, the parental right frontal cirrus, and the parental transverse cirri.
Eight frontal-ventral-transverse cirri primordia occur, from which the following number
of cirri develop: 1,3,3,3',3',3',4,4 (Fig. 219j). Omitting two of the streaks marked
with an asterisk results in the well-known I8-cirri pattern of typical oxytrichids.

o c cur r en c e and e colo g y: Very common in litter (species name!) of beech

forests and often associated with Sterkiella cavicola and Rigidocortex octonucleatus
(own observations); very likely a character species of (Eurasian?) beech forests! Locus
classicus is the Gaisberg (at about 800 m above sea level on the western ridge), a moun-
tain near the city of Salzburg, Austria, where BERGER & FOISSNER (1988a) discovered Ter-
ritricha stramenticola in the above mentioned habitat. BONKOWSKI (1996) found it in the
same biotope in Gennany. Feeds on fungal spores of different shape and size and ciliates
like Colpoda sp. Biomass of 106 individuals about 140 mg.

Pseudouroieptus HEMBERGER, 1985

1985 Pseudouroleplus HEMBERGER, Arch. Protistenk., 130: 398 - Type (original designation): Pseudourolep-
Ius caudatus HEMBERGER, 1985.

C h a r act e r i sat ion: Body flexible? Adoral zone of membranelles fonned like a
question mark. Undulating membranes in Oxytricha pattern. Very many frontal-ventral
cirri, most of them fonn two frontoventral rows. Postoral ventral cirrus (usually) present.
Transverse cirri lacking or strongly reduced in number. Four dorsal kineties. Caudal cirri
present. Left frontoventral row composed of cirri from streaks IV-VI. Right row origi-
nates from streak V, that is, right frontoventral row not involved in primordia fonnation.
Dorsal morphogenesis with fragmenting kinety 3; dorsomarginal kinety lacking.

T a x 0 nom y: Five species are assigned to Pseudouroleptus: Pseudouroleptus cauda-

tus HEMBERGER, 1985 (type; see below); Pseudouroleptus buitkampi (FOISSNER, 1982)
BERGER & FOISSNER, 1987 (basionym: Paraurostyla buitkampi FOISSNER, Arch. Protis-
tenk., 126: 40, Abb. 4a-f, Tabelle 7); Pseudouroleptus humicola (GELLERT, 1956) HEM-
BERGER, 1985 (basionym: Uroleptus humicola GELLERT, Acta bioI. hung., 6: 245, Abb.
10); Pseudouroleptus procerus BERGER & FOISSNER, 1987, ZooI. Jb. Syst., 114: 195, Fig.
1-6, Table 2; Pseudouroleptus terrestris HEMBERGER, 1985, Arch. Protistenk., 130: 399,
Abb. 2. However, I have some doubt that these 5 species fonn a monophyletic group be-
cause the dorsal ciliary pattern is rather different: 2 kineties in P. procerus, 3 kineties in
P. buitkampi, unknown number in P. humicola, and 4 in P. terrestris and P. caudata.
Such a high variability is unusual in oxytrichid genera. Furthennore, fragmentation of
 Pseudouroleptus 889

dorsal kinety 3 - the sole synapomorphy of the species treated in the present book - is de-
scribed only for the type species. Thus, I confme Pseudouroleptus to P. caudatus, how-
ever, without formally transferring the other species to a different taxon.
Pseudouroleptus was originally classified in the Amphisiellidae (HEMBERGER 1982,
1985) and later transferred to the Kahliellidae by TUFFRAU (1987, p 115; see also TUFFRAU
& FLEURY 1994, P 137). EIGNER (1997) assigned it to the Oxytrichidae, because it shows
"neokinetaI3" anlagen development (for explanation, see EIGNER 1997, P 555).

Pseudouroleptus caudatus HEMBERGER, 1985 (Fig. 219.la-j)

1985 Pseudouroleptus caudatus HEMBERGER, Arch. Protistenk., 130: 398 (type slides are deposited in the In-
stitut filr landwirtschaftliche Zoologie, Bonn, Germany).

T a x 0 nom y: See also same chapter in genus section. Pseudouroleptus caudatus is as-
signed to the Oxytrichidae because dorsal kinety 3 shows simple fragmentation (Fig.
219.lj), that is, the posterior portion of primordium 3 splits off from the anterior portion
to form kinety 4 of the postdivider. Furthermore, morphogenesis of the ventral ciliature
closely resembles that of Gastrostyla, because the left frontoventral row of Pseudouro-
leptus caudatus consists of the same parts as the frontoventral row of Gastrostyla (cp.
Fig. 219.1 i and Fig. 202k, m, 0): the anterior portion is the anterior part of streak VI; the
middle portion is streak IV, except the prominent postoral ventral cirrus (likely homolo-
gous to cirrus IV/2 of the 18-cirri oxytrichids, see Fig. 6a); the posterior portion is streak
V. As in Gastrostyla, cirri of the anlage VI are not involved in primordia formation
which is an important difference to Apoamphisiel/a.
Pseudouroleptus terrestris has 4 buccal cirri and 2 transverse cirri. Pseudouroleptus
buitkampi has 4 macronuclear nodules, 2-4 buccal cirri, and 2-3 cirri behind the right
frontal cirrus. Pseudouroleptus humicola has many (about 32) macronuclear nodules.
Pseudouroleptus procerus has 3-4 buccal cirri and only 2 dorsal kineties.

M 0 r p hoi 0 g y: In life(?) about 250 x 50 /lm. Body elongate, slightly spiralled, ante-
rior end broadly rounded, posterior tapered. Body not contractile (HEMBERGER 1985), but
very likely rather flexible. Two ellipsoidal macronuclear nodules, 2-3 micronuclei. Con-
tractile vacuole in "standard position", that is, on left body margin slightly behind buccal
vertex. Cytoplasm brownish (possibly cortical granules are present). Adoral zone of
membranelles about 25 % of body length, composed of 33-50 (mean = 40) membra-
nelles. Invariably 3 distinctly enlarged frontal cirri, 1 buccal cirrus near anterior end of
undulating membranes which are in Oxytricha pattern, and 1 enlarged cirrus behind right
frontal cirrus (homologous to cirrus HII2 of the I8-cirri oxytrichids). One cirrus immedi-
ately behind buccal vertex (sometimes lacking). Left frontoventral row composed of
about 42 cirri, right row consists of about 55 cirri; both rows begin near distal end of ado-
ral zone of membranelles, left row terminates distinctly subterminally, right at posterior
 890 SYSTEMATIC SECTION

I;
- =/

~\- - -\.-
-,
... ~.
:;
-I
:/
- :/
. . :1
. :7
-.
\'. }
Vg
Fig. 219.1a-g Pseudouro/eptus caudatus (from HEMBERGER 1982. a-g, protargol impregnation). a) Ventral in-
fraciliature and nuclear apparatus of a non-dividing specimen, 233 Ilm. Arrow marks postoral ventral cirrus
which is very likely homologous to cirrus IV12 of the 18-cirri oxytrichids. b-g) Ventral ciliature of early (b-d)
and middle (e-g) morphogenetic stages. The primordia V and VI originate from the middle portion of the left
frontoventral row. Note that the right frontoventral row is not involved in primordia formation. OP = oral pri-
mordia; p 889.
 Pseudouroleptus 891

\\

Fig. 219.1h-j Pseudouroleptus caudatus (from HEMBERGER 1982. b-j, protargol impregnation). b) Ventral in-
fraciliature and nuclear apparatus of a late morphogenetic stage. Arrows mark the 6 frontal-ventral cirri pri-
mordia of the opisthe. i) Ventral infraciliature and nuclear apparatus of a very late morphogenetic stage. Ante-
rior arrow marks the anterior portion of primordium VI which migrates to the left to form the anterior portion
of the left frontoventral row in postdividers. The posterior arrow denotes the posterior-most cirrus of primor-
dium IV which becomes the conspicuous postoral ventral cirrus of the interphasic specimen; the anterior cirri
of primordium IV form the middle portion of the left frontoventral row. When morphogenesis is finished the
left frontoventral row appears as continuous cirra1 row, although composed of three parts (Fig. 219.1 a). j) Dor-
sal infraciliature of middle morphogenetic stage showing fragmentation (arrows) of kinety 3. Obviously no
dorsomarginal kineties are formed in Pseudouroleptus caudatus; p 889.

end. Distinct transverse cirri lacking. About 55 right and 45 left marginal cirri, about
20 J.lm long. Dorsal cilia about 6 /lm long, arranged in 4 kineties. Caudal cirri not dis-
tinctly longer than marginal cirri, that is, about 20 /lm; each one cirrus on dorsal kineties
1,2, and 4.

M 0 r p hog e n e sis is described and illustrated by HEMBERGER (1982, P 36; see Fig.
219.1b-j). It commences with the formation of two oral primordia (Fig. 219.1b). As in
many other oxytrichids the buccal cirrus, the cirrus behind the right frontal cirrus, and the
postoral ventral cirrus are involved in primordia formation (Fig. 219.1 c-e). The middle
portion of the left frontoventral row is transformed to the primordia V and VI of the pro-
ter and the opisthe (Fig. 219.ld-f). Later, these primary primordia divide (Fig. 219.1g,
h). As most other oxytrichids, Pseudouroleptus caudatus forms 6 frontal-ventral cirri
892 SYSTEMATIC SECTION

primordia (Fig. 219.lh, arrows). No cirrus of the right frontoventral row is involved in
primordia fonnation. A late morphogenetic stage shows that the left frontoventral row
originates, as in Gastrostyla, of cirri of streaks VI, IV, and V (from anterior; details see
Fig. 219.Ii). This strongly indicates a close relationship of Pseudouroleptus and Gas-
trostyla; by contrast EIGNER (1997) classified Pseudouroleptus in the Oxytrichidae and
Gastrostyla in the Parakahliellidae. Dorsal morphogenesis shows a simple fragmentation
in dorsal kinety 3 (Fig. 219.lj, arrows) resulting in 4 kineties during interphase because
no dorsomarginal kineties are fonned.

o c cur r e n c e and e colo g y: Terrestrial and limnetic. Locus classicus is the

suburb of Puerto Maldonado (about lat 12°3(')', long 69°12'), Peru, where HEMBERGER
(1985) discovered Pseudouroleptus caudatus in a forest soil (details see HEMBERGER
1982, P 2). He found it also in the plankton of a river (Rio Tambopata) from this region,
possibly due to soil erosion. No further records known. Biomass of 106 specimens about
375 mg (FOISSNER 1987a).
Taxa not Considered
The following taxa, although included by some authors in the Oxytrichidae or Oxytrichi-
dae sensu lato, were not considered in the present book because they lack the typical 18
frontal-ventral-transverse cirri and/or fragmentation of a dorsal kinety or they are species
indeterminata.

Balladinopsis GHoSH, 1921, 1. R. microsc. Soc., year 1921: 248 - Type (monotypy): Balladinopsis nuda
GHOSH, 1921. Remarks: Classified in the Oxytrichinae by JANKOWSKI (1979). Bal/adynopsis and Bal/adynopsis
nuda in KAHL (1932, P 592) are incorrect subsequent spellings. Contractile vacuole on right side (misobserva-
tion?). A species indeterminata

Dipleurostyla Roux, 1899, Revue suisse Zoo!., 6: 615 - Type (monotypy): Dipleurostyla acuminata Roux,
1899. Remarks: Both the type species and D. tristyla TAGUANI, 1922 are species indeterminata According to
KAHL (1932, P 650) both species are regenerating stylonychias.

Drepanina SAVI, 1913, Monitore zoo!. ital., 24: 99 - Type (monotypy): Drepaninafalcata SAVI, 1913 (Fig.
220c). Remarks: Species indeterminata. According to KAHL (1932, p 650) the anterior end of a large Stylony-
chiao

Engelmanniella FOISSNER, 1982, Arch. Protistenk., 126: 66 - Type (original designation): Uroieptus mobilis
ENGELMANN, 1862. Remarks: Engelmanniella was only provisionally assigned to the Oxytrichidae. Later, mor-
phogenetic data revealed that it is very likely a kahliellid (EIGNER 1995, WIRNSBERGER-AEscHT & FOISSNER
1989).

Hemisincirra HEMBERGER in FOISSNER, 1984, Stapfia, 12: 119 - Type (original designation): Uroleptus kahli
BUITKAMP, 1977a. Remarks: FOISSNER (1984) used the name Hemisincirra before the original description was
published (HEMBERGER 1985, p. 408). Hemisincirra comprises several, usually very slender soil inhabiting spe-
cies, which do not, however, have a very uniform infraciliature (for example, BERGER & FOISSNER 1987a,
1989a, FOISSNER 1982, 1984, HEMBERGER 1982, 1985). I do not include it in the Oxytrichidae because both the
fragmentation of a dorsal kinety and the typical 18 frontal-ventral-transverse cirri pattern are lacking. Morpho-
genetic data are needed to clarifY the phylogenetic relationships of Hemisincirra. However, two species (Hemi-
sincirra polynucleata, Perisincirra similis) have the frontoventral cirri arranged very similarly to Urosoma and
were thus transferred to this taxon in the present book.

Lamtostyla BurrKAMp, 1977, Acta Protozoo!., 16: 270 - Type (original designation): Lamtostyla lamottei BUIT-
KAMP, 1977. Remarks: Lamtostyla was originally classified in the Holostichidae and later transferred to the
Cladotrichidae (SMALL & LYNN 1985), however, without detailed explanation. Previously, I proposed a classifi-
cation in the Oxytrichidae because of morphogenetic similarities with typical oxytrichids (BERGER & FOISSNER
1987a, 1988a). PETZ & FOISSNER (1996) provided morphogenetic evidence that Lamtostyla belongs to the am-
phisiellids. However, some species (for example, L. longa, L. raptans) have the 18 frontal-ventral-transverse
cirri pattern and possibly belong to Tachysoma, where they were originally established.

Prosopsenus ANDRE, 1916, Revue suisse Zool., 24: 632 - Type (monotypy): Prosopsenus sinuatus ANDRE,
1916. Remarks: Classified in the Oxytrichinae by JANKOWSKI (1979). A species indeterminata, as already sup-
posed by KAHL (1932, P 650). No records available.

Psammomitra BORROR, 1972,1. Protozool., 19: 15 - Type (monotypy): Mitra radiosa QUENNERSTEDT, 1867.
Remarks: Psammomitra is the replacement name for Micromitra KAHL, 1933, Tierwe\t N.- u. Ostsee,23: 112,
which was the replacement name for Mitra QUENNERSTEDT, 1867, Acta Univ. lund., 4: 41. Some authors (for
894 SYSTEMATIC SECTION

example, KARL 1935, BORROR 1972a) classified it in the Oxytrichidae because it has a ventral ciliature alleg-
edly very similarly to 18-cirri oxytrichids. SONG & WARREN (1996), however, proved that it belongs to Urolep-
Ius EHRENBERG, 1831.

Psilolricha STEIN, 1859, Lotos, 9: 5 - Type (original designation and monotypy, respectively): Psilolricha
acuminala STEIN, 1859. Remarks: For a detailed description of this curious species, see STEIN (l859b) and
FOISSNER (1983b) and references therein. The rigid body strongly suggests an inclusion in the Stylonychinae.
However, neither ventral and dorsal infraciliature nor morphogenesis are reminiscent of that of oxytrichids. I
have thus not considered it in the present book. FOISSNER (1983b) proposed a close relationship with Uroleploi-
des, Pseudouroleptus, Kahliella, Cladolricha, and Kerona. EIGNER (1997) classified it in the Orthoamphisielli-
dae.

Terricirra BERGER & FOISSNER, 1989, Bull. Br. Mus. nat. Hist., 55: 35 - Type (original designation): Perisin-
ci"a viridis FOISSNER, 1982. Remarks: Te"icirra includes three soil inhabiting species with green to blue-
green cortical granules and spindle-shaped food vacuoles, short undulating membranes which form an acute
angle, and one short frontoventral row. The ventral infraciliature is reminiscent of that of Lamtosty/a, which is
now classified in the Amphisiellidae (PETZ & FOISSNER 1996). Because fragmentation of a dorsal kinety and the
typical 18 frontal-ventral-transverse cirri pattern are lacking, r do not include it in the Oxytrichidae. Morphoge-
netic data are needed to clarify the phylogenetic relationships of Te"ici"a.

Trachelochaela SRAMEK-HuSEK, 1954, Arch. Protistenk., 100: 265 - Type (original designation): Trachelocha-
eta bryophila SRAMEK-HuSEK, 1954. Remarks: Classified in the Oxytrichidae by BORROR (1972a). Trache-
lochaela sphagni on p 256 ofSRAMEK-HUSEK (1954) is probably a different (invalid) name for the type species.
Ventral and dorsal ciliature not known in detail; thus, Trachelochaeta is not considered in the present book.

Trachelostyla BORROR, 1972,1. Protozool., 19: 15 - Type (original designation): Slichochaela pediculi/ormis
COHN, 1866. Remarks: Classified in the Oxytrichidae by BORROR (1972a). Trachelostyla was originally estab-
lished by KARL (1932, Tierwelt Dtl., 25: 596) with two species. Since he did not determine any as type, the
taxon is invalid according to Article 13 (b) of the rCZN (1985); a fact overlooked by CORLISS (1979). I consider
BORROR (1972a) as the author of Trachelostyla (although he did not realize that KARL's taxon was invalid) be-
cause he fixed the type species. Slichochaela CLAPAREDE & LACHMANN, 1858 is not a synonym of Trache-
lostyla, as erroneously stated by CORLISS (1979, P 310), but of Slichotricha PERTY, 1849 (see BORROR 1972a,
P 12). Neither the ventral nor the dorsal ciliature of Trachelostyla pediculi/ormis are known in detail; thus,
Trachelostyla is not considered in the present book.

Tricholeptus FROMENTEL, 1875, Microzoaires, p 177 - Type (monotypy): Tricholeplus aculeatus FROMENTEL,
1875 (species description in FROMENTEL 1876, P 294). Remarks: Classified in the Oxytrichinae by JANKOWSKI
(1979). Nomen oblitum (see p 397). LoNGHI (1895, P 81) described the variety T. aculeatus telracu/eatus,
which is also a species indeterminata
Illustrations of Species Indeterminata and Insufficient
Redescriptions

The following plates show species indeterminata - species that cannot be identified from
the original description - and illustrations from redescriptions of valid species which
deviate significantly from the original description or the authoritative redescription.
Names are given as written in the original literature, that is, incorrect spellings or deviat-
ing combinations (for example, Kerona pustulata instead of Stylonychia pustulata) have
not been corrected.

Fig. 220a-m Insufficient redescriptions and species indeterminata (ventral view from life unless otherwise
indicated). a) Sty/onychia sp. (from WfEf & LUNDIN 1963), 200 ~m; p 605. b) Pleurotricha sp. (from WfEf &
LUNDIN 1963), 175 Jim; p 717. c) Drepaninafa/cata (from SAVI 1913), ? Jim; p 893. d) Kerona pustu/ata (after
CARTER 1859), size not indicated; p 600. e) Oxytrichajurcatus (after SMl1lI 1897b), ventral surface in dorsal
view, 140 Jim; p 248. 1) Sty/onychia sp. (from LEPSI 1965),90-100 Jim; p 605. g) Sty/onychia sp. (from Vux-
ANOVICI 1963), 120 Jim; p 605. h-j) Stylonychia sp. (from VUXANOVICI 1963), 130 Jim (h), conjugation pair (i),
arrangement of transverse cirri 0); p 605. k) Oxytrichafallax (from WF1)T & LUNDIN 1963), 150 Jim; p 605.1)
Oxytricha saprobia (from VUXANOVICI 1963), 140 Jim; p 259. m) Oxytricha aeruginosa (from PATSaI 1974),
ventral infraciliature after protargo) impregnation, 140 Jim; p 256. Figures on p 901.

Fig. 221a-i Insufficient redescriptions (ventral view from life unless otherwise indicated). a, b) Oxytricha
aeruginosa (from AGAMALlEV 1974), ventral and dorsal infraciliature, wet nitrate silver impregnation, 135 Jim;
p 256. c) Oxytrichafallax (after CONN 1905), 140 Jim; p 257. d) Oxytricha hymenostoma (after CONN 1905),
55 ~m; p 259. e) Oxytrichafallax (from SMl1lII978), 110 Jim; p 257.1) Oxytrichafallax (from BISHOP 1943),
size not indicated; p 257. g) Oxytricha /udibunda (from CURDS 1969), 100 Jim; p 259. h) Opisthotricha similis
(from CURDS 1969), 100 Jim; p 255. i) Oxytricha chlorigella (after HORVATH 1939b), ventral view after
opalblue stain after BRF1)sLAu, 140 Jim; p 256. Figures on p 902.
Fig. 221j Oxytricha hymenostoma (after ESPOSITO et aI. 1978). Ventral view after haematoxylin stain, size not
indicated; p 150. Figure on p 902.

Fig. 222a-t Insufficient redescriptions and species indeterminata (ventral view from life unless otherwise
indicated). a) Holosticha oxytrichoidea (from <JELEl 1950), ventral infraciliature, method and size not indi-
cated; p 254. b) Opistotricha terrico/a (from GELLERT 1957), ventral infraciliature, sublimate fixation and
opalblue stain after BRF1)sLAu, 70 Jim; p 245. c, d) Oxytricha sp. (from SF1)HACHAR & KAsTURl BAl 1963),
nuclear apparatus after Feulgen stain, sizes not indicated; p 245. e) Oxytricha sp. (after MOTE 1954), ventral
surface in dorsal view, stain (?), size not indicated; p 260. 1) Histrio acuminatus (after MAsKELL 1886),
140 ~m; p 666. g) Histrio complanatus (after LEPsI 1965), 70 Jim; p 667. h) Oxytricha ovalis (after SCHMARDA
1854), ventral and dorsal view from life, about 90 Jim; p 252. i) Oxytricha striata (after SCHMARDA 1854), ven-
tral, dorsal, and lateral views from life, about 145 ~m; p 253. j) Stylonchia mytilus (after CIENKOWSKY 1855, Z.
wiss. Zool., 6: 302), 260 Jim; p 517. k, I) Stylonychia pustulata (after CIENKOWSKY 1855), 123 Jim (k), cyst
from life, 45 Jim (I); p 603. m, n) Stylonychia lanceolata (after CIENKOWSKY 1855),275 Jim (m), cyst from life,
75 ~m (n); p 717. o-t) Oxytricha truncata (from VUXANOVICI 1963. o-t, from life). 0, p) Ventral views,
140 Jim. q, r) Ventral and lateral view of forma dilatata, 90 Jim. s, t) Ventral and lateral view of forma piri-
formis, 120 !-1m; p 254. Figures on p 903.
896 SYSTEMATIC SECTION

Fig. 223a-n, p Insufficient redescriptions and species indeterminata (ventral view from life unless otherwise
indicated). a) Pleurotricha grandis (after GOODEY 1911), 120 J.lm; p 716. b) Onychodromopsisjlexilis (from
LUNDIN & WEST 1963), ventral surface in dorsal view, size not indicated; p 479. c) Steinia inquieta (from DING-
FElDER 1962), 150 J.lm; p 324. d) Steinia candens (after CHARDEZ 1981), 135 J.lm; p 323. e-g) Oxytrichafer-
ruginea (after PENARD 1922), size not indicated (e), overall view and detail of cyst (t; g), size not indicated;
p 491. b) Stylonychia putrina (from BARWICK et al. 1955), 100 11m; p 604. i, j) Oxytricha (7) parvula (from
VUXANOVlCl 1963), ventral and lateral view from life, i = 40 J.lm; p 252. k) Oxytricha acuminata (from Vux-
ANOVlCl 1963), 100 J.lm; p 246. I) Oxytricha monstrosa (from VUXANOVlCl 1963), 85 11m; p 250. m, n) Oxytri-
chaformosa (from ALEKPEROV 1984a), ventral infraciliature after wet nitrate silver impregnation, size not indi-
cated (n), nuclear apparatus (m); p 248. p) Oxytrichafallax (from CHARDEZ 1967), 150 Jlm; p 258. Figures on
p904.

Fig. 224a-m Insufficient redescriptions and species indeterminata (ventral view from life unless otherwise
indicated). a, b) Oxytricha immemorata (from ALEKPEROV 1984a), ventral infraciliature after wet silver nitrate
impregnation, size not indicated (a), nuclear apparatus (b); p 249. c) Oxytricha jurcata (from DINGFELDER
1962), 110 11m; p 258. d) Oxytricha lata (after STERK! 1878), part of the fan-shaped adoral zone of mem-
branelles, size not indicated; p 250. e) Oxytricha oltenica (from LEPSI 1965), size not indicated; p 251. t) His-
trio complanatus (from LEPSI 1965), size not indicated; p 667. g) Opisthotricha sp. (after SUDZUKI 1964b), size
not indicated. Possibly identical with Sterkiella thompsoni (see there); p 679. b) Opisthotricha sordis (from
LEPSI 1965), size not indicated; p 245. i, j) Oxytricha sp. (after CONN 1905), i = 55 J.lm, j = 70 J.lm; p 260. k)
Opisthotricha procera (after NAIDU 1965), ventral surface in dorsal view from life, 70 J.lm; p 255. I) Stylony-
chia pustulata (after NAIDU 1965), 140 11m; p 604. m) Oxytricha ludibunda (after NAIDU 1965), dorsal view
from life, 85 11m; p 259. Figures on p 905.

Fig. 22Sa--u Insufficient redescriptions and species indeterminata (ventral view from life unless otherwise
indicated). a) Tachysoma parvistylum (after STOKES 1887b), 65 J.lm; p 466. b, c) Oxytricha parvistyla (after
CONN 1905), sizes not indicated; p 467. d) Tachysoma parvistyla (from LUNDIN & WEST 1963), size not indi-
cated; p 468. e) Tachysoma sp. (after UPSI 1957), dorsal view from life, 70 11m; p 469. t) Oxytricha agilis
(after CONN 1905), size not indicated; p 467. g) Oxytricha pel/ionella (after SAMANO & SOKOLOFF 1931),
70 11m; p468. b) Urosoma sp. (from LEPSI 1965), 190 11m; p 431. i) Urosoma cienlwwski (after CONN 1905),
size not indicated; p 431. j) Oxytricha pellionella (from LEPINIS et al. 1973), size not indicated; p 468. k) P/eu-
rotricha sp. (from Nll(QUUK & GELUER 1972), size not indicated; p 717. I) Oxytricha pel/ionella (after CONN
1905), size not indicated; p 468. m) Oxytricha pellionella (from SMITH 1978), 80 11m; p 467. n) Oxytricha pel-
Iionella (after QUENNERSTEDT 1865), size not indicated; p 467.0) Gonostomum sp. (after MOTE 1954), 80 11m;
p 395. p, q) Steiniafenestrata (after LEPSI 1957), ventral views after a "preserved object", sizes not indicated;
p 644. r) Oxytricha pellionella (after FELLERS & ALLISON 1920), 70-100 11m; p 468. s) Steinia platystoma
(after BIERNACKA 1959), 70 11m; p 645. t) Oxytricha pel/ionella (after BIERNACKA 1959), 106 11m; p 468. u)
Ciliate sp. (after MOTE 1954), size not indicated; p 367. Figures on p 906.

Fig. 226a-l Insufficient redescriptions (ventral view from life unless otherwise indicated). a, c) P/eurotricha
sp. (a, after CONN 1905; c, after CHAUDHURl 1929), a = 95 Jlm, c = 80 11m; p 716, 717. b, g) Pleurotricha gran-
dis (b, after KOFFMAN 1926; g, from SCHMALL 1976, protargo! impregnation), b = 100 /lffi, g = 115 11m; p 716.
d) Oxytricha marina (from BIERNACKA 1967), 120 J.lm; p 259. e) Opistotricha ovata (from BIERNACKA 1967),
100-120 Jlm; p 255. t) Oxytricha pel/ionella (from BRODSKY & YANKOWSKAYA 1929), size not indicated;
p 467. b) Stylonychia notophora (from EDMONDSON 1906), 120-160 11m; p 603. i) Oxytricha sp. (from JACOB-
SON 1931), 100 Jlm; p 260. j) Urosoma sp. (from LEPSI 1927), 146 11m; p 431. k) Gastrostyla sp. (from LEPSI
1927), 76 Jlm; p 824. I) Stylonichia mytilus (from RAMIREZ DE GUERRERO 1970), 150 11m; p 603. Figures on
p907.
 SPECIES INDETERMINATA 897

Fig. 227a-p Species indetenninata (from DUMAS 1929. a-p, from life, sizes not indicated). a) Kerona tremula;
p 841. b) Kerona caudata; p 834. c) Kerona diademata; p 840. d) Kerona truncata; p 841. e) Kerona
curvata; p 840. f, g) Kerona hecci/ormis; p 839. b) Kerona ohlonga; p 840. i) Kerona suhrotondata; p 841. j)
Kerona anfracta; p 839. k) Kerona ovata; p 840. I, m) Kerona urostyla, an insufficient redescription ofOxy-
tricha urostyla CLAPAREDE & LACHMANN; p 244, 873. n) Kerona mamillata; p 840. 0) Kerona silurus (con-
gyrans); p 839. p) Kerona arcuata; p 839. Figures on p 908.

Fig. 228a-i Insufficient redescriptions and species indetenninata (ventral view from life). a) Oxytricha
marina (from GANAPATI & RAo 1958), 175 11m; p259. b) Oxytricha hi/aria (from SOKOLOFF 1931), size not
indicated; p 601. c) Oxytricha pel/ionel/a (from BELTRAN 1928), size not indicated; p 467. d) Gonostomum
affine (from DELHEZ & CHARDFZ 1970), 75 11m; p 395. e) Stylonychia putrina (from CONN 1905), 80 11m;
p 604. t) Stylonychia pustulata (from CONN 1905), 125 11m; p 603. g) Oxytricha hi/aria (from CONN 1905),
185 11m; p 601. b) Gonostomum parvum (from LEPSI 1947),60-80 J1m; p 393. i) Oxytricha sp. (from CZAPIK
1959), size not indicated; p 260. Figures on p 909.

Fig. 229a-b Insufficient redescriptions and species indetenninata (ventral view from life unless otherwise
indicated). a) Stylonichia mim. 1 (from izQUIERDO 1906), 190 11m; p 599. b) Stylonychia notophora (from
CONN 1905), size not indicated; p 603. c) Stylonychia pustulata var. (from CONN 1905), size not indicated;
p 603. d) Histriculus polycirratus (from AGAMALIEV 1978), ventral infraciliature after wet silver nitrate impreg-
nation and nuclear apparatus after Feulgen stain, 80-90 11m; p 666. e) Oxitricha IlUm. 3 (from izQUIERDO
1906), 92 11m; p 245. t) Oxitricha mim. 2 (from izQUIERDO 1906), 140 11m; p 245. g) Oxitricha num. 1 (from
IzQUIERDO 1906), 130 11m; p 245. b) Oxytrichafallax (from HEBERER 1928), size not indicated; p 257. Figures
on p910.

Fig. 230a-b Insufficient redescriptions (ventral view from life unless otherwise indicated). a, b) Oxytricha
(after MAUPAS 1888), 170 11m (a), cyst, 40 11m (b); p 256. c, d) Stylonychia mytilus (from AGAMALIEV 1978),
ventral and dorsal infraciliature, wet silver nitrate impregnation, 90 11m; p 602. e) Stylonychia pustulata (from
LUNDIN & WFSr 1963), 150 11m (size from WEST & LUNDIN 1963); P 604. C, g) Stylonychia muscorum (from
MATIS & DANISKovA 1972), sizes not indicated; p 602. b) Oxytricha similis (from PATSCH 1974), ventral infra-
ciliature, protargol impregnation, 120 11m; p 260. Figures on p 911.

Fig. 231a-j Insufficient redescriptions (ventral view from life unless otherwise indicated). a) Pleurotricha
grandis (after RAo 1928), size not indicated; p 716. b, c) Gonostomum sp. (after CHAUDHURI 1929), dorsal and
ventral view from life, 50 11m, 46 11m; p 395. d) Oxytrichafallax (after KOFFMAN 1926), ventral view ofa fIxed
specimen, 125 11m; p 257. e) Stylonychia pustulata (after KOFFMAN 1926), ventral view of a fIxed specimen,
115 11m; p 603. t) Oxytricha hi/aria (from BANCHETIl & RICCI 1986), schematic drawing of the ventral surface,
100 11m; p 568. g) Stylonychia sp. (from ALONSO & PERFZ-SILVA 1963b), stain method?, 80 11m; p 604. b) Oxy-
tricha minor (from CHORIK 1968), 85 11m; p 259. i, j) Oxytricha hi/aria (after CoNN 1905), 110-132 11m;
p 601. Figures on p 912.

Fig. 232a-p Insufficient redescriptions and species indetenninata (ventral view from life unless otherwise
indicated). a) Histrio complanatus (after CONN 1905),70 11m; p 667. b) Histrio erethisticus (after CoNN 1905),
100 11m; p 667. c) Histrio sp. (after CONN 1905), 135 11m; p 667. d, e) Holosticha setigera (after CoNN 1905),
ventral and lateral view from life, 40 11m. Certainly not identical with Oxytricha setigera because midventral
cirri present; possibly a new species. t) Stylonychia histrio (after BORGER 1905),250 11m; p 668. g) Oxytricha
complanatus (from CHORIK 1968),80 11m; p 668. b) Stylonychia pustulata (after EcKHARD 1846), dorsal view
from life, size not indicated; p 603. i) Oxytrichafallax (after BARKER 1949), 85 11m (?); p 257. j, k) Tachy-
soma pellionella (from TCHANG et al. 1984), ventral and dorsal infraciliature, protargol impregnation, sizes not
indicated. (j) shows an early morphogenetic stage; p 469. I) Histriculus similis (from NIETO et aI. 1984), ventral
898 SYSTEMATIC SECTION

infraciliature (schematic), protargol impregnation, 160 11m; p 666. m) Oxytricha ha/ophila (from CZAPIK &
JORDAN 1976a), ventral infraciliature, wet silver nitrate impregnation (?), size not indicated; p 258. 0) Oxytri-
chajal/ax (from CELA 1972), ventral view from life (?), 62 11m (?); p 258.0) Oxytricha sa/tans (from VUXANO-
VICI 1963),45 11m; p 253. p) Oxytricha strenua (after KOFFMAN 1926), ventral (?) view from life, 55 11m; p 395.
Figures on p 913.

Fig.233a-m Insufficient redescriptions (ventral view from life unless otherwise indicated). a) Tachysoma pel-
lione//a (from ROSA 1957b), 80 11m; p 468. b) Gonostomum qffine (from ROSA 1957a), 60 11m; p 395. c)
Tachysoma pe//ione//a (from ROSA 1957a), 100 11m; p 469. d) Tachysoma pe//ione//a (from BUISAN 1944),
80-100 11m; p 467. e) H%sticha mystacea (from CHORIK 1968), 130 11m; p 824. f) Oxytricha bifaria (after
SMl1lII914), 200 11m; p 601. g) Oxytricha pe//ione//a (from SMl1lII914), 160 J.lm; p 468. h) Gastrosty/a steini
(from SCHNEIDER 1930), dorsal view from life after addition of iodine-ink, about 200 J.lm; p 824. i) Uroleplus
agi/is (after SMl1lI 1914), 160 J.lm; p 367. j) Sty/onichia muscorum (from CiiARDEZ 1967), 88 J.lm; p 601. k)
Sty/onychia putrina (from BURKOVSKY 1984), 200 J.lm; p 604. I, m) Oxytricha ha/ophila (from AGAMALIEV
1978), ventral and dorsal infraciliature after wet silver nitrate impregnation, I = 110 J.lm, m = 100 J.lm; p 258.
Figures on p 914.

Fig. 234a, b, d-I, n-p Insufficient redescriptions and species indeterminata (ventral view from life unless oth-
erwise indicated). a) Sty/onychia mytilus (from LUNDIN & WEST 1963), size not indicated; p 602. b) Oxytricha
ja//ax (from FORMISANO 1957), about 140 J.lm; p 258. d) Oxytricha acuminata (from MAlwAN 1977),
100-115 11m; p 246. e) Uro/eplus agilis (from EDMONDSON 1906),300 11m; p 367. f-h) Steinia inquieta (from
VUXANOVICI 1961), ventral view, cortical granules, lateral view, 120-160 J.lm; p 324. i) Oxytricha pellionella
(from EDMONDSON 1906),80-100 J.lm; p 468. j) Tachysoma pe//ionel/a (from SOKOLOFF & ANCONA 1937), size
not indicated; p 468. k) P/eurotrieha lanceo/ata (from EDMONDSON 1906),250 J.lm; p 716. I) Steinia eaudens
(from CiiARDEZ 1971), 145 J.lm; p 324.0) Tachysoma parvistyla (from CHARDEZ 1971),53 J.lm; p 468.0) P/eu-
rotricha sp. (from OUVEIRA et al. 1954),85 J.lm; p 717. p) Sty/onichia sp. (from OUVEIRA et al. 1954), 125 J.lm;
p 602. Figures on p 915.
Fig.234c Sty/onychia pustulata (from EDMONDSON 1906). Ventral view from life, 150-170 J.lm; p 565. Figure
on p 915.
Fig. 234m Sty/onychia curvata (from CHARDEZ 1971). Ventral view from life, 73 J.lm; p 593. Figure on p 915.

Fig. 235A, B Oxytrieha nova (from PRESCOIT 1983). A) Stained for DNA by the Feulgen procedure and coun-
terstained with fast green. Four micronuclei and 2 macronuclear nodules are present. Bar = 30 J.lm. This figure
was already published in PRESCOTT (1977). B) SCarIDing electron micrograph of the ventral side. Recent studies
by FOISSNER et al. (in preparation) strongly indicate that 0. nova is a species of the Sterkie//a histriomuscorum
group. Bar = 30 J.lm; p 250, 685. Figures on p 916.

Fig. 236a-r Insufficient redescriptions and species indeterminata (a-p, from DUMAS 1929; q, r, from DUMAS
1930. Ventral or dorsal views from life; sizes uncertain). a, b, r) Oxytrieha eautiata, 200 J.lm; p 430. c) Oxytri-
eha ovalis, 120 J.lm; p 248. d) Oxytrieha eornipes, 35 J.lm; p 247. e) Oxytrieha platystoma, 88 J.lm; p 645. f)
Oxytricha laerimu!a, 70 11m; p 249. g) Oxytricha laeerata, 115 11m; p 249. b) Oxytricha pistil/oides, 55 11m;
p 252. i) Oxytrieha subcy/indrica, 85 11m; p 253. j) Oxytrieha oblonga, 55 11m; p 251. k) Oxytricha curta,
57 11m; p 247. I) Oxytrieha becciformis, 52 11m; p 247. m) Oxytricha; p 256. 0) Oxytricha cornuta, 103 11m;
p 247. 0) Oxytricha pleuronectes, 70 11m; p 252. p, q) Oxytricha avata, (q) is reminiscent of O. setigera;
p 251. Figures on p 917.

Fig. 237a-j Insufficient redescriptions and species indeterminata (from DUMAS 1929. Ventral or dorsal views
from life; sizes uncertain). a) Stylonichia pustu/ata, ? J.lm; p 60l. b) Sty/onichia trochiformis, 112 J.lm; p 599.
c) Sty/onichia rastrata, 85 J.lm; p 599. d) Sty/aniehia virgu/a (in the legend designated as S. auriformis),
 SPECIES INDETERMINATA 899

102 11m; p 549. e) Stylonichia massula, 162 11m; p 598. f, i) Stylonichia mytilus, 175 11m, 142 11m; p 601. g)
Stylonichia dupla, 112 11m; p 598. h) Stylonichia regularis, 200 11m; p 600; j) Stylonichia e//ipsoides, ? 11m;
p 598. Figures on p 918.

Fig. 238a-r Species indeterminata und insufficient redescriptions (a-d, from DUMAS 1929; e-r, from DUMAS
1930. Ventral, dorsal, and lateral views from life; sizes uncertain). a) Stylonichia anfracta, 140 11m; p 597. b)
Stylonichia cornifrons, 155 11m; p 597. c) Stylonichia mamillata, 212 11m; p 598. d) Stylonichia cornicuJata;
p 597. e) Stylonichia globifrons; p 598. f-h) Stylonichia mutabunda; p 599. i, q) Stylonichia Iimbiformis;
p 598. j) Stylonichia rec/inis; p 599. k) Stylonichia labiata; p 598. I) Stylonichia excavata; p 598. m) Stylo-
nichia capucinus; p 597. n) Stylonichea bistyl/us; p 597. 0) Stylonichia truncata; p 599. p) Stylonichia gyrini-
jormis; p 598. r) Stylonichia unguiculus; p 599. Figures on p 919.

Fig. 239a-w Species indeterminata and insufficient redescriptions (a-o, r-t, from DUMAS 1930; p, q, u-w,
from DUMAS 1937. Ventral, dorsal, and lateral views from life; sizes uncertain). a) Oxytricha obtusa; p 251. b,
c) Oxytricha fimbriata; p 248. d, e) Oxytricha arcuata; p 246. f, n, u) Oxytricha praeceps; p 252. g) Oxytri-
cha caudata; p 430. h) Oxytricha stratiformis; p 253. i) Oxytricha longipes; p 250. j) Oxytricha labiata;
p 249. k) Oxytricha barbula; p 247. I) Oxytricha exociformis; p 248. m) Oxytricha rostrata; p 252. 0) Oxytri-
cha anca; p 246. p) Stylonichia viridis; p 599. q) Oxytricha luteolucens; p 250. r, s) Oxytricha proboscis;
p 252. t) Oxytricha acuminata; p 246. v) Stylonichia fronte-albens; p 598. w) Stylonichia membranaceus;
p 599. Figures on p 920.

Fig. 240a-n Species indeterminata and insufficient redescriptions (ventral view from life unless otherwise
indicated). a) Stylonichia auriformis (from DUMAS 1937); P 597. b) Stylonichiafrontosus (from DUMAS 1937);
p 598. e, d) Stylonichia bicaudatus (from DUMAS 1937); p 597. e) Stylonichia octonistylus (from DUMAS
1937); P 599. t) Stylonychia sp. (from NIKOUUK & GELTZER 1972), 96 11m; p 605. g) Oxytricha tetranucleata
(after GELEI 1950; from life?); p 253. h) Hypotrichen (nov. spec.?) (from FORTNER 1934),225-270 )lffi; p 597.
i) Opisthotricha paral/ela (from SCHUSTER 1899), dorsal view?, 250 11m; p 255. j) Stylonichia tricornis (from
MADRAZO-GARIBAY & li>PEZ-OCHOTERENA 1973), 128 11m; p 602. k) Abnormal reproduction in Stylonychia
(from GREENAWAY 1916), 200 11m; p 28. I, m) Oxytricha hengshanensis (from SHEN et al. 1992; I, silver
impregnation), 124-153 11m. Arrow marks postoral ventral cirri; p 249. 0) Oxytricha jal/ax (after GREGORY
1923; haematoxylin stain), 85 Ilm; p 257. Figures on p 921.

Fig. 241a-i Insufficient redescriptions (ventral view from life unless otherwise indicated). a, b) Oxytricha
crassistilata (from ALEKPEROV 1993). Ventral and dorsal infraciliature after wet silver nitrate impregnation and
nuclear apparatus after Feulgen stain, 20 11m; p 257. c) Tachysoma sp. (from LEPSI 1965), 100-160 Ilm; p 469.
d) Tachysoma pe//ionel/a (from ALADRO LUBEL et al. 1990), 75 Ilm; p 469. e) Gonostomum strenuum (from
MADRAZO-GARIBAY & li>PEZ-OcHOTERENA 1973),170 x 35 Ilm; p 395. t) Pleurotricha num. 1 (from IzQUIERDO
1906),280 Ilm; p 716. g, h) Pleurotricha num. 2 (from IzQUIERDO 1906), g = 112 Ilm, h = 100-140 11m; p 716.
i) Sty[onychiaputrina (from SAMANO & SOKOLOFF 1931), 140 11m; p 604. Figures on p 922.

Fig. 242a-i Insufficient redescriptions (ventral view from life unless otherwise indicated). a, b) Oxytricha sp.
(after CHAUDHURI 1929), dorsal (7) and ventral (7) view, a = 90 Ilm, b = 80 Ilm; p 260. c) Stylonychia noto-
phora (after WANG 1925), II 0 Ilm; p 601. d) Oxytricha pel/ionel/a (after WANG 1925), 105 11m; p 468. e) Sty-
lonychia sp. (after WANG 1925), 150 11m; p 602. t) Stylonychia putrina (after WANG 1925),95 11m; p 602. g)
Historio sp. (after WANG 1925), 85 11m; p 666. h) Oxytricha bifaria (after WANG 1925), 70 11m; p 601. i) Stylo-
nychia pustulata (from CHEN 1944), ventral view after haematoxylin stain, 80 11m; p 603. j) Gastrostyla pul-
chra (from JONES 1974), 185 11m; p 824. Figures on p 923.
900 SYSTEMATIC SECTION

Fig. 243a-p Species indetenninata (a, b, q-u, from DUMAS 1929; c-h, from DUMAS 1930; i-p, from DUMAS
1937. a-p, from life, sizes not indicated). a) Kerona cylindrica; p 840. b) Kerona calvaeformis; p 839. c)
Kerona cunea/a; p 840. d) Kerona subr%nda/a; p 841. e) Kerona curva/a; p 840. f) Kerona extuberans;
p 840. g) Kerona ondulata; p 840. b) Kerona absidata; p 839. i) Kerona rostrata; p 840. j) Kerona bipes;
p 839. k) Kerona bicaudata; p 839. 1) Kerona anjracta; p 839. m) Kerona epalxijormis; p 840. n) Kerona
unguicularis; p 841. 0, p) Kerona mamillata; p 840. q) Oxytricha gyrinioides; p 249. r) Oxytricha ephip-
pioides; p 248. s) Oxytricha galeata; p 249. t) Oxytricha cypris; p 247. u) Oxytricha pisciunculijormis; p 252.
Figures on p 924.
 SPECIES INDETERMINATA 901

Fig.220a-m Legend on p 895.

 902 SYSTEMATIC SECTION

.Y

a . Fk;;:tA{:/·'·

Fig. 221a-j Legend on p 895.

 SPECIES INDETERMINATA 903

Fig.222a-t Legend on p 895.

 904 SYSTEMATIC SECTION

.......
. '.': "'<~i;-.(
. : .
... t.: ...•....,'
.:-.~

..... .

...
-.- .':'.:.'---~)

Fig. 223a-p Legend on p 896.

 SPECIES INDETERMINATA 905

Fig. 224a-m Legend on p 896.

 906 SYSTEMATIC SECTION

p q r
Fig. 225a-i Legend on p 896.
 SPECIES INDETERMINATA 907

~~: . .
~
.

. .'
",.- .

Fig. 226a-1 Legend on p 896.

908 SYSTEMATIC SECTION

Fig. 227a-p Legend on p 897.

 SPECIES INDETERMINATA 909

Fig. 228a-i Legend on p 897.

910 SYSTEMATIC SECTION

Fig. 229a-b Legend on p 897.

 SPECIES INDETERMINATA 911

Fig. 230a-h Legend on p 897.

912 SYSTEMATIC SECTION

. 23la-j Legend on p 897.

FIg.
 SPECIES INDETERMINATA 913

.
~ ...
··.
...•.
~

! •

..••'() ••...... I
•\.

Fig. 232a-p Legend on P 897.

914 SYSTEMATIC SECTION

Fig. 233a-m Legend on p 898.

 SPECIES INDETERMINATA 915

h
Fig. 234a-p Legend on P 898.
916 SYSTEMATIC SECTION

MACRONUCLEUS

. . . . t _ MICRONUCLEUS

Fig. 235A, B Legend on p 898.

 SPECIES INDETERMINATA 917

Fig. 236a-r Legend on p 898.

918 SYSTEMATIC SECTION

Fig.237a-j Legend on P 899.

 SPECIES INDETERMINATA 919

Fig. 238a-r Legend on p 899.

920 SYSTEMATIC SECTION

Fig. 239a-w Legend on p 899.

 SPECIES INDETERMINATA 921

Fig. 240a-n Legend on p 899.

922 SYSTEMATIC SECTION

.• .
;
~
~
,:"":.?-"

:.~
"
~
~

~
.<::
.. ,.'.Y.
.J~::
~ 'j

.}
~
;.. ~~:
I ~

·.;;f
~
,
~

../·f
;
~

t.r. ·~
. ·. '. ~. . . -. '. ~
\U,
...~.-:
ri.··
.. .
9

Fig. 241a-i Legend on p 899.

 SPECIES INDETERMINATA 923

9
e

Fig.242a-i Legend on p 899.

924 SYSTEMATIC SECTION

F·Ig. 243a-p Legend on p 900.

Addenda
The data by EIGNER (1997), PETZ & ForSSNER (1997a), and SONG & WILBERT (1997a) be-
came available too late for inclusion in the main text and thus are added here at the end of
the book.

1. Oxytricha opisthomuscorum FOISSNER, BLATIERER, BERGER & KOH-

MANN, 1991 (see p 217; Fig. 244a-j, Table 52)

Rem ark s: PETZ & ForSSNER (1997a, Polar Record, 33: 318) and PETZ & FOISSNER
(1998) found this species in terrestrial habitats of Wilkes Land, Antarctica. Their popula-
tion matches the brief descriptions based on living specimens given by KAHL (1932; Fig.
73a) and ForSSNER (1980a; Fig. 73b) in size, position of buccal cirrus, shape of peristo-
miallip, and habitat, but the marked contractility noted by ForSSNER (1980a) was not ob-
served. However, Antarctic specimens soon became rounded and slowed down when

•
--•,
# ~

#
,
,
,
"
f

I-I t .
a b c
Fig.244a--(: Oxytricha opisthomuscorum (from PElZ & FOISSNER 1997a a, b, from life; c, protargol impregna-
tion). a, b) Ventral and right lateral view, a = 70 ~m. c) Ventral infraciliature, 63 ~m. Large arrow marks
slightly postorally located posterior-most frontoventral cirrus (cirrus IV/3), small arrow denotes posterior-most
postoral ventral cirrus. M = hook-shaped anterior margin of buccal cavity; p 217, 925.
 926 ADDENDA

4
..
; ;

,.. 9

Fig. 244d-j Oxytricha opisthomuscorum (from PETz & FOissNER 1997a d-j, protargol impregnation). d) Dor-
sal infraciliature of same specimen as shown in Figure 244c. e--g) Dorsal views showing more or less distinct
connections between dorsal kineties 3 (arrow) and 4 (arrowheads). b-j) Variability of body shape and nuclear
apparatus, especially number and position of micronucleus. Ma = macronuclear nodule, Mi = micronucleus,
1-6 = dorsal kineties (5 and 6 are very likely dorsomarginal kineties); p 217, 925.
 ADDENDA 927

Table 52 Morphometric data of Oxytricha opisthomuscorum (opi) and Sterkiella histriomuscorum (his). All
data (from PETZ & FOISSNER 1997a) are based on protargol-impregnated specimens from raw cultures. All
measurements in micrometres. CV = coefficient of variation (in %), Max = maximum value, mean = arithme-
tic mean, Min = minimum value, n = sample size, SD = standard deviation

Character Species mean SD CV Min Max n

Body, length opi 60.8 5.3 8.6 49 74 31
his 83.7 8.4 10.1 66 102 31
Body, width opi 26.6 3.1 11.7 21 38 31
his 44.3 6.0 13.6 35 57 31
Anterior macronuclear nodule, length opi 12.8 1.8 14.1 9 17 30
his 20.1 3.1 15.3 14 29 31
Anterior macronuclear nodule, width opi 6.9 1.1 15.6 5 9 30
his 10.4 1.5 14.3 8 15 31
Micronuclei, number opi 1.0 1 2 32
his 1.8 1 2 31
Micronucleus, length opi 3.8 0.4 11.1 2.5 4.5 30
his 3.2 0.5 14.3 2.5 4.5 30
Adoral zone of membranelles, length opi 22.5 1.5 6.7 20 26 31
his 37.4 3.5 9.4 31 45 31
Adoral membraneIles, number opi 20.3 0.9 4.5 18 22 33
his 27.9 1.1 4.0 26 31 31
Paroral, length opi 9.7 1.0 10.0 7 12 32
his 20.7 1.8 8.7 17 25 31
Endoral, length opi 9.4 0.8 8.1 8 11 32
his 19.6 2.1 10.7 14 23 31
Apex to buccal cirrus, distance opi 9.9 1.1 11.1 8 12.5 31
Apex to paroral, distance opi 8.3 0.9 11.4 6.5 10.5 31
Frontoventral cirri, number opi 4.1 4 5 29
his 4.0 4 4 31
Postoral ventral cirri, number opi 3.3 0.7 22.5 2 5 31
his 3.0 0.3 8.6 2 4 31
Transverse cirri, number opi 5.1 5 6 31
his 5.0 5 5 31
Right marginal row, number of cirri opi 12.1 1.0 8.5 11 16 26
his 20.5 1.1 5.3 18 22 31
Left marginal row, number of cirri opi 13.1 1.0 7.6 11 15 23
his 18.1 1.3 7.1 15 21 31

transferred from the refrigerator (5°C) to the microscope (beginning encystment?). PETz
& FOISSNER (l997a) provided the following improved diagnosis: Size in life about
55-80 x 20-40 Jim. hlVariably (n = 31) 2 macronuclear nodules and one micronucleus in
between. Buccal cirrus near anterior end of paroral. 20 adoral membranelles, 12 right
marginal cirri, 13 left marginal cirri, 5 transverse cirri, and 3 caudal cirri on average. Six
dorsal kineties with about 10 Jim long cilia.
Redescription by PETZ & FOISSNER (l997a; slides are deposited in the OberOsterrei-
chische Landesmuseum in Linz, Upper Austria): In life 65-70 x 25-30 Jim, outline ellip-
928 ADDENDA

soidal to fusifonn because ends often narrowly rounded or bluntly pointed (Fig. 244a,
h-j), dorsoventrally flattened up to 2: 1 (Fig. 244b). Flexible, especially in anterior por-
tion. Macronuclear nodules ellipsoidal to almost globular, in life about 16 x 10 ~m, with
nuclei 0.S-2.0 ~m across. Micronucleus globular, in life about 4-S ~m in diameter, in
71 % of specimens (n = 31) between macronuclear nodules or slightly displaced laterally,
in 16 %, respectively, 13 % adjacent to anterior or posterior nodule; one specimen had
two micronuclei (Fig. 244a, d, h-j). Contractile vacuole near left margin in or slightly be-
hind mid-body, with inconspicuous collecting canals. No cortical granules. Cytoplasm
colourless but specimens rather dark at low magnification due to bright fat globules
3--4 ~m long, and food vacuoles up to 10 ~m across containing bright and dark green al-
gae and heterotrophic flagellates. Crawls moderately fast in straight line interrupted by
short backward jerks; rotates about main body axis when swimming.
Adoral zone of membranelies extends over about 37 % of body length; longest bases
about 6-7 ~m in life, cilia IS-23 ~m long. Buccal cavity inconspicuous, narrow and flat,
anterior margin curved hook-like in live cells, right margin covered by inconspicuous
hyaline lip (Fig. 244a). Undulating membranes almost of same length,· slightly curved,
double-rowed paroral optically intersects apparently single-rowed endoral (Fig. 244c);
paroral cilia 3--4 ~m long. Pharyngeal fibres about 22 ~m in life. Invariably (n = 31) 3
frontal cirri, 1 buccal cirrus, and 2 pretransverse ventral cirri. Frontal cirri each consisting
of3--4, transverse cirri of up to S, and other cirral bases of3 basal body rows. Buccal cir-
rus near anterior end of paroral. Posterior-most frontoventral cirrus usually slightly be-
hind buccal vertex; 3, rarely 4 or S (19 %, n = 31) postoral ventral cirri in mid-body and
2 pretransverse ventral cirri in usual position; transverse cirri distinctly enlarged,
20--28 ~m long in life and usually motionlessly trailing behind. Marginal cirral rows al-
most confluent posteriorly, cirri in life about 16 ~m long, slightly elongated in rear por-
tion of cell, bases composed of two basal body rows each. Invariably (n = 2S) 6 dorsal ki-
neties, kinety pattern as in other oxytrichids, that is, kineties 1 and 2 nearly as long as
body; row 3 slightly to distinctly curved posteriorly and often indistinctly separated from
its offspring, the very short kinety 4, which bears the rightmost caudal cirrus; row S ex-
tends from anterior end to mid-body; row 6 consists of2--4 cilia only (Fig. 244d-g). Dor-
sal cilia 8-12 ~m long and stiffly spread in life, do not beat like nonnal cilia but become
soft and fleXIble under cover-slip pressure. Invariably (n = 26) 3 caudal cirri, fine, about
20-24 ~m long, associated with dorsal kineties 1,2, and 4 (Fig. 244d, g).
PETZ & FOISSNER (1997a) found Oxytricha opisthomuscorum in moss from Casey
Station, Wilkes Land, Antarctica. They counted IS2 active cells per gram dry mass of
soil, comprising about 9 % of the ciliate community. Edaphic parameters: soil tempera-
ture 2.8-13.7 °C, water content 6S-87 % of wet mass, loss of ignition 16.6-34.7 % of dry
mass, pH S.2-S.4. Grew in the laboratory at room temperature (about 19°C) and in dried
and moistened samples at SoC. Biomass of 106 medium-sized specimens about 18 mg.
Recently, I recognised that O. opisthomuscorum is possibly a synonym of (or at least
very closely related with) 0. balladyna (p 126), as suggested by the redescriptions by
Song & Wilbert (1989, Fig. 33b) and Petz & Foissner (1997a, Fig. 244c).
 ADDENDA 929

2. Sterkiella histriomuscorum (FOISSNER, BLATIERER, BERGER & KOH-

MANN, 1991) FOISSNER, BLATIERER, BERGER & KOHMANN, 1991 (see p 683;
Table 52)

Rem ark s: PETZ & FOISSNER (1997a, Polar Record, 33: 323) found this common spe-
cies in a fell field, south-west of Casey Station, Wilkes Land, Antarctica. They provided
the following additional observations (Table 52): In life 70-110 x 40-60 ~m, that is,
slightly smaller than other well studied populations. Marginal cirri about 15 ~m long,
frontal cirri about 17 ~m, transverse and caudal cirri 20-24 ~m, and dorsal cilia about
3 ~m. Adoral zone of membranelles extends over about 45 % of body length, longest
bases 7-9 ~m; cilia 17-20 ~m long. Biomass of 106 medium-sized specimens about
60 J.Ull. Several-days-old resting cysts spherical and entirely filled by cel~ in life
33-44 ~m (mean = 38.9 ~m, n = 11) in diameter; wall about 2.5/lm thick, surface
smooth or slightly uneven. Macronuclear nodules fused; however, when cyst wall is rup-
tured two (disintegrated?) nodules sometimes stain with methyl green-pyronin. Cyto-
plasm contains numerous granules and some globules up to 4 ~m across.
PETz & FOISSNER (1997a) also studied the morphogenesis (see this paper for details
and illustrations). They found some differences to the population described by BERGER et
al. (1985a), indicating that populations with 4 transverse cirri (BERGER et al. 1985a) are
not conspecific with those having 5 transverse cirri, as in the type population described
by KAHL (1932, Fig. 184k). Note that Sterkiella histriomuscorum is a sibling species
complex (FOISSNER & BERGER 1999).

3. Classification of the Oxytrichidae by EIGNER (1997)

EIGNER (1997) analysed the detailed descriptions of divisional morphogenesis of 40 hy-

potrichous species to establish a natural classification. He made schematised drawings
which show the interphase and its morphogenesis in one image. EIGNER'S classification
differs significantly from my classification, simply because he used other characters.
Here I discuss only some doubtful points of this most recent attempt to classify the hy-
potrichs.
• EIGNER (1997, P 553) stated that his results are in accordance with those obtained
from molecular techniques (SCHLEGEL & STEINBROCK 1986, SCHLEGEL et al. 1991).
However, Abb. 11 in SCHLEGEL & STEINBROCK (1986) shows that Stylonychia mytilus
is more closely related to S. pustulata than to Oxytricha granulifera and not vice
versa as suggested by EIGNER, who classified Stylonychia mytilus and Oxytricha
granulifera in the Oxytrichidae and Stylonychia pustulata (as Clara pustulata) in the
Parakahliellidae (see EIGNER'S Fig. 3). The dendrogram proposed by SCHLEGEL &
STEINBROCK (1986) matches our (BERGER & FOISSNER 1997) separation of the Oxytri-
chidae in the Stylonychinae and Oxytrichinae (note that Oxytricha nova, 0. bifaria,
930 ADDENDA

and 0. Ja/lax in SCHLEGEL & S1EINBROCK 1986 are in fact Sty/onychia or Sterkiella
species!). Figure 2 in SCHLEGEL et al. (1991) matches both our (BERGER & FOISSNER
1997) and EIGNER'S classification.
• EIGNER (1997, p 553) assumed that the conspicuous 18 frontal-ventral-transverse cir-
ral pattern (see my Fig. 6a) originated from at least two phylogenetic lineages. Ac-
cording to him, this pattern is the result of transformation series from little evolved
Oxytrichidae and Parakahliellidae to their highly evolved state, and this pattern is
therefore not an autapomorphy of a monophyletic group. Instead, he used differences
in the neokinetal anlagen development to separate the Oxytrichidae from the
Parakahliellidae. This hypothesis is the main reason why EIGNER'S classification is
very different from my proposal. However, I am rather certain that such a highly
characteristic cirral pattern (Fig. 6a), including many morphogenetic details, is not
the product of convergent evolution. Furthermore, EIGNER (1997, p 570) stated that
fragmentation of a dorsal kinety cannot be used to characterised the most highly
evolved hypotrich group (= oxytrichids). Conversely, fragmentation of a dorsal ki-
nety (primarily kinety 3) is considered as the sole autapomorphy of the last common
ancestor of all species treated in my book. Certainly, this character was modified
several times (for example, multiple fragmentation in kineties 1-3 or loss of frag-
mentation) during the origin of the oxytrichids 300-400 million years ago (WRIGHT
& LUIN 1997). But, as in the case of the 18-cirri pattern, I am certain that this curi-
ous dorsal pattern did not evolve convergently. In addition, I do not exclude that
groups like the amphisiellids or kahliellids also belong to the oxytrichids, that is,
they possibly evolved from oxytrichids in that they lost the fragmentation of dorsal
kineties and changed the ventral ciliature more (for example, Kahliella) or less (for
example, Lamtostyla) distinctly.
The cladogram proposed by EIGNER (1997, his Fig. 2) is not constructed according to
the original cladistic method and, is thus somewhat confusing, showing several con-
vergences. (i) EIGNER used the Holostichidae as outgroup. Many species of this group
have transverse cirri indicating that the presence of transverse cirri is a plesiomorphy
at the base of EIGNER'S cladogram. Consequently, the loss of transverse cirri in the
Orthoamphisiellidae would be an apomorphyand not a plesiomorphy (EIGNER'S char-
acter 6). Furthermore, according to EIGNER'S cladogram this character evolved con-
vergently in the Oxytrichidae and Parakahliellidae because it is applied at the wrong
level (it can be used to separate the Orthoamphisiellidae from the Oxytrichidael
Parakahliellidae group). (ii) In the holostichids, the outgroup used by EIGNER, the
rightmost cirral row (= migratory cirri and right transverse cirrus and its correspond-
ing pretransverse ventral cirrus) is not involved in primordia formation (for example,
Holosticha similis in HEMBERGER 1982, P lOS). Thus, the lack of an anlage in the
rightmost ventral cirral row is very likely a synapomorphy at the base of EIGNER'S
cladogram. Consequently, the lack of an anlage in this row (character I in EIGNER'S
cladogram) is not an apomorphy but a plesiomorphy.
 ADDENDA 931

• According to EIGNER (1997, his Fig. 2), the lack of old (parental) cilia and cirri in the
interphase cells of both the Oxytrichidae and Orthoamphisiellidae is an apomorphy,
that is, a further convergence. In spite of this, he assigned Kahliella simplex, a spe-
cies where many parental marginal cirri are retained, to the Oxytrichidae and Engel-
manniella mobilis, which also has many parental marginal cirri, to the Orthoam-
phisiellidae. Thus, the Kahliellidae became a junior synonym of the Oxytrichidae
and EIGNER had to establish a new (superfluous?) taxon, the Parakahliellidae.
• According to Figure 32 in EIGNER (1997, P 567), the buccal cirrus (cirrus 11/2) of
Coniculostomum monilata originates from the undulating membrane primordium. I
do not understand this misinterpretation of the original data (KAMRA & SAPRA 1990).
Presumably EIGNER confuSed the left frontal cirrus (cirrus Ill) and the buccal cirrus.
• According to Figure 41 in EIGNER (1997, P 571), the pretransverse ventral cirrus VI2
of Oxytricha granulifera is involved in primordia formation (indicated by a black
square = symbol h in his Fig. 1). However, this is certainly incorrect because this cir-
rus is not involved in primordia formation in Oxytricha granulifera or other species.
This cirrus is even present in an early postdivider (see Fig. 66a, d, f).
• According to Figure 42 in EIGNER (1997, P 571), Urosoma macrostyla has 3 pre-
transverse ventral cirri, which is due to a misinterpretation of the original illustra-
tions. This species has, as is usual in oxytrichids, only 2 pretransverse ventral cirri
(see p 411).

4. Oxytricha saltans (COHN, 1866) REES, 1881 (see p 236, Fig. 245a-e)

1997 Oxytricha saltans (ColIN, 1866) - SONG & WILBERT, Arch. Protistenk., 148: 420 (Fig. 24Sa-e; 1 nootype
slide of protargol-impregnated specimens is deposited in the ProtozooIogicaI Laboratory, College of
Fisheries, Ocean University of Qingdao, Qingdao, China).

Rem ark s: Oxytricha saltans (CoHN, 1866) REEs, 1881 is a secondary homonym of
Oxytricha saltans VUXANOVICI, 1963, a species determinata (see p 253). Thus, I do not re-
place the name OfVUXANOVICI'S species. SONG & WILBERT (1997a) found O. saltans many
times in marine fish- and mollusc-farming ponds in Qingdao, China (morphological stud-
ies mainly based on a population collected on October 27, 1993). They provided the fol-
lowing additional description: Cell size varies considerably depending on diet or different
populations, however, body shape almost constant, generally as shown in Figure 245a
with inconspicuous tapering anterior end. Pellicle flexible, no cortical granules. Cyto-
plasm colourless to slightly greyish, often with a few large food vacuoles filled with
green flagellates in specimens newly collected or containing many to numerous small
(2-4 Jlm) shining globes mostly within posterior half of cells when cultured (fed with
bacteria). Movement quite unique, usually very fast crawling (even jump-like) on bottom
of Petri dish with frequent jerks. When resting, more or less thigmotactic, adhering on
932 ADDENDA

substrate with the aid of

transverse cirri. When
swimming (only occa-
sionally) always slowly
rotating about main
body axis. Adoral zone
of membranelles about
33 % of body length,
generally bipartite by
3-5 flm wide gap in an-
terior third, 5 (seldom 4)
stiff membranelles (over
20 flm in length) form-
b
ing a crown-shaped api-
50pm
cal style radiating from
anterior border of cell
a (Fig. 245a) in anterior
,, ~~ ~ portion, others in poste-
,§
,s

.....
.t).. rior with last 3 ones

,,
1/ # '{00' :;.
close together, often
forming another separate
•••
I

-0 ,
., , It .- -OK
group. Undulating mem-
branes short and straight,

:~.~;..
It

, • ,
Ma-
parallel, almost equally
.
-0 ,, ,
. long. Buccal field very
Mi-
narrow (Fig. 245a, b);
, f buccal lip covers almost

-
If,
, .1 entire proximal part of
, •, e
adoral zone of mem-
branelles, with conspicu-
Fig. 245a-e Oxytricha sa/tans (from SONG & WILBERT 1997a a, b, from ous, flexible, spur-like
life; c-e, protargol impregnation). a) Ventral view. Note the very narrow protrusion (Fig. 245b,
buccal field. b) Ventral view of anterior portion showing spur-like (thorn- arrow) of unknown func-
shaped) protrusion (arrow) of the buccal lip. c) Left lateral view showing
tion, overlooked by pre-
nuclear apparatus. d, e) Ventral and dorsal infraciliature. Arrow in (d) de-
notes distinctly posteriorly displaced anterior end of right marginal row. vious authors. Cirral pat-
The arrangement of the frontal and frontoventral cirri is not as in a tern very particular: in-
"typical" Oxytricha species. Explanation of original labels DK = dorsal ki- variably 7 frontal cirri,
neties, Ma = macronuclear nodule, Mi = micronucleus; p 236, 931. conspicuously shifted
posteriorly with only
about 4 within frontal
field; cirri generally fme except transverse ones, which are 20-30 flm long; right mar-
ginal row begins behind mid-body (Fig. 245d, arrow). Usually 5 (rarely 6) dorsal
 ADDENDA 933

kineties, leftmost row distinctly shortened posteriorly or with large gap (Fig. 245e). Dor-
sal cilia 8-10 ~m long, conspicuous in life (Fig. 245a).

5. Hemigastrostyia SONG & WILBERT, 1997

1997 Hemigastrostyla SONG & WILBERT, Arch. Protistenk., 148: 421- Type (original designation): Oxytricha
(Urosoma) stenocephala BORROR, 1963.

C h a r act e r i sat ion: Marine or brackish water Oxytrichidae with slightly cephal-
ised body shape; mostly 8-10 frontal, 5 ventral and a few extra lateroventral cirri which
possibly derive from the 6th (rightmost) frontal-ventral-transverse cirral anlage during
morphogenesis; caudal cirri present.
Rem ark s: I did not change the characterisation proposed by SONG & Wll.BERT
(l997a). Unfortunately, they did notfix Oxytricha enigmatica (which they redescribed in
detail) as type species, but Oxytricha stenocephala (now Hemigastrostyla stenocephala
(BORROR, 1963) SONG & WILBERT, 1997), an incompletely known species previously clas-
sified in Gastrostyla (see p 822). Hemigastrostyla has two apomorphies, namely the ce-
phalised anterior end and the additional cirri right of the transverse cirri ("LVC" in Fig.
246e). According to SONG & WILBERT (l997a), an additional frontal-ventral-transverse
cirri primordium (that is, in total 7 against 6 in other 18-cirri oxytrichids) is a further apo-
morphy. However, this assumption is obviously based only on two specimens in physio-
logical reorganisation (Fig. 246i, j). Thus, divisional morphogenesis has to be studied to
clarify the origin of the curious cirri right of the transverse cirri. Oxytricha discifera also
has a cephalised anterior end (p 227, Fig. 81); however, KAlIL (1932) did not describe the
cirri right of the transverse cirri; thus, redescription is needed for fmal classification of
this species, which also lives in the sea.

Hemigastrostyia enigmatica (DRAGESCO & DRAGESCO-KERNEIS, 1986)

SONG & WILBERT, 1977 (see p 229, Fig. 246a-j, Table 53)

1997 Hemigastrostyla enigmatica (DRAGESCO & DRAGESco-KERNEIS, 1986) - SONG & WILBERT, Arch. Protis-
tenk., 148: 421 (Fig. 246a-j, Table 53)

Rem ark s: SONG & WILBERT (1997a) collected H. enigmatica twice in a marine
mollusc-farming water in Qingdao, China. The redescription clearly shows that Hemigas-
trostyla enigmatica is an 18-cirri oxytrichid with a somewhat unusual arrangement of the
frontal and frontoventral cirri, possibly due to the cephalisation. SONG & WILBERT (1997a)
provided a detailed redescription, which is given word for word (see also Table 53):
"Cell shape elongate with both ends widely rounded, frontal portion more or less head-
like; left margin only slightly convex while portion posterior to distal end of adoral zone
of membranelles on right evidently convex (Fig. 246a). When viewed ventrally, one in-
934 ADDENDA

Fig.246a-e Hemigastrostyla enigmatica (from SONG & WIlBERT 1997a lHI, from life; e, protargol impregna-
tion). a) Ventral view of a representative specimen, 105 Ilm. Note the cephalised anterior end. b) Detail ofleft
and right marginal groove. c) Detail of cortical granulation. Arrowheads mark cortical granules, <I Ilm across.
d) Detail of dorsal side showing the dorsal cilia within the cup-shaped depression. e) Ventral infraciliature,
112 Ilm. Tiny arrows denote the 4 frontoventral cirri, arrowhead marks the anterior-most postoral ventral
cirrus. Explanation of original labels: Be = buccal cirrus, ee = caudal cirri, Lve = curious cirri right of
transverse cirri (an apomorphy of Hemigastrostyla), ve = pretransverse ventral cirri; p 229, 933.

conspicuous indentation (marginal groove) formed by right marginal row at about ante-
rior 2/5 of cell length. Dorsoventrally significant flattened (about 1:3-4). Dorsal side un-
even, with central part thicker than posterior and anterior ends, right and left often form-
ing a projection-like border (Fig. 246g, h). Pellicle as shown in Fig. 246c, cortical gran-
ules very fine «1 ~m), generally grouped rosette-like and close to cilia (Fig. 246c, ar-
rowheads). At high magnification (1250 x ), always numerous small (ca. 2 ~m in size)
 ADDENDA 935

vacuole-like, ellipsoid glob-

ules at deeper level (beneath
\ \ cortical granules) recognis-
\ \
\ \ .I able (Fig. 246c). Cytoplasm
hyaline and colourless; cells
DKS-- - \- )-~ / frequently with many lipid
\ ) I.L -DKF droplets, 5-8 Jim across,
)) ! II
\ ~ ! •\ Food vacuoles rarely to rec-
t ,.. / '\ ognise, contain mostly flag-
) ~ 'II
./ ellates and diatoms. Con-

'}
'}
•J
tractile vacuole not ob-
served Constant two large
macronuclear nodules, ellip-
! soid, located in about mid-
body and left to cell margin,
9 h containing many spherical
nucleoli; replication band
often visible in non-dividing
individuals (Fig. 246f). Mi-
cronuclei several to ''many'',
small and near macronu-
clear nodules (Table 53).
f Movement slow to
moderate crawling on bot-
Fig. 246C-b Hemigastrostyla enigmatica (from SoNG & WIlBERT tom of Petri dish. Some-
1997a ( protargol impregnation; g, h, from life). f) Dorsal infracilia-
ture and nuclear apparatus, 112 }1m. Morphogenetic data are needed to
times more or less thigmo-
"understand" this kinety pattern. g, b) Left lateral and dorsal view. Ex- tactic, adhering with the aid
planation of original labels DKI,~, s = dorsal kineties, DKF = dorsomar- of transverse cirri on sub-
ginal kinety; p 229, 933. strate. When swimming
slowly rotating around its
longitudinal axis. Character-
istically 2-3 (usually 2) right marginal cirri arising from border and slowly waving up
and down, while all other ciliary organelles stationary.
Oral area narrow, posterior portion of adoral zone of membranelles always half cov-
ered by cytoplasmic lip on right margin of buccal field (Fig. 246a). Adoral zone ofmem-
branelles about half of cell length with distal end of zone extending back far onto right
side (Fig. 246a, e). Paroral and endoral membranes slightly curved, almost in parallel to
each other (Fig. 246e).
Three evidently enlarged cirri in anterior cell portion: 2 frontal cirri at most apical
area of cell, one buccal cirrus at about middle way of frontal region, other 5 frontal cirri
always grouped together, near distal end of adoral zone. Ventral cirri in two groups as in
many other oxytrichids: 3 anterior ventral cirri located anterior to cytostome level and
936 ADDENDA

Fig. 246i, j Hemigastrostyla enigmatica (from SONG & Wn.BERT 1997a i, j, protargol impregnation). Physio-
logical reorganisation. Dashed lines in (j) show possible cirra1 differentiation. However, detailed morphoge-
netic data are needed to elucidate the ontogenetic origin of the conspicuous cirri right of the transverse cirri
("LVC" in Fig. 246e). Explanation of original labels: CA = cirra1 anlagen, DKF = dorsomarginal kinety, UM =
undulating membranes, UMA = primordium of undulating membranes, I, 11, VI = cirra1 anlagen (note that an-
lage I corresponds to anlage II of Fig. 6a, because SoNG & Wn.BERT 1997a did not count the anlage of the undu-
lating membrane; p 229,933.

close to frontal cirri; 2 posterior ventral cirri near to transverse ones. Except those 5 nor-
mal ventral cirri, always 2 extra lateroventral cirri (called "fimbriate caudal cirri" by BOR-
ROR) right to transverse cirri and posterior end of right marginal row (Fig. 246a). Trans-
verse cirri strong, about 20/lm long, located subterminally. Marginal rows extending
slightly spirally from upper-right to lower-left. Right row shortened anteriorly with its
posterior end extending near to rightmost transverse cirrus (Fig. 246e). Left row turning
onto dorsal side (at least on margin of cell) and almost confluent with caudal cirri posteri-
orly. All marginal cirri densely spaced, laying "firmly" on cell surface along marginal
grooves (Fig. 246a, b). Bases of all cirri associated with well impregnated fibres (Fig.
246e).
 ADDENDA 937

Constant 5 dorsal kineties, kinety 1 shortened posteriorly while dorsal kinety 4 ex-
tending anteriorly only about halfway of cell (Fig. 246t). Apart from these 5 rows, one
ciliary fragment anterior of right marginal cirral row consisting of several basal bodies.
Dorsal cilia about 3 J.1m long, very characteristically located within cup-shaped depres-
sions (Fig. 246d)."

Table 53 Morphometric data of Hemigastrostyla enigmatica (from SONG & WILBERT 1997a). All data are
based on protargol-impregnated specimens. All measurements in micrometres. CV = coefficient of variation
(in %), Max = maximum value, mean = arithmetic mean, Min = minimum value, n = sample size, SD = stan-
dard deviation

Character mean SD CV Min Max n

Body, length 131.2 17.7 13.5 108 165 20
Body, width 63.4 11.2 17.7 45 82 20
Adoral zone of membranelles, length 64.6 7.5 11.6 54 77 20
Adoral membranelles, number 42.4 4.2 9.9 34 50 20
Macronuclear nodules, number 2.0 0 0 2 2 20
Macronuclear nodules, length 21.5 4.2 19.5 16 30 20
Macronuclear nodules, width 15.8 2.0 12.9 13 18 20
Micronuclei, number 4.5 1.3 28.7 2 6 20
Cirri, number! 15.0 0 0 IS IS 20
Transverse cirri, number 5.0 0 0 5 5 20
Left marginal cirri, number 20.3 1.0 5.1 17 21 18
Right marginal cirri, number 16.7 2.0 11.6 13 19 18
Dorsal kineties, number 5.0 0 0 5 5 12
Caudal cirri, number 3.0 0 0 3 3 18

! Including frontal cirri, buccal cirrus, frontoventral cirri, postoral ventral cirri, pretransverse ventral cirri, and
cirri right of transverse cirri.
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Note: All references listed below (about 2500) and more than 2000 further references (including keywords)
concerning hypotrichs are available as database from the author.

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Systematic Index
The index contains all names of hypotrichous ciliates mentioned in the book, including vernacular names (for
example, amphisiellids); food organisms and other ciliates are usually not included. The index is two-sided,
that is, species appear both with the generic name first (for example, Stylonychia mytilus) and with the species
name (epithet) first (mytilus, Stylonychia). Valid (in my judgement) oxytrichid species and genera are in bold-
face italics print. Invalid taxa Gunior homonyms, synonyms, outdated combinations, incorrect spellings, spe-
cies indeterminata, nomina nuda) are given in italics. Homonyms (for example, Plagiotricha BORY and
Plagiotricha KENT) are not included separately, that is, author names are omitted. Parentheses, correctly used
to mark a subgenus (ICZN 1985, Article 6a), are also omitted because in most cases they have not been applied
correctly. Suprageneric taxa are given in normal type (for example, Amphisiellidae), valid oxytrichid supra-
generic taxa in boldface (for example, Stylonychinae). Boldface page number indicates the location of a
valid taxon in the systematic section, "T" indicates the location of the table with the morphometric characteri-
sation.

absidata, Kerona 839 aestuari~ Cyrtohymena 280, 281,308,317,629

acarus, Himantopus 514 aestuarii, Oxytricha Steinia candens 308
acarus, Kerona 514 aestuarii, Oxytricha candens 317
acarus, Trichoda 514, 566 affine, Gastrostyla 370
Actinotricha 80, 81, 115,236,239 affine, Gonostomum Oxytricha 370
Actinotricha hyalin 236 affine, Trachelostyla 370
Actinotricha hyalina 236, 241 affine, Gonostomum 9, 33, 59, 61, 88, 103, 106,
Actinotricha saltans 115,236,253,255 143, 368, 369, 382T, 384, 389, 391, 392, 393,
Actinotricha saltans, Oxytricha 236 394,395,449
Actinotricha saltans, Tachysoma 236 affinis, Euplotes 450
aculeatus tetraculeatus, Tricholeptus 894 affinis, Oxytricha 367, 368, 369, 373
aculeatus, Tricholeptus 397, 894 affinis, Plagiotricha Gonostomum 370
acuminaba, Laurentia 753 affinis, Stichochaeta 370
acuminata, Dipleurostyla 893 affinis, Gonostomum 373
acuminata, Laurentia 753, 756, 757, 761, 763, 764T afxfine, Gonostomum 379
acuminata, Laurentiel/a 25,753 agile, Tachysoma 434, 444
acuminata, Oxytricha 246, 414 agiliformis, Urosomoida 35, 36, 57, 64, 68, 158,
acuminata, Oxytricha Urosoma 414 159,170, 184, 186,345,347,349,352,354T,356
acuminata, Psilotricha 894 agili/ormis, Urosomoides 359
acuminata, Urosoma 4, 11, 396, 397, 401, 405T, agilis, Oxytricha 36,347,467
414,495 agilis, Tachysoma 434
acuminata, Urosoma Oxytricha 414 agilis, Uroleptus 345, 346, 347, 348, 367
acuminatus, Histriculus 666 agilis, Urosomoida 8, 11, 35, 64, 106, 224, 345,
acuminatus, Histrio 646, 666 347,352,354T,407,409,817
acuminatus, Onychodromus 753 agilis, Urpleptus 367
acuminatus, Oxytricha Histrio 666 agilis, Utroleptus 347
acuminatus, Onycchodromus 753 Alaster polyporum 825
acuta, Urosoma 404,407,409 Alastor 825
admirabilis, Histriculus 646, 677 Alastor polyporum 825, 826
admirabilis, SterkieUa 4,669,671,677 alba, Oxytricha 242
aeroginosa, Oxytricha 122 alfredi, Oxytricha 120,125, 151, 167
aeruginosa, Oxytricha 9, 120, 122, 166, 169, 209, alfredkah/~ Oxytricha 121,227
224,254,256,480,488 algicola, Gonostomum 370, 373
aeruginosa, Oxytricha Oxytricha 123 algicolum, Gonostomum 370
1050 SYSTEMATIC INDEX

Allotricha 17, 18,56,62,66,67,69,71, 73, 75, 76, aurijormis, Stylonychia 549

77, 80, 81, 105, 114, 117, 205, 261, 359, 476, auripunctata, Oxytricha 10, 121, 123, BIT, 194,
478,607,699,716,782,848 224,346,347,364,684
Allotricha antarctica 9, 33, 62, 64, 68, 106, 205, australis, Cyrtohymena 9, 279, 281, 291T, 294, 296
262, 266, 267T, 268, 476, 704 australis, Notohymena 9, 324, 325, 334, 339T, 340
Allotricha lanceolata, Pleurotricha 700 australis, Oxytricha 340
Allotricha mo/lis 9, 33, 261, 262, 267T, 268, 270, Australocirrus 114,470, 717
272,704,782,848 Australocirrus octonucleatus 470, 717, 718
Allotricha spp. 782 Australocirrus oscitans 17, 106, 113, 470, 474T,
Allotricha, Pleurotricha 261, 698 717
alpestris, Epistylis 84
ambigua, Oxitricha 116 Bakuella 843
ambigua, Oxytricha 246 balatonica, Tachysoma 433, 454, 456, 457
ambigua, Trichoda 116,246 balladina, Steinia 177
ambigua, Urosoma 404, 407,409 Balladinella 128
ambiguum, Spirostomum 116,246 Balladinopsis 80, 893
Amphisia 808 Balladinopsis nuda 893
Amphisiella 243 Balladinopsis sphaericus 738
Amphisiella dorsicirrata 803 Balladyna 128
Amphisiella marioni 790 Balladynajusijormis 128
Amphisiellidae 81, 103, 104,491,781,889,894 Balladyna similis 126, 128
amphisiellids 61,607,790,815,842,884,893,930 balladyna, Oxytricha 4, 119, 126, 131T, 177, 178,
anca, Oxytricha 246 183,457,659
Ancystropodiinae 80, 778 Balladynella 80, 128
Ancystropodium 1,79,80,81,103, 104, 163,777 Balladynopsis 893
Ancystropodium maupasi 4, 14, 105, 164, 568, 777, Balladynopsis nuda 893
778, 790 balladynula, Cyrtohymena 175
andoi, Gonostomum 370, 384 balladynula, Oxytricha 177, 178
anfracta, Kerona 839 balladynula, Oxytricha Steinia 177
anfracta, Stylonichia 597 balladynula, Steinia 175, 177
antarctica, Allotricha 9, 33, 62, 64, 68, 106, 205, balladynulla, Oxytricha Steinia 177
262,266, 267T, 268, 476, 704 barbula, Oxytricha 247
antarctica, Notohymena 9, 324, 325,334, 339T becciformis, Kerona 839
antarctica, Urosomoida 121,346,347, 354T, 365 becciformis, Oxytricha 247
aper, Kerona 839 bicaudata, Kerona 839
Apoamphisiella 104, 113,476, 777, 781, 843, 844, bicaudatus, Stylonichia 597
882,889 bicirrata, Oxytricha 457
Apoamphisiella hymenophora 781, 782, 786, 872 bicirratum, Tachysoma 4, 86, 87, 173, 433, 452,
Apoamphisiella tihanyiensis 11, 781, 782, 786T, 454,456,457
787, 788 bijaria, Diophrys 557
appendiculata, Diophrys 596 bijaria, Oxytricha 14, 26, 37, 250, 557, 565, 568,
appendiculata, Sty/onychia 596 571,578,579,581,584,601,929
arcuata, Kerona 839 bijaria, Oxytricha Oxytricha 557
arcuata, Oxytricha 246 bi/aria, Stylonychia 34, 64, 67, 68, 69, 75, 76, 77,
Aspidisca 87, 840 78, 502, 503, 534T, 553, 556,557,571,577,601,
Aspidisca cicada 254 602,685,731
Aspidisca Iynceus 737,839 bilobata, Oxytricha 247
Aspidisca turrita 723 bimarginata, Coniculostomum 607, 608, 618T, 620,
Aspidiscidae 79 621,624
auricularis, Oxytricha 243 bimembranata, Oxytricha 121, 138,196,221,683
aurijormis, Stylonichia 597 bipes, Kerona 839
 SYSTEMATIC INDEX 1051

bistyllus, Stylonichea 597 Cerona lanceolata 699

bivacuolata, Oxytricha 641 Cerona mytilus 504
bivacuolata, Steinia 625,641 Cerona pustulata 565
bryonicolum, Trachelostyla 370 charon, Euplotes 450
bryonicolum, Gonostomum 370, 375 chilensis, Oxytricha 452
bryophila, Trachelochaeta 894 chilensis, Oxytricha pellionella 451
buitkampi, Oxytricha 175,178 chilensis, Tachysoma 432, 451
buitkampi, Paraurostyla 843, 888 chloreHigera, Oxytricha 8, 86, 87, 119, 129, 256,
buitkampi, Pseudouroleptus 843, 888, 889 605
bulla, Oxitricha 116 chlorelligera, Oxytricha Oxytricha 129
bulla, Trichoda 116 chlorigella, Oxytricha 129, 256
chonotrichids 4
calva, Stylonychia 599 cicada, Aspidisca 254
calvaeformis, Kerona 839 cicada, Trichoda 254
calvitium, Kerona 700 cienckowski, Urosoma 401
calvitium, Trichoda 566, 700 cienkowekii, Urozoma 404
camerounensis, Holosticha 769 cienkowski, Urosoma 398, 431
canadensis, Trachelostyla 370 cienkowskii, Oxytricha Urosoma 400
candens aestuarii, Oxytricha 317 cienkowskii, Urosoma 396, 398, 400, 401, 403
candens aestuarii, Oxytricha Steinia 308 cienkowskii, Urozoma 396
candens depressa, Cyrtohymena 316, 317 cienkowsky, Urosoma 401
candens depressa, Steinia 314, 315, 316 cienkowskyi, Urosoma 398
cantiens, Cyrtohymena 281,308,314,317,629,631 ciliata, Kerona 839
candens, Oxytricha 314 Ciliate sp. 367
candens, Oxytricha Steinia 309 Ciliate sp. No.1 683
candens, Steinia 305, 307, 308, 309, 323 Ciliophora 79, 80, 81
capitata, Oxytricha 243 ciliophorum, Trachelostyla 370
capucinus, Stylonichia 597 ciliophorum, Gonostomum 370
carnea, Oxytrichajlava 243 cimex, Stylonychia 501
carnea, Pseudokeronopsis 243 cimex, Trichoda 501
caudata, Kerona 839 cinekowski, Urosoma 398
caudata, Oxytricha 396, 398, 401, 430 citrina, Cyrtohymena 9, 281, 291T, 293, 296, 314,
caudata, Oxytricha Urosoma 400 340,459
caudata, Trichotaxis 398 citrina, Steinia 293
caudata, Urosoma 11, 106, 396, 398, 405T, 411, Cladotricha 894
414,691 Cladotrichidae 893
caudata, Urosoma Oxytricha 398 Clara 502, 565, 571, 591
cauda tum, Urosoma 400 Clara pustulata 565, 929
caudatus, Paruroleptus 84 Clara vorax 591
caudatus, Pseudouroleptus 105, 888, 889 c1avata, Stylonychia 511
caudatus, Uroleptus 23 c1avi/ormis, Oxytricha Stylonychia 505
caudens, Oxytricha 309 c1avi/ormis, Stylonychia 505, 511, 517, 537
caudens, Steinia 324 coei, Urostyla 845, 846, 848, 855, 870
cavicola, Histriculus 669 collini, Podophrya 584
cavicola, Oxitricha 677 colpodids 885
cavicola, Oxytricha 668, 669 companatus, Histrio 658
cavicola, Sterkiella 4, 16, 18, 20, 22, 23, 34, 46, 57, complanatus, Histriculus 647, 656, 657
60,64,71,168, 169,207, 667, 668,669, 674T complanatus, Histrio 657, 667
Ceratidium 597 complanatus, Oxytricha Histriculus 668
Ceratidium cuneatum 597 complanatus, Oxytricha Histrio 658
Cerona histrio 647 complex, Paraurostyla weissei Ill, 843,844, 868T
1052 SYSTEMATIC INDEX

complex, Sterkiella histriomuscorum 668, 669 cypris, Kerona 540

complex, Stylonychia mytilus 1,2,6, 8, 14,35,39, cypris, Oxytricha 247
40,41,42,43, 71, 81, 85, 87, 88, 106, 143,248, cypris, Trichoda 514
403,450,502,503,504,549,550,551,554,555, Cyrtohymena 17,56,57,62,66,67,68,70,73,75,
562,568,577,588,592,598,599,600,601,602, 76, 77, 112, 178, 218, 234, 248, 249, 263, 279,
603,604,607,611,617,640,704,755,759,761 324,325,338,340,346,349,459,480,605,625,
complex, Tetrahymena pyriformis 510 643,645,646
congyrans, Kerona 839 Cyrtohymena aestuarii 280,281,308,317,629
congyrans, Kerona silurus 839 Cyrtohymena australis 9, 279, 281, 291T, 294, 296
Coniculostomum 1,2,4, 18,20,28, 56, 62, 67, 68, Cyrtohymena bal/adynula 175
70, 71, 73, 75, 76, 77, 78, 81, 104, 105, 502, 571, Cyrtohymena candem 281, 308, 314, 317, 629, 631
606,625,640,699,706,755,761,775 Cyrtohymena candens depressa 316, 317
Coniculostomum bimarginata 607, 608, 618T, 620, Cyrtohymena citrina 9, 281, 29lT, 293, 296, 314,
621,624 340,459
Coniculostomum indica 608,611, 618T, 619, 620 Cyrtohymena fenestrata 645
Coniculostomum monilata 1, 33, 46, 48, 49, 50, 51, Cyrtohymenaferruginea 487
64, 106, 606, 607,608, 618T,621,624, 706, 763, Cyrtohymena gracilis 4,9, 125,280,295,305
931 Cyrtohymenagranulata 280,281,295,306
Conicyciostomum 607 Cyrtohymena inquieta 280,281,294,309,313,317,
corniculata, Stylonichia 597 320,324,625,626,629
cornifrons, Stylonichia 597 Cyrtohymena macrostoma 314
cornipes, Oxytricha 247 Cyrtohymena marina 9, 234, 281, 295,323
cornuata, Oxytricha 247 Cyrtohymena muscorum 33,64,86, 106, 117, 123,
cornuta, Oxytricha 247 217,224,279,280,281, 291T, 305, 314, 325,
corona, Himantopus 514 331,344,480,488,490
corona, Kerona 514 Cyrtohymena avalis 317
crasistillata, Oxytricha 135 Cyrtohymena primicirrata 9, 59, 281, 291T, 294,
crasistylata, Oxytricha 135 296,300,629,632
crassa, Holosticha Trichotaxis 243 Cyrtohymena quadrinucleata 2, 4, 56, 60, 207, 279,
crassa, Oxytricha 116,243 280, 291T,317,320, 500, 677
crassa, Trichotaxis Oxytricha 243 Cyrtohymena saprope/ica 4, 157, 280, 281, 295,
crassistilata, Opisthotricha 135 306
crassistilata, Oxytricha 4, 119,135, 137,257 Cyrtohymena simplex 309
crassistilata, Oxytricha Opisthotricha 135 Cyrtohymena sp. 4, 281, 307,310,317,631
crassistylata, Opisthotricha 135 Cyrtohymena sphagnicola 632
cristata, Pseudourostyla 23, 26, 865 Cyrtohymena tetracirrata 280, 281, 29lT, 308, 317,
cultriformis, Oxytricha 253 320,643
cuneata, Kerona 840 Cyrtohymena torrenticola 280, 281, 295, 307
cuneatum, Ceratidium 597 cyrtophorids 4
Cuniculostomum monilata 607
curta, Oxytricha 247 decumana, Oxytricha 247
curvata, Kerona 840 deformis, Oxytricha 247
curvata, Oxytricha Stylonychia 593 depressa, Cyrtohymena candens 316, 317
curvata, Stylonchia 502 depressa, Steinia candens 314,315,316
curvata, Stylonichia 593 diademata, Kerona 840
curvata, Stylonychia 502, 503, 568, 588, 591, 593, diana, Plagiotricha 514
683 dilatata, Oxytricha truncata 254
Cyclidium pediculus 825, 826, 827 Diophrys appendiculata 596
Cyclidium radians 566 Diophrys bifaria 557
cyciidium, Trichoda 566 Diophrys scutum 44
cylindrica, Kerona 840 Dipleurostyla 893
 SYSTEMATIC INDEX 1053

Dip/eurosty/a acuminata 893 enigmatica, Oxytricha 121, 131T, 229, 232, 824,
Dip/eurosty/a tristy/a 242 933
discifera, Oxytricha 122,227,933 enigmatica, Paraurosty/a 843
discijera, Oxytricha Oxytricha 227 enigmatica, Ponturostyla 843
Discode//a mu/tipes 577 epaixijormis, Kerona 840
dispar, Paraurostyla 842 ephippioiaes, Oxytricha 248
dispar, Urostyla 842 Epiclintesfelis 243
dorsicirrata, Amphisie//a 803 Epistylis alpestris 84
dorsicirrata, Gastrostyla 20, 33, 36, 791, 802T, 803, erecthisticus, Histrio 650
817 erethesticus, Histriculus 650, 653
dorsiincisura, Urosomoida 4, 11, 194, 207, 345, erethesticus, Histrio 64 7
346, 354T, 362 erethisicus, Oxytricha Histrio 650
Drepanina 893 erethisticus, Histriculus 650
Drepaninafalcata 893 erethisticus, Histrio 647, 650, 667
dragescoi, Oxytricha 150,151 erethisticus, Oxytricha Histrio 650
dragescoi, Tachysoma 11,432,433, 442T, 463 erethysticus, Histrio 650
dubia, Oxytricha 247 erosa, Kerona 514
dubia, Parurosoma 492 erosa, Trichoda 514
dubia, Steinia 281, 282, 293 Euciliata 79
dubia, Tachysoma 281 euglenivora, Opisthotricha 136
dubium, Holosticha Parurosoma 491, 492 euglenivora, Oxytricha 4,119, 125, 135,136
dubium, Paruro/eptus 492 euglenivora, Oxytricha Opisthotricha 136
dubium, Parurosoma 11, 105, 106,492 Euhypotrichina 32
dubium, P/eurotricha 492 Eup/otes 87, 501, 603, 676
dubius, Kerona 504, 517 Eup/otes affinis 450
dujardiniana, Oxytricha 247 Eup/otes charon 450
dupla, Sty/onichia 598 Euplotes patella 296, 450, 692
dupla, Stylonychia 598 Euplotes vannus 839
durhamiensis, Oxytricha 10, 120,228,233,695 Euplotidae 79
euplotids 1,23,26,32,56,87,253,499,500,723
echinata, Oxytricha Tachysoma 435 eurystoma, Oxytricha 626
echinata, Pleurotricha 434 excavala, Stylonichia 598
echinata, Sty/onychia 431,434,444 exocijormis, Oxytricha 248
echinata, Tachysoma 434 extuberans, Kerona 840
ehrenbergiana, Oxytricha 247
eissei, Paraurostyla 845 falax, Oxytricha 138
e//ipso/des, Stylonichia 598 falcata, Drepanina 893
e/liptica, Oxytricha 150, 152, 156 fa//ax, Oxitricha 138,256
ellipticus, Uro/eptus 787 fa/lax, Oxytricha 6, 23, 25, 32, 84, 87, 121, 137,
elongata, Kerona 840 150, 152, 186, 191, 196,221,254,257,258,259,
eiongata, Opisthotricha 136, 242 565,650,684,930
elongata, Opistotricha 242 fa//ax, Oxytricha Oxytricha 138
elongata, Oxytricha 119, 136,164 fallox, Oxytricha 138
e/ongata, Oxytricha Opisthotricha 136, 242 fastigiata labiata, Oxytricha 246
emarginata, Opisthotricha 407, 409 fastigiata, Stylonichia 598
emarginata, Urosoma 35, 163, 369, 396, 397, 402, fastigiata, Sty/onychia 598
405T, 407, 409, 414 faurei, Oxytricha 10, 104, 120, 131T, 146
Enge/mannie//a 79, 423, 607, 893 faurei, Oxytricha Opisthotricha 146
Engelmannie/la mobilis 931 faurei, Peritromus 839
enigmatica, Hemigastrostyla 11,229,933, 937T felis, Epiclintes 243
enigmatica, Mixotricha 843 felis, Oxitricha 116
1054 SYSTEMATIC INDEX
jelis, Trichoda 116
jenestrata, Cyrtohymena 645
jenestrata, Steinia 644
jerruginea, Cyrtohymena 487
jerruginea, Oxytricha 484, 487, 489, 491
jerruginea, Oxytricha Steinia 487
jerruginea, Rubrioxytricha 11, 87, 123, 479, 480,
481,484,486T,487,491
jerruginea, Steinia 487, 625
jerruginea, Steinia Oxytricha 487
jerrujinea, Oxytricha 487
jeruginea, Oxytricha 487
jimbriata, Kerona 514
jimbriata, Oxytricha 248, 514
jimbriata, Trichoda 248,514
jissieta, Stylonychia 505
jissiseta, Oxytricha Stylonychia 566
jissiseta, Stylonychia 565, 566, 580
flXa, Podophrya 81
jlava carnea, Oxytricha 243
jlava, Oxytricha 243
jlava, Pseudokeronopsis 243
jlavicans, Urostyla 844, 845, 848, 870
j1exilis, Onychodromopsis 62, 68, 106, 261, 268,
475,476,478,479,782,787
j1exilis, Onychodromopsis sp. 479
jlexilis, Pleurotrycha 478
jormosa, Oxytricha 248
jossicola, Holosticha Trichotaxis 876
jossicola, Paraurostyla 876
jossicola, Trichotaxis 848, 874, 875
jranzi, Kahliella 369,391
jranzi, Orthoamphisiella 369
jranzi, Gonostomum 369
jromenteli, Oxytricha 227, 248
jronte-albens, Stylonichia 598
jrontosus, Stylonichia 598
jurcata, Oxytricha 248, 258, 457
jurcata, Oxytricha Tachysoma 248, 457
jurcata. Tachysoma 173,457
jurcatas, Orytricha 248
furcatas, Oxytricha 248
jurcatum, Tachysoma 457
jurcatus. Oxytricha 248, 457
fusca, Oxytricha 243
fusiformis, Balladyna 128
fusiformiS, Tachysoma 175, 178
ga/eata, Oxytricha 249
gallina, Oxitricha 116
gallina, Trichoda 116
gallina, Uroleptus 116
Gasterostyla setifera 816
Gasterostyla steini 789
Gasterostyla steinii 789
Gastostyla steini 789
Gastrostila 789
Gastrostyla 15, 26, 28, 56, 73, 78, 79, 80, 81, 103,
104, 105, 163, 260, 476, 601, 605, 699, 777, 778,
789,797,827,879,889,892,933
Gastrostyla affine 370
Gastrostyia dorsici"ata 20, 33, 36, 791, 802T, 803,
817
Gastrostyla hebbalica 818, 824
Gastrostyia minima 33, 791, 797, 802T, 809, 810,
817
Gastrostyla muscororum 808
Gastrostyla muscorum 2, 4, 186,790,808,818
Gastrostyla mystacea 87, 791, 808, 810, 811
Gastrostylaparasteinii 791, 792, 797
Gastrostylaphilippinensis 791,792,797
Gastrostyla Pleurotricha setifera 816
Gastrostyla pulchra 9,790, 791,818
Gastrostyia sedjera 790, 816, 818, 824
Gastrostyla sp. 824
Gastrostyla spp. 790
Gastrostyla steini 791,792,824
Gastrostyla stein;; 1,2,4,6, 19,25,26,33,54, 87,
88,111,646,694,789,790,791,802T,808,817
Gastrostyla steinni 792
Gastrostyla stenocephala 9,789, 791, 822
Gastrostyla sterkii 818, 820, 821
Gastrostyla Stylonychia pulchra 818, 824
Gastrostyla vorax 791,792,797
Gastrostylus 789
Gastrosyla 789
Gastrosyla stenocephala 789
Gastrotricha mystacea 808
Gastrotyla stenocephala 789
geleii, Holosticha 232
geieii, Oxytricha 4, 33, 119, 13IT, 230, 232
geleii, Trachelostyla 370
geleii, Gonostomum 370
germanica, Oxytricha 282
gibba, Oxytricha 243, 248
gibba, Paraurostyla 842
gibba, Trichoda 116, 842
gibbosa, Oxitricha 116
gigantea, Oxytricha 36, 425
gigantea, Oxytricha Urosoma 425
gigantea, Urosoma 11,35, 164,396,398, 405T, 425
gigantea, Urosoma fur Oxytricha 425
 SYSTEMATIC INDEX 1055

globifrons, Stylonichia 598 grandis, Onychodroma 722

Gonostomatidae 81,103,368 grandis, Onychodromas 722
Gonostomidae 102 grandis, Onychodromis 722
gonostomoida, Trachelochaeta 392 grandis, Onychodromus 4, 106, 669, 671, 676, 677,
gonostomoida, Gonostomum 105,368, 369, 382T, 722, 723, 724, 736T, 752, 753, 755
392 grandis, Onycodromus 724
Gonostomum 15, 17,20,32, 56, 57, 62, 67, 68, 69, grandis, Orynchodromus 723
73, 74, 75, 76, 77, 78, 79, 80, 81, 103, 104, 112, grandis, Oxychodromus 723
114,223,260,346,367,368,396,397,419,432, grandis, Oxytricha Stylonychia 671
840 grandis, Pleurotricha 1, 86, 87, 476, 698, 699, 709,
Gonostomum aJJine 9, 33, 59, 61, 88,103,106, 143, 712,716,717
368, 369, 382T, 384, 389, 391, 392, 393, 394, grandis, Pleurotricha Pleurotricha 713
395,449 grandis, Stylonychia 671,676,697
Gonostomum affinis 373 grandis, Stylonychia Onychodromus 669
Gonostomum affinis, Plagiotricha 370 grandis, Urostyla 23, 243, 487, 841, 848, 870
Gonostomum afxfine 379 granulata, Cyrtohymena 280,281,295,306
Gonostomum algicola 370, 373 granulata, Oxytricha Steinia 306
Gonostomum algicolum 370 granulata, Steinia 306
Gonostomum andoi 370,384 granulifera quadricirrata, Oxytricha 197,223
Gonostomum bryonicolum 370, 375 granulifera, Oxytricha 2, 10,33,44, 60, 64, 68, 71,
Gonostomum ciliophorum 370 75, 86, 106, 115, 116, 120, l3lT, 142, 159, 182,
GonostomumJranzi 369 191,194,197,207,223,271,335,352,359,430,
Gonostomum geleii 370 472,625,687,723,929,931
Gonostomum gonostomoido 105, 368, 369, 382T, granull/era, Paraurostyla 10, 844,874, 876T
392 granulifera, Parurosoma 261, 262, 263
Gonostomum kuehnelti 1,4,9,369, 382T, 391 granulifera, Tachysoma 459
Gonostomum mereschkowskii 368 granulifera, Urosomoida 11, 120, 346, 347, 352,
Gonostomum Oxytricha affine 370 354T
Gonostomum Oxytricha strenuum 384 granuliferum, Lamtostyla 432
Gonostomum parvum 369, 393 granuliferum, Tachysoma 11,432,433, 442T, 459
Gonostomum sp. 395 granulosa, Oxytricha 4, 10, 119,148
Gonostomum spirotrichoides 370, 375 grelli, Podophrya 81, 82, 84, 551
Gonostomum spp. 500 gyriniformis, Stylonichia 598
Gonostomum stenuum 384 gyrinioiaes, Oxytricha 249
Gonostomum sternuum 384
Gonostomum strenua 384 haematoplasma, Oxytricha 36, 479, 481
Gonostomum strenum 384 haematoplasma, Rubrioxytricha 8, 11,34,87, 106,
Gonostomum strenuum 9, 33, 242, 368, 369, 370, 123,125, 138,480, 481,486T,488
373, 375, 382T,384, 395 halophila, Opisthotricha 232, 256, 258
Gonostomun stenuum 395 halophila, Oxytricha 119, 170,228,232,258
Gonyostomum semen 395 halophi/a, Oxytricha Opisthotricha 232
gracilis, Cyrtohymena 4,9, 125,280,295,305 halophila, Oxytricha Opistotricha 232
gracilis, Oxytricha Steinia 305 halophyla, Opisthotricha 232
gracilis, Steinia 305 Halteria 23
gradus, Onychodromus 724 Halteria grandinella 2
grandinella, Halteria 2 harbinensis, Stylonychia 599
grandis simplex, Onychodromus 724 haustellum, Kerona 514
grandis simplex, Pnychodromus 724 haustrum, Kerona 514
grandis, Onchodromus 722 hebbalica, Gastrostyla 818, 824
grandis, Onichodromus 722 Hemigastrostyla 105,230,822,933
grandis, Onychodomus 722 Hemigastrostyla enigmatica 11,229,933, 937T
1056 SYSTEMATIC INDEX
Hemigastrostyla stenocephala 933
Hemisincirra 79, 397, 421, 893
Hemisincirrapolynuc!eata 36,397,419,893
Hemisincirra similis 419
henegui, Oxytricha 259
hengshanensis, Oxytricha 249
hennegui, Opisthonecta 259
Himantophorus 723
Himantopus acarus 514
Himantopus corona 514
Himantopus ludio 514
Himantopus sannio 515
Himantopus volutator 116
himenophora, Holosticha 787
Historia 646
Historio 646
Historio sp. 666
histria, Stylonychnia 650
Histrichulus vorax 646
Histriculus 14, 18,20,28,54,56,61,67,68, 73, 74,
75, 76, 77, 79, 80, 81, 103, 112,250,314,432,
456,645,668,695,699,716,771,827
Histriculus acuminatus 666
Histriculus admirabilis 646, 677
Histriculus cavicola 669
Histriculus complanatus 647,656,657
Histriculus complanatus, Oxytricha 668
Histriculus erethesticus 650, 653
Histriculus erethisticus 650
Histriculus histrio 8, 17, 18,33,57,63,66,67, 106,
140,629, 646,647, 654T,657, 667, 668, 691, 705
Histriculus histrio, Oxytricha 650
Histriculus hyalinus 650, 653
Histriculus lemani 791
Histriculus minimus 4,647,656,658
Histriculus muscorum 36, 646, 669, 683
Histriculus polycirratus 646, 666
Histriculus simi/is 25, 36, 568, 647, 660, 663, 665,
666,667,682,683,684,693,694,792
Histriculus similis tricirratus 36, 646, 682
Histriculus sp. 33
Histriculus sp. nov.? 667
Histriculus sphagni 647, 650, 655, 658
Histriculus triccirratus 682
Histriculus tricirratus 646, 682
Histriculus vorax 87,647,656,659,667
Histriculux vorax 646
histrie, Stylonychia 650
Histrio 261, 282, 314, 645, 646, 650, 684, 694
Histrio acuminatus 646, 666
Histrio acuminatus, Oxytricha 666
Histrio companatus 658
Histrio complanatus 657, 667
Histrio complanatus, Oxytricha 658
Histrio erecthisticus 650
Histrio erethesticus 647
Histrio erethisicus, Oxytricha 650
Histrio erethisticus 647,650,667
Histrio erethisticus, Oxytricha 650
Histrio erethysticus 650
Histrio histrio 647
Histrio histrio, Oxytricha 650
Histrio hyalinus 650
Histrio inquieta, Steinia 293
Histrio inquietus 313,646
Histrio Kerona histrio 647
Histrio lemani 646,791,792,797
Histrio macrostoma 646,683,684,691
Histrio muscorum 557, 646, 683, 684, 685, 689
Histrio muscorum, Oxytricha 116,684
histrio, Oxytricha Histriculus 650
Histrio pelionella 435
Histrio pellionella 650
Histrio polycirratus 666
Histrio quadrinucleatus 646, 665, 694
Histrio similis 665, 694, 696
Histrio similis, Oxytricha 183,665,694
Histrio sp. 667
Histrio sphagni 654
Histriosphagni, Oxytricha 190, 657
Histrio steini 650
Histrio steinii 605,645,646,647
Histrio vorax 659
Histrio vorax, Oxytricha 591,659
histrio, Cerona 647
histrio, Histriculus 8,17,18,33,57,63,66,67, 106,
140, 629,646,647,654T,657,667,668,691,705
histrio, Histrio 647
histrio, Histrio Kerona 647
histrio, Kerona 647,667
Histrio, Oxytricha 79, 116,282,646,684
histrio, Oxytricha Histrio 650
histrio, Paramaecium 501, 646, 647
histrio, Stylonichia 646, 650
histrio, Stylonychia 501, 647, 668
histrio, Stylonychnia 650
histrio, Trichoda 647
histrioides, Oxytricha 683, 684, 689
histriomuscorum complex, Sterkiella 668, 669
histriomuscorum, Oxytricha 683, 684
histriomuscorum, Sterkiella 16,26,28,30,31,34,
59, 64, 71, 86, 87, 106, 117, 118, 121, 122, 125,
 SYSTEMATIC INDEX
151,152,168,196,250,254,257,468,578,583,
593, 603, 604, 646, 658, 665, 666, 668, 669,
674T, 679, 681, 682, 683, 693, 695, 716, 723,
927T,929
histrix, Stylonichia 650
hologama, Paraurostyla 848
hologama, Urostyla 845, 848, 855, 870
hologamma, Urostyla 845
Holosticha 61, 128, 232, 243, 244, 250, 491, 492,
699,787,808,809,876,884
Holosticha camerounensis 769
Holosticha geleii 232
Holosticha himenophora 787
Holosticha Holosticha hymenophora 787
Holosticha hymenophora 781, 786, 787
Holosticha hymenophora, Holosticha 787
Holosticha kessleri 243
Holosticha macrostoma 769,884
Holosticha monilata 296
Holosticha mononucleata 492, 493
Holosticha multistilata 340, 785
Holosticha mystacea 808, 818, 824, 825
Holosticha mystacina 809
Holosticha oxytrichoidea 254
Holosticha Parurosoma 491
Holosticha Parurosoma dubium 491, 492
Holosticha pullaster 116, 242, 243, 252, 435, 444
Holosticha setigera 897
Holosticha similis 126, 128,930
Holosticha stueberi 470
Holosticha Trichotaxis 116,243,876
Holosticha Trichotaxis crassa 243
Holosticha Trichotaxisfossicola 876
Holostichidae 80,81,368,475,491,893,930
holostichids 56, 66, 67, 71, 73, 77, 432, 435, 766,
809,930
Holostischa oxytrichoidea 254
hortualis, Parentocirrus 4, 10, 113, 781, 878, 879,
883T
humicola humicola, Tachysoma 433, 442T, 452,
454,455,456
humicola longisetum, Tachysoma 433, 442T, 452,
455,456
humicola, Pseudouroleptus 888, 889
humicola, Tachysoma 4, 253, 432, 433, 452, 456,
458
humicola, Tachysoma humicola 433, 442T, 452,
454,455,456
humicola, Uroleptus 888
hyalin, Actinotricha 236
hyalina, Actinotricha 236, 241
1057
hyalina, Tachysoma 432
hyalinus, Histriculus 650, 653
hyalinus, Histrio 650
hydrarum, Peritromus 826, 827, 837
hymatostoma, Oxytricha 151
hymenophora, Apoamphisiel/a 781, 782, 786,872
hymenophora, Holosticha 781, 786, 787
hymenophora, Holosticha Holosticha 787
hymenophora, Paraurostyla 25,781,787,843
hymenostoma, Oxytricha 34, 36, 87, 117, 122, 138,
139,140,141,150,257,259
hymenostomata, Oxytricha 150, 151
Hypotricha 79
Hypotrichen nov. spec.? 597
Hypotrichida 79,80,81
Hypotrichidium 723
hystrio, Stylonichia 650
immemorata, Oxytricha 249
incrassata, Oxytricha 249
indica, Coniculostomum 608, 611, 618T, 619, 620
indica, Onychodromus 738,740,741,743,749,752
indica, Pleurotricha 607, 620, 623, 624, 699, 816
Infusoria 79
inqueta, Steinia 315
inquieta, Cyrtohymena 280, 281, 294, 309, 313,
317,320,324,625,626,629
inquieta, Oxytricha 313
inquieta, Oxytricha Steinia 294
inquieta, Steinia 281, 282, 300, 303, 314, 324
inquieta, Steinia Histrio 293
inquietus, Histrio 313, 646
islandica, Oxytricha 2, 4,118, BIT, 207, 346, 364
joblotii, Oxitricha 116, 245, 433
kahli, Onychodromopsis 476,477, 478
kahli, Stylonychia 521, 549
kahli, Uroleptus 421,892
Kahliella 74, 369, 607, 894, 930
Kahliellajranzi 369, 391
Kahliella simplex 931
KahlieIlidae 73, 78, 879, 889,931
kahlieIlids 75, 607, 843, 882, 893, 930
kahlovata, Oxytricha 122, 157
karini, Urosoma 11,396,398, 405T, 422
Kerona 15, 79, 81, 103, 104, 244, 597, 777, 825,
826,894
Kerona absidata 839
Kerona acarus 514
Kerona anjracta 839
1058 SYSTEMATIC INDEX

Kerona aper 839 Kerona triangularis 841

Kerona arcuata 839 Kerona truncata 841
Kerona beccijormis 839 Kerona unguicularis 841
Kerona bicaudata 839 Kerona urostyla 897
Kerona bipes 839 Kerona vannus 839
Kerona calvaejormis 839 Kerone pustulata 601
Kerona calvitium 700 Keronidae 80, 81,102,827
Kerona caudata 839 Keronina 102
Kerona ciliata 839 Keronopsis 88, 128, 663, 827
Kerona congyrans 839 Keronopsis longicirrata 155, 163
Kerona corona 514 kessleri, Holosticha 243
Kerona cuneata 840 kessleri, Oxytricha 243
Kerona curvata 840 Komplex, Stylonychia mytilus- 506
Kerona cylindrica 840 kuehnelti, Gonostomum 1,4,9,369, 382T, 391
Kerona cypris 540
Kerona diademata 840 labiata, Oxytricha 247, 249
Kerona dubius 504, 517 labiata, Oxytrichajastigiata 246
Kerona elongata 840 labiata, Stylonichia 598
Kerona epalxijormis 840 lacerata, Kerona 840
Kerona erosa 514 lacerata, Oxytricha 249
Kerona extuberans 840 lacrimula, Oxytricha 249
Kerona jimbriata 514 laevis, Pseudourostyla 39
Kerona haustellum 514 lamottei, Lamtostyla 893
Kerona haustrum 514 Lamtostyla 79,81,104,432,893,894,930
Kerona histrio 647, 667 Lamtostyla granulijerum 432
Kerona histrio, Histrio 647 Lamtostyla lamottei 893
Kerona lacerata 840 Lamtostyla longa 432
Kerona lanceolata 699 Lamtostyla perisincirra 432
Kerona ludio 514 Lamtostyla rap tans 432, 893
Kerona mamillata 840 lanccolata, Pleurotricha 700
Kerona multipes 873 lanceolata, Cerona 699
Kerona mytilus 504, 517, 549 lanceolata, Kerona 699
Kerona oblonga 840 lanceolata, Oxytricha 34,73, 121, 131T, 209
Kerona ondulata 840 lanceolata, Oxytricha Oxytricha 209
Kerona ovata 840 lanceolata, Pleurotricha 106, 261, 262, 268, 449,
Kerona pediculus 33, 86, 87, 103, 104, 105, 111, 505,517,698,699, 710T, 716,816
723,741,748,751,826 lanceolata, Pleurotricha Allotricha 700
Kerona polymorphum 826 lanceolata, Stylonychia 698, 699, 717
Kerona polypora 826 lanceolatum, Pleurotricha 700
Keronapolyporum 825, 826, 827, 837, 841 lata, Oxytricha 250
Kerona pustulata 502, 565, 600, 825 latissima, Paraurostyla 842
Kerona rastellum 840 latissima, Urostyla 842
Kerona rostrata 515,840 Laurentia 79,80,752,755
Kerona rotunda 839 Laurentia acuminaba 753
Kerona rotundata 841 Laurentia acuminata 753, 756, 757, 761, 763, 764T
Kerona sannio 515 Laurentia macrostoma 752, 753, 757, 763
Kerona silurus 504, 566, 600, 601 Laurentia monilata 606, 608, 755, 763
Kerona silurus congyrans 839 Laurentiella 15, 56, 80, 81, 103, 104,470,607,608,
Kerona subrotondata 840 699,717,752
Kerona subrotundata 840 Laurentiella acuminata 25, 753
Kerona tremula 841 Laurentiella Laurentiella 753
 SYSTEMATIC INDEX
Laurentiella Laurentiella macrostoma 753
Laurentiella Laurosticha 606
Laurentiella Laurosticha monilata 608
Laurentiella macrostoma, Laurentiella 753
Laurentiella monilata 608
LaurentieHa strenua 4, 105, 106, 500, 741, 753,
764T, 769, 773
Laurentiella, Laurentiella 753
Laurosticha 80
Laurosticha monilata, Laurentiella 608
Laurosticha, Laurentiella 606
Lautosticha 607
lemani, Histriculus 791
lemani, Histrio 646, 791, 792, 797
lemnae, Stylonchia 502
lemnae, Stylonychi 502
lemnae, Stylonychia 5, 6, 12, 37, 82, 84, 205, 502,
503,506,509,510,511,513,521,524,525,526,
529,533,534T,548,549,550, 676, 752
Leptocirra tekirghiolica 492
lepus, Oxitricha 115
lepus, Oxytricha 116
lepus, Trichoda 115
limbiformis, Stylonichia 598
limbifrons, Stylonichia 598
lingua, Oxytricha 250
longa, Lamtostyla 432
longa, Oxytricha 34, 64, 68, 121, 13lT, 158, 170,
184,186,346,347,356,359,436
longa, Tachysoma 432
longi-caudata, Oxytricha 243
longicirrata, Keronopsis 155, 163
longicirrata, Oxytricha 4, 119, 163,495,778
longicirrata, Oxytricha Urosoma 163
longicirrata, Urosoma 163
longigranulosa, Oxytricha 10,34, 120, BIT, 194,
213
longipes, Oxytricha 250
longisetum, Tachysoma humicola 433, 442T, 452,
455,456
iongissima, Oxytricha 104, 119, 136, 164,470
ludibunda, Oxytricha 150, 156,259
ludio, Himantopus 514
ludio, Kerona 514
ludio, Trichoda 514
lundi, Oxytricha 150, 152
lundibunda, Oxytricha 151
luteolucens, Oxytricha 250
Iynceus, Aspidisca 737, 839
lynchi, Urostyla 845, 848, 855, 870
1059
macrostoma, Cyrtohymena 314
macrostoma, Histrio 646,683,684,691
macrostoma, Holosticha 769, 884
macrostoma, Laurentia 752, 753, 757, 763
macrostoma, Laurentiella Laurentiella 753
macrostoma, Opisthotricha 152
macrostoma, Opistotricha 150, 156
macrostoma, Paraurostyla 843
macrostoma, Pleurotricha 699
macrostoma, Steinia 314, 315
macrostoma, Stienia 314
macrostoma, Trachelostyla 370
macrostoma, Urosoma 370, 373
macrostya, Urosoma 404
macrostyla, Oxytricha 404, 407
macrostyla, Oxytricha Stylonychia 600
macrostyla, Oxytricha Urosoma 404, 600
macrostyla, Stylonichia 600
macrostyla, Stylonychia 600
macrostyla, Urosoma 36, 125, 398, 404, 405T, 409,
931
macrostyla, Urosoma Oxytricha 404, 409
maeceps, Oxytricha 252
magna, Oxytricha 669, 671, 676
makrostyla, Sty/onychia 600
mamillata, Kerona 840
mamillata, Stylonichia 598
marina, Cyrtohymena 9, 234, 281, 295, 323
marina, Oxytricha 233, 256, 259
marina, Oxytricha Oxytricha 234, 323
marina, Oxytricha Steinia 234, 323
marina, Paraurostyla 842
marina, Steinia 323
marina, Urostyla 842
marioni, Amphisiella 790
massula, Stylonichia 598
matritensis, Oxytricha 122, 165, 169, 170
maupasi, Ancystropodium 4, 14, 105, 164, 568,
777,778,790
membranaceus, Stylonichia 599
mereschkowskii, Gonostomum 368
merula, Oxytricha 250
Mesotricha 431
micans, Oxytricha 243
Micromitra 893
minima, Gastrostyla 33, 791, 797, 802T, 809, 810,
817
minima, Oxytricha 150, 151
minima, Urosomoida 35, 345, 346, 347, 351, 354T,
366
minimum, Parahistriculus 658
1060 SYSTEMATIC INDEX
minimus, Histriculus 4,647,656,658
minimus, Parahistriculus 645, 658
minimus, Parahistricus 646
minor, Opisthotricha parallela 166
minor, Oxytricha 122, 125, 137, 150, 151, 156, 166,
172,259,683
minor, Oxytricha Opisthotrichaparalella 166
minor, Oxytricha Oxytricha 125
mirabile, Tachysoma 434, 444
mirabilis, Oxytricha Tachysoma 435
mirabilis, Tachysoma 434
mistacea, Oxytricha 809
mitilus 511
mitilus, Styloichia 511
mitilus, Stylonichia 511
Mitra 893
Mitra radiosa 893
Mixotricha 843
Mixotricha enigmatica 843
mobilis, Engelmanniella 931
mobilis, Uroleptus 893
mol/is, Allotricha 9, 33, 261, 262, 267T, 268, 270,
272,704,782,848
monilata, Coniculostomum 1, 33, 46, 48, 49, 50, 51,
64, 106,606, 607,608,618T, 621,624, 706, 763,931
monilata, Cuniculostomum 607
monilata, Holosticha 296
monilata, Laurentia 606, 608, 755, 763
monilata, Laurentiella 608
monilata, Laurentiella Laurosticha 608
monilata, Pleurotricha 607
mononuc/eata, Holosticha 492, 493
mononuc/eata, Pleurotricha 492
monostylus, Stylonychia 600
monostylus, Stynolychia 502
monspessulana, Opisthotricha 36, 168,669
monspessulana, Oxytricha 122, 168
monstrosa, Oxytricha 250
monstrosa, Tachysoma 250
multipes, Discodella 577
multipes, Kerona 873
multipes, Oxytricha 848, 873
multipes, Urostyla 873
multipes, Urostyla Oxytricha 873
multipes, Urostyla trichogaster 873
multiseta, Oxytricha 122,166,169, 184, 190
multistilata, Holosticha 340, 785
muscororum, Gastrostyla 808
muscorum, Cyrtohymena 33,64,86,106,117,123,
217, 224, 279, 280, 281, 29lT, 305, 314, 325,
331,344,480,488,490
muscorum, Gastrostyla 2, 4, 186,790,808,818
muscorum, Histriculus 36,646,669,683
muscorum, Histrio 557, 646, 683, 684, 685, 689
muscorum, Opisthotricha 217
muscorum, Opithotricha 117
muscorum, Oxytricha 217,281,344,595,684
muscorum, Oxytricha Histrio 116,684
muscorum, Oxytricha Opisthotricha 116, 217, 684
muscorum, Oxytricha Steinia 116,279,282,684
muscorum, Oxytricha Stylonychia 116, 595,684
muscorum, Steinia 281, 282
muscorum, Stylonichia 601
muscorum, Stylonychia 595, 602
mutabunda, Stylonichia 599
mutilis, Stylonychia 511
mutilus, Stylonichia 511
mutilus, Stylonychia 511
mysilus, Stylonychia 511
mystacea, Gastrostyla 87,791,808,810,811
mystacea, Holosticha 808, 818, 824, 825
mystacea, Oxytricha 808
mystacea, Gastrotricha 808
mystacina, Holosticha 809
mystacina, Oxytricha 808
mystilus, Stylonychia 511
mythilus, Stylonychia 504, 505
mytilis, Stylonichia 505
mytilis, Stylonychia 511
mytillus, Stylonichia 505, 511
mytillus, Stylonychia 513
mytiluns, Stylonychia 513
mytilus-Komplex, Stylonychia 506
mytilus complex, Stylonychia 1, 2, 6, 8, 14, 35, 39,
40,41,42,43,71,81,85,87,88, 106, 143,248,
403, 450, 502, 503, 504, 549, 550, 551, 554,
555, 562, 568, 577, 588, 592, 598, 599, 600,
601, 602, 603, 604, 607, 611, 617, 640, 704,
755, 759, 761
mytilus pusilla, Stylonychia 555
mytilus syngen 1, Stylonychia 506
mytilus syngen 2, Stylonychia 510
mytilus Varietat 1, Stylonychia 505,550
mytilus Varietat 2, Stylonychia 505,549
mytilus, Cerona 504
mytilus, Kerona 504, 517, 549
mytilus, Oxytricha Stylonychia 505, 511
mytilus, Stilonychia 501
mytilus, Stylonchia 502, 895
mytilus, Stylonichia 504, 505, 506, 565, 601
mytilus, Stylonichnia 502
mytilus, Stylonichus 502
 SYSTEMATIC INDEX 1061

mytilus, Stylonychia 2,4, 17,18,20,21,23,25,26, num. 3, Oxitricha 245

34,44,49,51, 52, 54, 55, 64, 66, 67, 68, 70, 75, nuptacina, Oxytricha 809
76,77,78,84,248,251,256,501,502,503,504,
505,506,509,510,511,513,514,517,519,521, oblonga, Kerona 840
524, 525, 526, 534T, 537, 544, 549, 550, 551, oblonga, Oxytricha 251
552,553,555,560, 571,581,592,597,598,599, oblongatus, Oxytricha 251
600,602,603,606,625,687, 737, 755, 757, 870, ob/ongua, Oxytricha 251
929 obtusa, Oxytricha 251
mytilus, Stylonychna 502 octinostylus, Stylonichia 599
mytilus, Stylonychnia 502 octonucleata, Urosoma 2, 4, 396, 397, 405T, 417
mytilus, Styloychia 502 octonucleatus, Australocirrus 470, 717, 718
mytilus, Trichoda 501, 504, 549 octonucleatus, Rigidocortex 2, 4, 106, 113, 676,
mytylus, Stylonichia 511 718
mytylus, Stylonychia 513 oculata, Oxytricha 243
Ocytricha sp. 117
naine, Urostyla weissei 845 oligotrichid 253
naumanni, Paraurostyla 842 oligotrichs 4, 23
naumanni, Urostyla 842 oltenica, Oxytricha 251
nauplia, Oxytricha 121, 131T, 215 Onchodromus grandis 722
Neogeneia 607 ondulata, Kerona 840
New species of Oxytricha 245, 679 Onichadromus grandis 722
nodulinucleata, Stylonychia 2, 4, 503, 552 oniscus, Volvox 566
non-euplotids 1,23, 103, 104,500 Onycchodromus acuminatus 753
Nothopleurotricha 80,789,790 Onychdromus 722
Nothopleurotricha seti/era 816 Onychodomus grandis 722
Notocephalus 432 Onychodroma grandis 722
Notacephalus parvulus 432 Onychodromas 722
Notohymena 17,46,62,66,67,68,70,73,75,76, Onychodromas grandis 722
77,112,280,288,292,324,346,349,480,625 Onychodromis grandis 722
Notohymena antarctica 9,324,325,334, 339T Onychodromopsis 76, 79, 80, 81, 104, 106, 114,
Notohymena australis 9, 324, 325, 334, 339T, 340 261,268,475,607,781,782
Notohymena rubescens 9, 33, 59, 63, 64, 106,224, Onychodromopsis flexilis 62, 68, 106, 261, 268,
324,325, 339T,490 475,476,478,479,782,787
Notohymena selvatica 324,325, 339T, 344 Onychodromopsis kahli 476,477,478
notophora, Oxytricha Stylonychia 553 Onychodromopsis sp. 475, 476
notophora, Stylonichia 553, 601 Onychodromopsis sp.j1exilis 479
notophora, Stylonychia 36,205, 503, 525, 530, 549, Onychodromopsis tihanyiensis 476,781,782,843
553,562,565,568,578,588,592,594,603 Onychodromopsis variabilis 268
notophra, Stylonychia 554 Onychodromous quadricomutus 722
Notoxoma 885 Onychodromus 15, 56, 74, 79, 80, 81, 103, 104,
notrophora, Stylonychia 553 105,106,470,475,607,608,699,717,722,777,
nov. spec.?, Hypotrichen 597 842
nova, Oxytricha 6, 75,250, 578,685,723,929 Onychodromus acuminatus 753
n. sp., Stylonychia 605 Onychodromus gradus 724
nuda, Balladinopsis 893 Onychodromus grandis 4, 106, 669, 671, 676, 677,
nuda, Bal/adynopsis 893 722, 723, 724, 736T, 752, 753, 755
num. 1, Oxitricha 245 Onychodromus grandis simplex 724
num. 1, Pleurotricha 716 Onychodromus grandis, Stylonychia 669
num. 1, Stylonichia 599 Onychodromus indica 738,740,741,743,749,752
num. 2, Oxitricha 245 Onychodromus quadricornatus 740
num. 2, Pleurotricha 716 Onychodromus quadricomutis 740
1062 SYSTEMATIC INDEX

Onychodromus quadricornutus 4, 18, 75, 103, 578, Opisthotricha sphagni, Oxytricha 190, 657
723,724,736,738, 750T, 761, 773,827,833,884 Opisthotricha tericola 245
Onychodromus quedricomutus 740 Opisthotricha terrestris 683, 684
Onychodromus spp. 500,755,769 Opisthotricha, Oxytricha 79, 116, 146, 282
Onychodromus strenuus 755 Opistotricha 117
Onychodronum sp. 475 Opistotricha elongata 242
Onychodronus 722 Opistotricha halophila, Oxytricha 232
Onycodromus 722 Opistotricha macrostoma ISO, 156
Onycodromus grandis 724 Opistotricha ovata 157,255
Onyochodromus 722 Opistotricha parallela 171
Onytricha 117 Opistotricha similis 117
opisthomuscorum, Oxytricha 4, 119, 128, 135, 137, Opistotricha terrestris 683, 684, 689
217,925, 927T Opistotricha terricola 245
Opisthonecta hennegui 259 Opistrotricha similis 117
Opisthotricha 20, 80, 81, 115, 117, 135, 136, 166, Opithotricha muscorum 117
170, 171, 183, 190,217,220,232,242,255,282, Opithotricha parahalophila 170
683 Orthoamphisiella 369
Opisthotricha crassistilata 135 Orthoamphisiellajranzi 369
Opisthotricha crassistilata, Oxytricha 135 OrthoamphisielIidae 894, 930, 931
Opisthotricha crassistylata 135 Orynchodromus grandis 723
Opisthotricha elongata 136, 242 Orytrichafurcatas 248
Opisthotricha e/ongata, Oxytricha 136,242 oscitans, Australocirrus 17, 106, 113, 470, 474T,
Opisthotricha emarginata 407, 409 717
Opisthotricha eug/enivora 136 ottowi, Oxytricha 10, 118, BIT, 218
Opisthotrichajaurei, Oxytricha 146 ova/e, Sty/onychia 593
Opisthotricha halophila 232, 256, 258 ovalis, Cyrtohymena 317
Opisthotricha ha/ophila, Oxytricha 232 ova/is, Oxytricha 227, 248, 252
Opisthotricha halophyla 232 ovalis, Oxytricha Oxytricha 227
Opisthotricha macrostoma 152 ovalis, Steinia 314, 317
Opisthotricha monspessulana 36,168,669 ovalis, Stylonychia 591
Opisthotricha muscorum 217 ovata, Kerona 840
Opisthotricha muscorum, Oxytricha 116, 217, 684 ovata, Opisthotricha 157
Opisthotricha ovata 157 ovata, Opistotricha 157,255
Opisthotricha ovata, Oxytricha 157,251 ovata, Oxytrica 251
Opisthotricha ovata, Oxytricha 157,251 ovata, Oxytricha 157,251
Opisthotricha Oxytricha paralleta 171 ovata, Tachysoma 4, 11,432,433, 442T, 464
Opisthotricha Oxytricha similis 183 Oxitrica sp. 117
Opisthotricha parahalophila 170 Oxitricha 115, 116
Opisthotricha paraholophila 170 Oxitricha ambigua 116
Opisthotricha para/ella minor, Oxytricha 166 Oxitricha bulla 116
Opisthotrichaparalle/a 166,171,172,255 Oxitricha cavico/a 677
Opisthotricha paralle/a minor 166 OXitrichajallax 138,256
Opisthotricha parallela, Oxytricha 171 OXitrichajelis 116
Opisthotricha parallelis 171, 255 Oxitricha gallina 116
Opisthotricha procera 220, 255 Oxitricha gibbosa 116
Opisthotricha procera, Oxytricha 220 Oxitrichajob/otii 116, 245, 433
Opisthotricha similis 169, 183,255 Oxitricha lepus 115
Opisthotricha similis, Oxytricha 183,665,694 Oxitricha num. 1 245
Opisthotricha sordis 245 Oxitricha num. 2 245
Opisthotricha sp. 679, 682 Oxitricha num. 3 245
Opisthotricha sphagni 190 Oxitricha pelionella 115,433
 SYSTEMATIC INDEX
Oxitricha pUlex 115
Oxitricha pullaster 116
OxitricOO pullicina 115,246
Oxitricha spec. 117
Oxitricha transjUga 116
Oxitricha variabilis 116, 193,246
OxitricOO volutator 116
Oxitrycha sp .117
Oxychodromus grandis 723
oxymarina, Oxytricha 121,233
Oxythricha 117
OxyticOO 117
Oxytrica ovata 251
Oxytricha ? parvula 252
Oxytricha 1, 14, 17,24,26,28,54,55,56,61,62,
66,67,68,69,73,75,76,77,79,80,81,103,104,
113, 114, 115, 150,205,243,244,248,250,251,
254,256,261,279,282,305,306,309,323,331,
345,346,348,352,359,365,375,396,397,400,
404,414,425,431,432,435,457,461,466,468,
480,487, 495, 501, 505, 553, 555, 557, 566, 588,
591,593, 595, 597, 625, 626, 646, 650, 657, 658,
659,665,666,671,684,687,694,790,848,887
OxytricOO ActinotricOO saltans 236
Oxytricha acuminata 246, 414
Oxytricha acuminata, Urosoma 414
OxytricOO aeroginosa 122
Oxytricha aeruginosa 9, 120, 122, 166, 169, 209,
224,254,256,480,488
Oxytricha aeruginosa, Oxytricha 123
Oxytricha affine, Gonostomum 370
OxytricOO affinis 367, 368, 369, 373
Oxytricha agilis 36, 347, 467
Oxytricha alba 242
Oxytricha alfredi 120, 125,151,167
Oxytricha alfredkahli 121, 227
Oxytricha ambigua 246
OxytricOO anca 246
OxytricOO arcuata 246
Oxytricha auricularis 243
Oxytricha auripunctata 10, 121, 123, 131T, 194,
224,346,347,364,684
Oxytricha australis 340
Oxytricha ba//adyna 4, 119, 126, 131T, 177, 178,
183,457,659
Oxytricha balladynula 177, 178
Oxytricha barbula 247
Oxytricha becci/ormis 247
OxytricOO bicirrata 457
OxytricOO bifaria 14, 26, 37, 250, 557, 565, 568,
571,578,579,581,584,601,929
1063
OxytricOO bi/aria, OxytricOO 557
Oxytricha bilobata 247
Oxytricha bimembranata 121, 138,196,221,683
Oxytricha bivacuolata 641
Oxytricha bUitkampi 175, 178
Oxytricha candens 314
Oxytricha candens aestuarii 317
OxytricOO capitata 243
Oxytricha caudata 396, 398, 401, 430
Oxytricha caudata, Urosoma 398
OxytricOO caudens 309
Oxytricha cavicola 668, 669
Oxytricha chilensis 452
Oxytricha ch/orel/igera 8, 86, 87, 119, 129, 256,
605
Oxytricha chlorelligera, OxytricOO 129
Oxytricha chlorigella 129,256
Oxytricha comipes 247
Oxytricha comuata 247
Oxytricha comuta 247
Oxytricha crasistillata 135
Oxytricha crasistylata 135
Oxytricha crassa 116, 243
Oxytricha crassa, Trichotaxis 243
Oxytricha crassistilata 4, 119,135, 137,257
Oxytricha cultri/ormis 253
Oxytricha curta 247
Oxytricha cypris 247
Oxytricha decumana 247
Oxytricha deformis 247
Oxytricha disclfera 122, 227, 933
OxytricOO disci/era, Oxytricha 227
Oxytricha dragescoi 150, 151
Oxytricha dubia 247
Oxytricha dujardiniana 247
Oxytricha durhamiensis 10, 120,228, 233, 695
Oxytricha ehrenbergiana 247
Oxytricha elliptica 150, 152, 156
Oxytricha e/ongata 119, 136, 164
Oxytricha enigmatica 121, 131T, 229, 232,824,933
Oxytricha ephippio'ides 248
Oxytricha euglenivora 4, 119, 125, 135, 136
Oxytricha eurystoma 626
Oxytricha exoci/ormis 248
Oxytrichafalax 138
Oxytrichafallax 6,23,25,32, 84, 87, 121,137, 150,
152, 186, 191, 196,221,254,257,258,259,565,
650,684,930
Oxytrichafallax, OxytricOO 138
Oxytrichafallox 138
Oxytrichafastigiata labiata 246
1064 SYSTEMATIC INDEX

Oxytrichajaurei 10, 104, 120, BIT, 146 Oxytricha inquieta 313

Oxytrichajerruginea 484,487,489,491 Oxytricha islandica 2, 4, 118, BIT, 207, 346, 364
Oxytrichaferruginea, Steinia 487 Oxytricha kahlovata 122, 157
Oxytrichajerrujinea 487 Oxytricha kessleri 243
Oxytrichaferuginea 487 Oxytricha labiata 247, 249
Oxytrichajimbriata 248,514 Oxytricha lacerata 249
Oxytrichaflava 243 Oxytricha lacrimula 249
Oxytrichaflava carnea 243 Oxytricha lanceolata 34,73, 121, 131T, 209
Oxytricha formosa 248 Oxytricha lanceolata, Oxytricha 209
Oxytrichafromenteli 227,248 Oxytricha lata 250
Oxytrichafurcata 248,258,457 Oxytricha lepus 116
Oxytrichafurcatas 248 Oxytricha lingua 250
Oxytrichafurcatus 248, 457 Oxytricha longa 34, 64, 68, 121, 131T, 158, 170,
Oxytrichafusca 243 184,186,346,347,356,359,436
Oxytricha galeata 249 Oxytricha longi-caudata 243
Oxytrichageleii 4,33,119, BIT, 230, 232 Oxytricha longicirrata 4, 119,163,495, 778
Oxytricha germanica 282 Oxytricha longigranulosa 10, 34, 120, 13IT, 194,
Oxytricha gibba 243, 248 213
Oxytricha gigantea 36, 425 Oxytricha longipes 250
Oxytricha gigantea, Urosoma filr 425 Oxytricha longissima 104, 119, 136, 164,470
Oxytricha granulifera 2, 10, 33, 44, 60, 64, 68, 71, Oxytricha ludibunda 150, 156,259
75,86, 106, 115, 116, 120, 131T, 142, 159, 182, Oxytricha lundi 150, 152
191,194,197,207,223,271,335,352,359,430, Oxytricha lundibunda 151
472,625,687,723,929,931 Oxytricha luteolucens 250
Oxytricha granulifera quadricirrata 197, 223 Oxytricha macrostyla 404, 407
Oxytrichagranulosa4, 10, 119, 148 Oxytricha macrostyla, Urosoma 404, 409
Oxytricha gyrinioiaes 249 Oxytricha maeceps 252
Oxytricha haematoplasma 36, 479, 481 Oxytricha magna 669, 671, 676
Oxytricha ha/ophi/a 119, 170,228,232,258 Oxytricha marina 233, 256, 259
Oxytricha henegui 259 Oxytricha marina, Oxytricha 234, 323
Oxytricha hengshanensis 249 Oxytricha matritensis 122, 165, 169, 170
Oxytricha Histriculus complanatus 668 Oxytricha merula 250
Oxytricha Histriculus histrio 650 Oxytricha micans 243
Oxytricha Histrio 79, 116,282,646,684 Oxytricha minima 150, 151
Oxytricha Histrio acuminatus 666 Oxytricha minor 122, 125, 137, 150, 151, 156,166,
Oxytricha Histrio complanatus 658 172,259,683
Oxytricha Histrio erethisicus 650 Oxytricha minor, Oxytricha 125
Oxytricha Histrio erethisticus 650 Oxytricha mistacea 809
Oxytricha Histrio histrio 650 Oxytricha monspessulana 122, 168
Oxytricha Histrio muscorum 116, 684 Oxytricha monstrosa 250
Oxytricha Histrio similis 183,665,694 Oxytricha multipes 848, 873
Oxytricha Histrio sphagni 190,657 Oxytricha multipes, Urostyla 873
Oxytricha Histrio vorax 591,659 Oxytricha multiseta 122, 166,169,184,190
Oxytricha histrioides 683, 684, 689 Oxytricha muscorum 217, 281, 344, 595, 684
Oxytricha histriomuscorum 683, 684 Oxytricha mystacea 808
Oxytricha hymatostoma 151 Oxytricha mystacina 808
Oxytricha hymenostoma 34, 36, 87, 117, 122, 138, Oxytricha nauplia 121, 131T, 215
139, 140, 141,150,257,259 Oxytricha nova 6, 75, 250, 578, 685, 723,929
Oxytricha hymenostomata 150, 151 Oxytricha nuptacina 809
Oxytricha immemorata 249 Oxytricha oblonga 251
Oxytricha incrassata 249 Oxytricha oblangatus 251
 SYSTEMATIC INDEX 1065

Oxytricha oblongua 251 Oxytricha phytojaga 564

Oxytricha obtusa 251 Oxytricha phytophaga 562
Oxytricha oculata 243 Oxytricha piscis 116
Oxytricha oltenica 251 Oxytricha pisciunculiformis 252
Oxytricha opisthomuscorum 4, 119, 128, 135, 137, Oxytricha pistillo'ides 252
217,925, 927T Oxytricha Plagiotricha strenua 395
Oxytricha Opisthotricha 79, 116, 146,282 Oxytricha plana 175, 178
Oxytricha Opisthotricha crassistilata 135 Oxytricha planctonica 494
Oxytricha Opisthotricha elongata 136, 242 Oxytricha planctontica 494
Oxytricha Opisthotricha eugleni1lora 136 Oxytricha plastystoma 626
Oxytricha Opisthotrichajaurei 146 Oxytricha platistoma 650
Oxytricha Opisthotricha halophila 232 Oxytricha platysoma 626
Oxytricha Opisthotricha muscorum 116,217,684 Oxytricha platystoma 116,259,308,309,313,314,
Oxytricha Opisthotricha o1lata 157, 251 317,624,625,626,629,630,645
Oxytricha Opisthotricha paralella minor 166 Oxytricha platystoma, Steinia 626
Oxytricha Opisthotricha parallela 171 Oxytricha pleuronectes 252
Oxytricha Opisthotricha procera 220 Oxytricha plicata 252
Oxytricha Opisthotricha similis 183,665,694 Oxytricha praeceps 252
Oxytricha Opisthotricha sphagni 190,657 Oxytricha proboscis 252
Oxytricha Opistotricha halophila 232 Oxytrichaprocera 10,121,220
Oxytricha ottowi 10, 118, BIT, 218 Oxytricha protensa 252
Oxytricha o1lalis 227, 248, 252 Oxytrichaproximata 121,191,196,221,258
Oxytricha o1lalis, Oxytricha 227 Oxytrichapseudojurcata 4,119,173,457
Oxytricha O1Iata 157,251 Oxytrichapseudojusijormis 4,119, BIT, 222
Oxytricha oxymarina 121, 233 Oxytricha pseudosimilis 10, 34, 118, BIT, 222
Oxytricha Oxytricha 79, 116 Oxytricha pullaster 116
Oxytricha Oxytricha aeruginosa 123 Oxytricha pustulata 259
Oxytricha Oxytricha bifaria 557 Oxytricha pustulata, Stylonychia 565
Oxytricha Oxytricha chlorelligera 129 Oxytricha quadrici"ata 10, 121, BIT, 197,223,684
Oxytricha Oxytricha discifera 227 Oxytricha quadrinuc/eata 252
Oxytricha Oxytrichajallax 138 Oxytricha quercineti 252
Oxytricha Oxytricha lanceolata 209 Oxytricha radians 253
Oxytricha Oxytricha marina 234, 323 Oxytricha retractilis 244
Oxytricha Oxytricha minor 125 Oxytricha rigescens, Tachysoma 461
Oxytricha Oxytricha o1lalis 227 Oxytricha rostrata 253
Oxytricha Oxytricha tricornis 242 Oxytricha rubipuncta 224
Oxyrrichaparahalophila 122, 170,232,233 Oxytricha rubra 244,281,282,293,344,401
Oxytrichaparallela 115, 118, 167,171 Oxytricha rubripuncta 10, 34, 121, 123, BIT, 194,
Oxytricha parallela, Opisthotricha 171 224,325
Oxytricha parvistyla 259, 467 Oxytricha salmastra 424
Oxytricha pelionella 158, 434, 435 Oxytricha saltans 122, BIT, 236, 253, 931
Oxytricha pellinella 450 Oxytricha saprobia 86, 87, 120, 174,259
Oxytricha pellionela 435, 467 Oxytricha saprobica 174
Oxytricha pellionella 115, 116, 183, 184,433, 434, Oxytricha sardida 260
467,468 Oxytricha scutellum 244
Oxytricha pellionella chilensis 451 Oxytricha scutum 557
Oxytricha pellionella. Tachysoma 434, 468 Oxytricha sel1latica 344
Oxytricha pellionella, Taxysoma 432 Oxytricha setifera 177,816
Oxytricha pellionnella 435 Oxytricha setigera 1, 2, 4, 5, 17, 34, 57, 87, 117,
Oxytricha perionella 435 118, 128, l3lT, 173, 175, 185, 187,213,217,
Oxytricha pernix 244 447,457,659,816
1066 SYSTEMATIC INDEX
Oxytricha simi/is 87, 121, 128, l36, 159, 169, 183,
220,222,260,436
Oxytricha similis, Opisthotricha 183
Oxytricha siseris 62, 104, ll8, l31T, 187
Oxytricha sordida 123, 125,260
Oxytricha sordis 245
Oxytricha sp. 20,26,45,86, 122, l31T, 190,251,
260,679,682
Oxytricha sphagni 4, 119,190,657
Oxytricha spp. 403
Oxytricha Steinia 79, 116, 293, 305, 306, 309, 323,
625
Oxytricha Steinia balladynu/a 177
Oxytricha Steinia balladynulla 177
Oxytricha Steinia candens 309
Oxytricha Steinia candens aestuarii 308
Oxytricha Steiniajerruginea 487
Oxytricha Steinia gracilis 305
Oxytricha Steinia granu/ata 306
Oxytricha Steinia inquieta 294
Oxytricha Steinia marina 234, 323
Oxytricha Steinia muscorum 116,279,282,684
Oxytricha Steinia p/atystoma 626
Oxytricha Steinia sapropelica 306
Oxytricha stenocepha/a 822, 933
Oxytricha stephanocepha/a 822
Oxytricha stratiformis 253
Oxytricha strenua 367,368,384
Oxytricha strenuum, Gonostomum 384
Oxytricha striata 253
Oxytricha sty/omuscorum 595
Oxytricha Sty/onychia 79, 116,282,501,511,600
Oxytricha Sty/onychia c1aviformis 505
Oxytricha Sty/onychia curvata 593
Oxytricha Sty/onychiajissiseta 566
Oxytricha Sty/onychia grandis 671
Oxytricha Sty/onychia macrosty/a 600
Oxytricha Sty/onychia muscorum 116, 595, 684
Oxytricha Sty/onychia mytilus 505, 511
Oxytricha Sty/onychia notophora 553
Oxytricha Sty/onychia pusil/a 555
Oxytricha Sty/onychia pustu/ata 565, 566
Oxytricha Sty/onychia putrina 588
Oxytricha Sty/onychia vorax 591, 659
Oxytricha subcy/indrica 253
Oxytricha Tachysoma 79, ll6, 432
Oxytricha Tachysoma echinata 435
Oxytricha Tachysomajurcata 248,457
Oxytricha Tachysoma mirabilis 435
Oxytricha Tachysoma parvisty/a 466
Oxytricha Tachysoma pelionella 435
Oxytricha Tachysoma pellione/Ia 435
Oxytricha Tachysoma rigescens 461
Oxytricha tenella 10, 120, 129, l31T, 191, 221
Oxytricha tennella 191
Oxytricha terrestris 683
Oxytricha terruginea 487
Oxytricha tetracirrata 317
Oxytricha tetranucleata 253
Oxytricha tricirrata 682, 693
Oxytricha tricornis 119,241,602
Oxytricha tricornis, Oxytricha 242
Oxytricha trifallax 254, 683, 684, 685
Oxytricha truncata 10, 120,254
Oxytricha truncata dilatata 254
Oxytricha truncata piriforme 254
Oxytricha tubicola 244
Oxytricha Urosoma 79, ll6, 396,414
Oxytricha Urosoma acuminata 414
Oxytricha Urosoma caudata 400
Oxytricha Urosoma cienkowskii 400
Oxytricha Urosoma gigantea 425
Oxytricha Urosoma longicirrata 163
Oxytricha Urosoma macrosty/a 404, 600
Oxytricha Urosoma stenocephala 822, 933
Oxytricha urosty/a 244, 873
Oxytricha variabilis 122, 170, 190, 193, 246
Oxytricha velox 244
Oxytricha viridis 244, 254
Oxytricha wrzesniowskii 244
Oxytricha, New species of 245, 679
Oxytricha, Oxytricha 79,116
Oxytrichen 261
Oxytrichia 117
Oxytrichia pelione/la 435
Oxytrichia pellionella 435
Oxytrichidae 9, 56, 76, 77, 79, 80, 81, 102, 261,
368,395,492,499,677,766,777,778,781,841,
843,879,884,889,892,893,894,929,930,931,
933
oxytrichids 292, 393, 397, 401, 402, 432, 470, 476,
499,500,607,617,625,698,755,766,775,782,
788,789,790,801,809,815,816,827,828,832,
835, 842, 863, 879, 884, 888, 889, 890, 891, 1193,
894,930,933,935
Oxytrichina 102, 113
Oxytrichinae 1, 8, 14, 20, 74, 75, 76, 77, 78, 80,
104, 106,113, 151,250,303,312,317,320,463,
464,480,483,495,499,625,679,704,717,777,
790,842,893,929
Oxytrichoidea 80,102,766
oxytrichoidea, Holosticha 254
 SYSTEMATIC INDEX
oxytrichoidea, Holostischa 254
Oxytrichum 117
Oxytrocha sp. 117
Parabryophrya 885
paragrandis, Urostyla 844, 848, 870
parahalophila, Opisthotricha 170
parahalophila, Opithotricha 170
parahalophila, Oxytricha 122, 170, 232, 233
Parahistriculus 80, 81, 645, 646
Parahistriculus minimum 658
Parahistriculus minimus 645, 658
Parahistricus minimus 646
paraholophila, Opisthotricha 170
Paraholosticha 88, 663, 827
Parakahliella 32, 73, 607, 843
ParakahJiellidae 78, 102, 500, 502, 571, 632, 777,
827,842,879,892,929,930,931
parakahliellids 607
paralella minor, Oxytricha Opisthotricha 166
parallela minor, Opisthotricha 166
parallela, Opisthotricha 166, 171, 172, 255
parallela, Opisthotricha Oxytricha 171
parallela, Opistotricha 171
parallela, Oxytricha 115, 118, 167,171
parallela, Oxytricha Opisthotricha 171
parallela, Stylonychia 684
parallelis, Opisthotricha 171,255
parallella, Stylonychia 684
Paramaecium histrio 501, 646, 647
parasteinii, Gastrostyla 791, 792, 797
Parastrombidium 494
Parastrombidium pianctonicum 494
Parastrombidium planctonicum, Urosoma = 494
Parastrombidium planctonicum, Urosoma fur 494
Parastylonychia 74, 79, 80, 81,432,500,696
Parastylonychia pleuneouri 697
Parastylonychia plouneouri 4, 106, 112,696,697
Parauroleptus strenuus 753
Paraurostula sp. 841
Paraurostula we issei 841
Paraurostyla 14, 15, 18,26, 56, 68, 71, 73, 74, 78,
79,80,81,103,104,106,476,777,827,841,879
Paraurostyla buitkampi 843, 888
Paraurostyla dispar 842
Paraurostyla eissei 845
Paraurostyla enigmatica 843
Paraurostyla/ossicola 876
Paraurostyla gibba 842
Paraurostylagranulijera 10, 844,874, 876T
Paraurostyla hologama 848
1067
Paraurostyla hymenophora 25, 781, 787, 843
Paraurostyla latissima 842
Paraurostyla macrostoma 843
Paraurostyla marina 842
Paraurostyla naumanni 842
Paraurostyla polymicronucleata 843, 844, 846, 848,
851, 852, 854,855, 856, 859, 863, 868T, 872
Paraurostyla polynucleata 4, 844,877
Paraurostyla pulchra 843
Paraurostyla rubra 842
Paraurostyla terricola 843
Paraurostyla viridis 475,842,843
Paraurostyla weisei 845
Paraurostyla weissei 10,23,25,26,39,49,54, 87,
244,263,268,487,843,844,845,846,848,852,
854,855,856,857,858,859,862,863,864,865,
866,867, 868T, 870,871,873,874,877
Paraurostyla weissei complex Ill, 843, 844, 868T
Parentocirrus 73,104,777,878
Parentocirrus hortualis 4, 10, 113, 781, 878, 879,
883T
Paruroleptus 753
Paruroleptus caudatus 84
Paruroleptus dubium 492
Paruroleptus strenua 753
Paruroleptus strenuus 753, 755
Parurosomal, 18,67,80,81, 114,476,491,607
Parurosoma dubia 492
Parurosoma dubium 11, 105, 106,492
Parurosoma dubium, Holosticha 491, 492
Parurosomagranuli/era 261, 262, 263
Parurosoma tekirghiolica 492
Parurosoma, Hoiosticha 491
Parurostyla weissei 841, 845
parvistyla, Oxytricha 259, 467
parvistyla, Oxytricha Tachysoma 466
parvistyla, Tachysoma 259, 466, 468
parvistylum, Tachysoma 466
parvula, Oxytricha ? 252
parvulum, Tachysoma 432
parvulus, Notocephalus 432
parvum, Gonostomum 369, 393
pastulata, Stylonychia 566
patella, Euplotes 296, 450, 692
Pattersoniella 15,56,103,104,470,717,755,766
Pattersoniella vitiphila 4, Ill, 112, 500, 766, 776T,
884
patula, Stylonychia 603
pediculi/ormis, Stichochaeta 894
pediculi/ormis, Trachelostyla 894
pediculus, Cyclidium 825, 826, 827
 1068 SYSTEMATIC INDEX

pediculus, Kerona 33, 86, 87, 103, 104, 105, 111, piscis, Oxytricha 116
723,741,748,751,826 piscis, Uroleptus 116,450
pediculus, Trichodina 837 pisciuncuiijormis, Oxytricha 252
pelagica, Urosoma 494 pistil/oldes, Oxytricha 252
peiagica, Urotricha 494 Plagiotricha 80, 367, 368
pe/ione//a, Histrio 435 Piagiotricha diana 514
pelione//a, Oxitricha 115,433 Piagiotricha Gonostomum affinis 370
pelione//a, Oxytricha 158,434,435 Piagiotricha strenua 384
pelione//a, Oxytricha Tachysoma 435 Plagiotricha strenua, Oxytricha 395
pe/ione//a, Oxytrichia 435 plana, Oxytricha 175, 178
pe/ione//a, Tachysoma 435 pianctonica, Oxytricha 494
pe/ione//um, Tachysoma 435 planctonica, Urosoma 494
pe/line//a, Oxytricha 450 pianctonicum, Parastrombidium 494
pe//inne//a, Tachysoma 446 pianctonicum, Urosoma = Parastrombidium 494
pe/lionela, Oxytricha 435, 467 pianctonicum, Urosoma 494
pe/lione//a chiiensis, Oxytricha 451 pianctonicum, Urosoma filr Parastrombidium 494
pe/lione//a, Histrio 650 pianctonicum, Viosoma 494
pe/lione//a, Oxytricha 115, 116, 183, 184,433,434, planctonisum, Urosoma 494
467,468 planctontica, Oxytricha 494
pe//ione/la, Oxytricha Tachysoma 435 planctonticum, Pseudostrombidium 8, 14, 86, 106,
pel/ione/la, Oxytrichia 435 112,163,340,486T,494,564
pe/lione//a, Tachiosoma 432 planensis, Pleurotricha 699,715
pel/ione//a, Tachyosoma 432 pianktonicum, Urosoma 494
pel/ione/la, Tachysoma 435,469 piastystoma, Oxytricha 626
pe/lione/la, Tachysoma Oxytricha 434, 468 piatistoma, Oxytricha 650
pe/lione/la, Taxysoma Oxytricha 432 platyitoma, Steinia 626
pe/lione/la, Trichoda 115,431,433 platysoma, Oxytricha 626
pellionellum, Tachysoma 2, 4, 5, 35, 57, 68, 69, 73, piatysoma, Steinia 626
74, 86, 87, 106, 116, 125, 135, 143, 184, 187, platystoma, Oxytricha 116,259,308,309,313,314,
222,238,245,252,253,282,369,375,403,431, 317,624,625,626,629,630,645
432,433,442T,458,459,467,469,737 platystoma, Oxytricha Steinia 626
pellionne/la, Oxytricha 435 platystoma, Steina 625
perione/la, Oxytricha 435 platystoma, Steinia 34,87,280,300,310,625,626,
Perisincirra 397 632,643,645
Perisincirra simiiis 397,419,893 platystoma, Steinia Oxytricha 626
Perisincirra viridis 894 pleuneouri, Parastylonychia 697
perisincirra, Lamtostyia 432 pleuronectes, Oxytricha 252
perisincirra, Tachysoma 432 Pleurosticha 698
Peritromoides 723 Pleurotricha 1, 18,74,75,79,80,81,105,261,262,
Peritromoides simplex 723 432,475,492,500,517,607,620,646,698,816
Peritromus!aurei 839 Pleurotricha A/lotricha 261, 698
Peritromus hydrarum 826, 827, 837 Pieurotricha A/lotricha lanceolata 700
pemix, Oxytricha 244 Pleurotricha dubium 492
pemix, Pseudokeronopsis 244 Pleurotricha echinala 434
perthensis, Urosomoida 4,345,346, 354T, 362 Pleurotricha grandis 1, 86, 87, 476, 698, 699, 709,
petiionel/a, Tachysoma 435 712,716,717
philippinensis, Gastrostyla 791,792,797 Pleurotricha grandis, Pleurotricha 713
phyto!aga, Oxytricha 564 Pleurotricha indica 607,620,623,624,699,816
phytomastigophorean 395 Pleurotricha lanccolala 700
phytophaga, Oxytricha 562 Pleurotricha lanceolata 106, 261, 262, 268, 449,
pirijorme, Oxytricha truncata 254 505,517,698,699, 710T, 716,816
 SYSTEMATIC INDEX 1069

Pleurotricha lanceolatum 700 Ponturostyla enigmatica 843

Pleurotricha macrostoma 699 Postciliodesmatophora 81
Pleurotricha monilata 607 postulata, Stylonichia 566
Pleurotricha mononucleata 492 postulata, Stylonychia 566
Pleurotricha num. 1 716 praeceps, Oxytricha 252
Pleurotricha num. 2 716 praeceps, Trichoda 193,246
Pleurotricha planensis 699, 715 primicirrata, Cyrtohymena 9, 59, 281, 291T, 294,
Pleurotricha Pleurotricha 698 296,300,629,632
Pleurotricha Pleurotricha grandis 713 primicirrata, Steinia 300
Pleurotricha setifera 699,789,816 proboscis, Oxytricha 252
Pleurotricha setifera, Gastrostyla 816 procera, Opisthotricha 220, 255
P1eurotricha sp. 86,268,624,716,717 procera, Oxytricha 10, 121,220
Pleurotricha spec. 262,263,699,701 procera, Oxytricha Opisthotricha 220
Pleurotricha tchadensis 700,701,709 procerus, Pseudouroleptus 888, 889
Pleurotricha tchadiensis 700 Prosopsenus 80, 893
Pleurotricha tihanyensis 782 Prosopsenus sinuatus 893
Pleurotricha tihanyiensis 262 protensa, Oxytricha 252
Pleurotricha variabilis 261, 262, 263, 265, 268, 476, proximata, Oxytricha 121, 191, 196,221,258
699, 782 Psammomitra 79,81,244,778,893
Pleurotricha, P1eurotricha 698 Psammomitra retractilis 243
Pleurotrichen 261 pseudofurcata, Oxytricha 4, 119,173,457
Pleurotrichidae 80, 102 pseudofusij'ormis, Oxytricha 4,119, 131T, 222
Pleurotrichina 102 pseudograndis, Stylonychia 503, 564, 580, 594
Pleurotrichinae 102 Pseudohypotrichina 32
pleurotrichs 262 Pseudokeronopsis 491
Pleurotrychaflexilis 478 Pseudokeronopsis camea 243
Pleurtricha 698 Pseudokeronopsisflava 243
plicata, Oxytricha 252 Pseudokeronopsis pemix 244
plouneouri, Parastylonychia 4, 106, 112, 696, 697 Pseudokeronopsis rubra 23, 26, 244, 282, 401
Pnychodromus grandis simplex 724 pseudosimilis, Oxytricha 10,34, 118, 131T, 222
Podophrya collini 584 Pseudostrombidium 80, 114,494
Podophryafixa 81 Pseudostrombidium planctonticum 8, 14, 86, 106,
Podophrya grelli 81,82,84,551 112, 163, 340, 486T, 494, 564
Podophrya sol 81 Pseudouroleptus 104,777,843,888,894
Podophrya stylonychiae 81, 84, 85 Pseudouroleptus buitkampi 843, 888, 889
polycirratus, Histriculus 646, 666 Pseudouroieptus caudatus 105, 888, 889
polyci"atus, Histrio 666 Pseudouroleptus humicola 888, 889
Polyhymenophora 80 Pseudouroleptus procerus 888, 889
polymicronucleata, Paraurostyla 843, 844, 846, Pseudouroleptus terrestris 888, 889
848,851,852,854,855,856, 859,863, 868T,872 Pseudourostyla cristata 23, 26, 865
polymicronucleata, Urostyla 845,848,870 Pseudourostyla laevis 39
polymorphum, Kerona 826 Pseudourostyla urostyla 244
polynucleata, Hemisinci"a 36, 397, 419, 893 Psilotricha 104,499,598,894
polynucleata, Paraurostyla 4, 844, 877 Psilotricha acuminata 894
polynucleata, Urosoma 4, 11, 18,35,396,397,402, pulchra, Gastrostyla 9, 790, 791, 818
405T,411,417,419, 421 pulchra, Gastrostyla Stylonychia 818, 824
polypora, Kerona 826 pulchra, Paraurostyla 843
polyporum, Alaster 825 pulchra, Stilonichia 818
polyporum, Alastor 825, 826 pulchra, Stylonychia 818
polyporum, Kerona 825, 826, 827, 837, 841 pulex, Oxitricha 115
Ponturostyla 843 pulex, Trichoda 115,246,566
 1070 SYSTEMATIC INDEX

pullaster, Holosticha 116, 242, 243, 252, 435, quadrinucleata, Cyrtohymena 2, 4, 56, 60, 207,
444 279, 280, 291T, 317, 320, 500, 677
pullaster, Oxitricha 116 quadrinucleata, Oxytricha 252
pullaster, Oxytricha 116 quadrinucleata, Steinia 320
pullaster, Trichoda 116 quadrinucleata, Stylonychia 677, 678, 679
pullicina, Oxitricha 115, 246 quadrinucleatus, Histrio 646, 665, 694
Puraropeltus strenuus 753 quadrinucleatus, Steinia 320
pusilla, Oxytricha Stylonychia 555 quadrinucleatus, Sterkiella 4, 665, 666, 669, 677,
pusilla, Sphaerophrya 81 694, 792
pusilla, Stylonychia 555, 594 quedricomutus, Onychodromus 740
pusilla, Stylonychia mytilus 555 quercineti, Oxytricha 252
pusilus, Stylonychia 555
pussila, Stylonychia 549, 555 radians, Cyclidium 566
pustalata, Stylonychia 566 radians, Oxytricha 253
pustufata, Cerona 565 radiosa, Mitra 893
pustulata, Clara 565, 929 raptans, Lamtostyla 432, 893
pustulata, Kerona 502, 565, 600, 825 raptans, Tachysoma 432
pustulata, Kerone 601 rastellum, Kerona 840
pustulata, Oxytricha 259 reclinis, Stylonichia 599
pustulata, Oxytricha Stylonychia 565, 566 regularis, Stylonichia 600
pustulata, Stylometra 502 regularis, Stylonychia 600
pustulata, Stylonichia 565, 566, 598, 601 retractilis, Oxytricha 244
pustulata, Stylonychia 6, 20, 35, 43, 49, 59, 63, 64, retractilis, Psammomitra 243
68, 69, 75, 76, 77, 78, 87, 205, 245, 250, 259, retractilis, Uroleptus 244
296, 403, 446, 502, 503, 517, 521, 530, 534T, rigescens, Oxytricha Tachysoma 461
553,554,557,560,561,564,565,586,587,588, rigescens, Tachysoma 433,461
589,590,591,593,598,602,603,604,667,681, rigescens, Tachysoma Oxytricha 461
685,723,731,778,781,833,835,929 Rigidocortex470, 500,503,717
pustulata, Stylonychia Oxytricha 565 Rigidocortex octonucleatus 2, 4, 106, 113, 676, 718
pustulata, Stylonychnia 502 rostrata, Kerona 515,840
pustulatus, Stylonychia 566 rostrata, Oxytricha 253
pustulatus, Stylonychnia 566 rostrata, Stylonichia 599
pustullata, Stylonychia 603 rostrata, Stylonychia 599
pustutata, Stylonychia 566 rostrata, Trichoda 515
pusulata, Stylonychia 566 rotunda, Kerona 839
putrina, Oxytricha Stylonychia 588 rotundata, Kerona 841
putrina, Stylonichia 602 rubescens, Notohymena 9, 33, 59, 63, 64,106,224,
putrina, Stylonychia 35, 87, 502, 503, 505, 506, 324,325, 339T,490
517,546,553,555,565,568,571,573,584,586, rubipuncta, Oxytricha 224
591,592,593,604 rubra, Oxytricha 244, 281, 282, 293, 344, 401
putrinum, Sty/onychia 588 rubra, Paraurostyla 842
pyriformis complex, Tetrahymena 510 rubra, Pseudokeronopsis 23,26,244,282,401
rubra, Urostyla 842
quadricirrata, Oxytricha 10, 121, BIT, 197, 223, Rubrioxytricha 106, 113, 114, 118, 120, 279, 293,
684 325,479
quadricirrata, Oxytricha granulifera 197, 223 Rubrioxytricha /erruginea 11, 87, 123, 479, 480,
quadricornatus, Onychodromus 740 481, 484, 486T,487, 491
quadricornutis, Onychodromus 740 Rubrioxytricha haematoplasma 8, II, 34, 87, 106,
quadricornutus, Onychodromous 722 123, 125, 138,480,481, 486T,488
quadricornutus, Onychodromus 4, 18, 75, 103, 578, rubripuncta, Oxytricha 10,34,121,123, l31T, 194,
723,724,736,738, 750T, 761, 773,827,833,884 224,325
 SYSTEMATIC INDEX 1071

salmastra, Oxytricha 424 similis, Oxytricha Opisthotricha 183,665,694

sa/mastra, Urosoma 396, 398, 405T, 422, 424 similis, Perisincirra 397, 419,893
sa/tans, Actinotricha 115,236, 253, 255 similis, Stylonychia 665
saitans, Oxytricha 122, l3iT, 236, 253, 931 similis, Urosoma 18,245,396,398, 405T, 419
sa/tans, Oxytricha Actinotricha 236 simplex, Cyrtohymena 309
sa/tans, Tachysoma 236, 253 simplex, Kahliella 931
sa/tans, Tachysoma Actinotricha 236 simplex, Onychodromus grandis 724
sannio, Himantopus 515 simplex, Peritromoides 723
sannio, Kerona 515 simplex, Pnychodromus grandis 724
sannio, Trichoda 515 simplex, Steinia 308, 309, 312, 313
saprobia, Oxytricha 86, 87,120,174,259 sinuatus, Prosopsenus 893
saprobica, Oxytricha 174 siseris, Oxytricha 62,104, li8, l31T, 187
saprope/ica Oxytricha Steinia 306 siseris, Tachysoma 187
saprope/ica, Cyrtohymena 4, 157, 280, 281, 295, sol, Podophrya 81
306 sordida, Oxytricha 123, 125,260
sapropelica, Steinia 306 sordis, Opisthotricha 245
sardida, Oxytricha 260 sordis, Oxytricha 245
scutellum, Oxytricha 244 sp. ?, Stylonychia 504
scutum, Diophrys 44 sp.flexilis, Onychodromopsis 479
scutum, Oxytricha 557 sp. n. sp., Stylonychia 605
se/vatica, Notohymena 324, 325, 339T, 344 sp. No.1, Ciliate 683
selvatica, Oxytricha 344 sp. nov.?, Histriculus 667
semen, Gonyostomum 395 sp. nov?, Urostyla 844
setifera, Gasterosty/a 816 sp. Paraurostula 841
setifera, Gastrostyla 790, 816, 818, 824 sp., Ciliate 367
setifera, Gastrosty/a Pleurotricha 816 sp., Cyrtohymena 4,281,307,310,317,631
setifera, Nothopleurotricha 816 sp., Gastrostyla 824
setifera, Oxytricha 177, 816 sp., Gonostomum 395
setifera, Pleurotricha 699, 789, 816 sp., Historio 666
setigera, Holosticha 897 sp., Histriculus 33
setigera, Oxytricha 1,2,4,5,17,34,57,87, ll7, sp., Histrio 667
li8, 128, l3lT, 173, 175, 185, 187, 213, 217, sp., Ocytricha 117
447,457,659,81~ 897 sp., Onychodromopsis 475, 476
silurus congyrans, Kerona 839 sp., Onychodronum 475
silurus, Kerona 504, 566, 600, 601 sp., Opisthotricha 679, 682
silurus, Stylonychia 504, 517 sp., Oxitrica 117
silurus, Trichoda 504, 566 sp., Oxitrycha ll7
simi/is trici"atus, Histriculus 36, 646, 682 sp., Oxytricha 20, 26, 45, 86, 122, l3iT, 190,251,
similis, Balladyna 126, 128 260,679,682
simi/is, Hemisincirra 419 sp., Oxytrocha 117
similis, Histriculus 25,36,568,647,660,663,665, sp., Pleurotricha 86, 268, 624, 716, 717
666,667,682,683,684,693,694,792 sp., Stylonichia 602
similis, Histrio 665, 694, 696 sp., Stylonychia 28, 86, 503, 505, 604, 666
similis, Holosticha 126, 128, 930 sp., Stylonychia ? 605
similis, Opisthotricha 169, 183,255 sp., Stylonyehia 502
similis, Opisthotricha Oxytricha 183 sp., Styx 738
similis, Opistotricha 117 sp., Tachysoma 469
similis, Opistrotricha 117 sp., Urosoma 23,431
similis, Oxytricha 87, 121, 128, l36, 159, 169, 183, sp., Vorticella 84
220,222,260,436 spec., Oxitricha 117
simi/is, Oxytricha Histrio 183, 665, 694 spec., Pleurotricha 262, 263,699,701
 1072 SYSTEMATIC INDEX

spec., Stylonychia 505, 517 Steinia citrina 293

spec., Tachysoma 432 Steinia dubia 281, 282, 293
sphaerica, Stylonychia 600 Steiniajenestrata 644
sphaericus, Balladinopsis 738 Steiniajerruginea 487,625
Sphaerophrya pusilla 81 Steiniajerruginea, Oxytricha 487
sphagni, Histriculus 647,650,655,658 Steinia gracilis 305
sphagni, Histrio 654 Steinia gracilis, Oxytricha 305
sphagni, Opisthotricha 190 Steinia granulata 306
sphagni, Oxytricha 4, 119, 190, 657 Steinia granulata, Oxytricha 306
sphagni, Oxytricha Histrio 190, 657 Steinia Histrio inquieta 293
sphagni, Oxytricha Opisthotricha 190, 657 Steinia inqueta 315
sphagni, Trachelochaeta 894 Steinia inquieta 281, 282, 300, 303, 314, 324
sphagnicola, Cyrtohymena 632 Steinia inquieta, Oxytricha 294
sphagnicola, Steinia 18,44, 59, 61, 63, 64, 67, 106, Steinia macrostoma 314,315
280, 625,629, 632, 643, 644T, 705 Steinia marina 323
Spirostomum ambiguum 116, 246 Steinia marina, Oxytricha 234, 323
Spirotrich 506 Steinia muscorum 281,282
Spirotricha 79, 80 Steinia muscorum, Oxytricha 116,279,282,684
Spirotrichea 81 Steinia ovalis 314, 317
spirotrichoides, Gonostomum 370,375 Steinia Oxytrichajerruginea 487
spirotrichoides, Trachelostyla 370 Steinia Oxytricha platystoma 626
Sporadotrichina 80,81,261 Steinia platyitoma 626
spp., Allotricha 782 Steinia platysoma 626
spp., Gastrostyla 790 Steinia platystoma 34, 87, 280, 300, 310, 625, 626,
spp., Gonostomum 500 632, 643, 645
spp., Onychodromus 500, 755, 769 Steinia platystoma, Oxytricha 626
spp., Oxytricha 403 Steinia primicirrata 300
spp., Stylonychia 495 Steinia quadrinucleata 320
spp., Urosomoida 18, 159 Steinia quadrinucleatus 320
St. sylurus 511 Steinia sapropelica 306
Steina 625 Steinia saprope/ica Oxytricha 306
Steina platystoma 625 Steinia simplex 308,309,312,313
steini, Gasterostyla 789 Steinia sphagnicola 18, 44, 59, 61, 63, 64, 67, 106,
steini, Gastostyla 789 280, 625,629,632, 643, 644T, 705
steini, Gastrostyla 791, 792, 824 Steinia tetracirrata 317
steini, Histrio 650 Steinia torrenticola 307
steini, Stylonychia 605 Steinia ultricirrata 300
Steinia 17,20, 56, 61, 62, 66, 67, 68, 70, 73, 75, 76, Steinia, Oxytricha 79, 116, 293, 305, 306, 309, 323,
77,80, 103, 112, 116, 177, 178,248,279,280, 625
305,306,309,314,316,323,480,487,502,571, stein ii, Gasterostyla 789
605,624 steinii, Gastrostyla 1, 2, 4, 6, 19,25,26,33, 54, 87,
Steinia balladina 177 88, 111, 646, 694, 789, 790,791, 802T,808, 817
Steinia balladynula 175, 177 stein;;, Histrio 605, 645, 646, 647
Steinia balladynula, Oxytricha 177 stein ii, Stylonychia 506, 517, 605
Steinia balladynulla, Oxytricha 177 steinni, Gastrostyla 792
Steinia bivacuolata 625,641 stenocephala, Gastrostyla 9,789,791,822
Steinia candens 305, 307,308,309,323 stenocephala, Gastrosyla 789
Steinia candens aestuarii, Oxytricha 308 stenocephala, Hemigastrostyla 933
Steinia candens depressa 314,315,316 stenocephala, Oxytricha 822, 933
Steinia candens, Oxytricha 309 stenocephala, Oxytricha Urosoma 822, 933
Steinia caudens 324 stenocephala, Gastrotyla 789
 SYSTEMATIC INDEX 1073

stenuum, Gonostomum 384 strenuum, Gonostomum Oxytricha 384

stenuum, Gonostomun 395 strenuus, Onychodromus 755
stephanocephala, Oxytricha 822 strenuus, Parauroleptus 753
SterkieHa 14, 17,56,61,66,67,71,73,75,76,77, strenuus, Paruroleptus 753,755
106,113,117,168,503,617,646,650,662,665, strenuus, Puraropeltus 753
668,771,930 striata, Oxytricha 253
Sterkiella admirabilis 4,669,671,677 Strombidium 23
Sterkiella cavicola 4, 16, 18,20,22,23,34,46, 57, stueberi, Holosticha 470
60,64,71,168, 169,207, 667,668,669, 674T Stylochia 501
SterkieHa histriomuscorum 16,26,28,30,31, 34, Styloichia mitilus 511
59,64,71,86,87, 106, 117, 118, 121, 122, 125, Stylometra pustulata 502
151, 152, 168, 196,250,254,257,468,578,583, stylomuscorum, Oxytricha 595
593, 603, 604, 646, 658, 665, 666, 668, 669, stylomuscorum, Sty/onychia 4, 86, 87, 503, 594,
674T, 679, 681, 682, 683, 693, 695, 716, 723, 595,604
927T,929 Stylomychia 502
Sterkiella histriomuscorum complex 668, 669 Stylonchia 502
Sterkiella quadrinucleatus 4, 665, 666, 669, 677, Stylonchia curvata 502
694, 792 Stylonchia lemnae 502
Sterkiella terricola 4,34,669, 674T, 682, 690, 693 Stylonchia mytilus 502, 895
Sterkiella thompsoni 4, 113, 245, 668, 669, 674T, Stylonichea bistyllus 597
679 Stylonichia 502
Sterkiella tricirrata 668, 669, 674T, 693 Stylonichia anfracta 597
sterkii, Gastrostyla 818,820,821 Stylonichia auriformis 597
stemuum, Gonostomum 384 Stylonichia bicaudatus 597
Stichochaeta 894 Stylonichia capucinus 597
Stichochaeta affinis 370 Stylonichia comiculata 597
Stichochaeta pediculiformis 894 Stylonichia comifrons 597
Stichochaeta strenua 384 Stylonichia curvata 593
Stichotricha 894 Stylonichia dupla 598
Stichotricha tubicola 244 Stylonichia ellipsofdes 598
Stichotrichida 81 Stylonichia excavata 598
Stichotrichina 80, 81 Stylonichiajastigiata 598
Stienia macrostoma 314 Stylonichia fronte-albens 598
Stilonichia 501 Stylonichiafrontosus 598
Stilonichia pulchra 818 Stylonichia globifrons 598
Stilonychia 501 Stylonichia gyriniformis 598
Stilonychia mytilus 501 Stylonichia histrio 646, 650
stramenticola Territricha 11, 111, 112, 676, 694, Stylonichia histrix 650
720, 769,773,884, 887T Stylonichia hystrio 650
stratiformis, Oxytricha 253 Stylonichia labiata 598
strenua, Gonostomum 384 Stylonichia limbiformis 598
strenua, Laurentiella 4, 105, 106, 500, 741, 753, Stylonichia limbifrons 598
764T, 769, 773 Stylonichia macrostyla 600
strenua, Oxytricha 367, 368, 384 Stylonichia mamillata 598
strenua, Oxytricha Plagiotricha 395 Stylonichia massula 598
strenua, Paruroleptus 753 Stylonichia membranaceus 599
strenua, Plagiotricha 384 Stylonichia mitilus 511
strenua, Stichochaeta 384 Stylonichia muscorum 601
strenum, Gonostomum 384 Stylonichia mutabunda 599
strenuum, Gonostomum 9,33,242,368,369,370, Stylonichia mutilus 511
373, 382T,384, 395 Stylonichia mytilis 505
1074 SYSTEMATIC INDEX

Stylonichia mytillus 505, 511 Stylonychia grandis, Oxytricha 671

Stylonichia mytilus 504,505,506,565,601 Stylonychia harbinensis 599
Stylonichia mytylus 511 Sty/onychia histrie 650
Stylonichia notophora 553, 601 Stylonychia histrio 501, 647, 668
Stylonichia num. 1 599 Stylonychia kahli 521, 549
Stylonichia octinostylus 599 Stylonychia lanceolata 698, 699, 717
Stylonichia postulata 566 Stylonychia lemnae 5, 6, 12, 37, 82, 84, 205, 502,
Stylonichia pustulata 565, 566, 598, 601 503, 506, 509, 510, 511, 513, 521, 524, 525, 526,
Stylonichia putrina 602 529, 533, 534T, 548, 549,550, 676, 752
Stylonichia reclinis 599 Stylonychia macrostyla 600
Stylonichia regularis 600 Stylonychia macrostyla, Oxytricha 600
Stylonichia rostrata 599 Stylonychia makrosty/a 600
Stylonichia sp. 602 Stylonychia monosty/us 600
Stylonichia tricomis 242, 602 Sty/onychia muscorum 595, 602
Stylonichia trochijormis 599 Stylonychia muscorum, Oxytricha 116, 595, 684
Stylonichia truncata 599 Stylonychia mutilis 511
Stylonichia unguiculus 599 Stylonychia mutilus 511
Stylonichia virgula 549 Stylonychia mysilus 511
Stylonichia l'iridis 599 Stylonychia mystilus 511
Stylonichia vorax 591 Stylonychia mythilus 504, 505
Sty/onichnia mytilus 502 Stylonychia mytilis 511
Sty/onichus mytilus 502 Stylonychia mytillus 513
Stylonycha 502 Sty/onychia mytiluns 513
Stylonychi lemnae 502 Stylonychia mytilus 2, 4, 17, 18, 20, 21, 23, 25, 26,
Stylonychia ? sp. 605 34,44,49, 51, 52, 54, 55, 64, 66, 67, 68, 70, 75,
Stylonychia 1, 8, 14,20,24,26,43,49, 52, 54, 56, 76,77,78,84,248,251,256,501,502,503,504,
62, 71, 73, 75, 79, 80, 81, 103, 106, 112, 113, 505,506,509,510,511,513,514,517,519,521,
117,142,151,152,230,244,248,251,260,261, 524, 525, 526, 534T, 537, 544, 549, 550, 551,
282,495,499,501,511,560,571,597,617,646, 552,553,555,560,571,581,592,597,598,599,
650,668,671,728,741,755,778,818,825,827, 600, 602, 603, 606, 625, 687, 737, 755, 757, 870,
839,840,893,930 929
Stylonychia appendiculata 596 Stylonychia mytilus-Komplex 506
Sty/onychia aurijormis 549 Stylonychia mytilus complex I, 2, 6, 8, 14, 35, 39,
Sty/onychia hi/aria 34, 64, 67, 68, 69, 75, 76, 77, 40,41,42,43, 71, 81, 85, 87, 88, 106, 143,248,
78,502,503, 534T, 553, 556, 557, 571, 577,601, 403,450,502,503,504,549,550,551,554,555,
602,685,731 562,568,577,588,592,598,599,600,601,602,
Sty/onychia calva 599 603,604,607,611,617,640,704,755,759,761
Stylonychia cimex 501 Stylonychia mytilus pusilla 555
Stylonychia clavata 511 Stylonychia mytilus syngen 1 506
Sty/onychiaclavijormis 505, 511, 517, 537 Stylonychia mytilus syngen 2 510
Sty/onychia clavijormis, Oxytricha 505 Stylonychia mytilus Varietllt 1 505,550
Sty/onychia curvata 502, 503, 568, 588, 591, 593, Stylonychia mytilus Varietllt 2 505, 549
683 Stylonychia mytilus, Oxytricha 505, 511
Stylonychia curvata, Oxytricha 593 Stylonychia mytylus 513
Stylonychia dupla 598 Stylonychia n. sp. 605
Stylonychia echinata 431,434,444 Stylonychia nodulinucleata 2, 4, 503,552
Stylonychiajastigiata 598 Sty/onychia notophora 36, 205, 503, 525, 530, 549,
Stylonychiajissieta 505 553,562,565,568,578,588,592,594,603
Sty/onychiajissiseta 565, 566, 580 Stylonychia notophora, OxytriCha 553
Stylonychiajissiseta, Oxytricha 566 Stylonychia notophra 554
Stylonychia grandis 671, 676, 697 Stylonychia notrophora 553
 SYSTEMATIC INDEX 1075

Stylonychia Onychodromus grandis 669 Stylonychia vorax 36, 76, 86, 87, 502, 503, 557,
Stylonychia ovale 593 562,565,568,571,588,591,593,604
Stylonychia ovalis 591 Stylonychia vorax, Oxytricha 591, 659
Stylonychia Oxytricha pustulata 565 Stylonychia, Oxytricha 79,116,282,501,511,600
Stylonychia parallela 684 stylonychiae, Podophrya 81, 84, 85
Stylonychia parallella 684 stylonychias 893
Stylonychia pastulata 566 Stylonychinae 1, 8, 9, 14, 20, 74, 75, 76, 77, 78,
Stylonychia patula 603 106, 111, 114, 168,250,314,317,320,321,470,
Stylonychia postulata 566 495,499,638,679,697,717,723,736,766,771,
Stylonychia pseudograndis 503, 564, 580, 594 773,790,827,834,894,929
Stylonychia pulchra 818 Stylonychna mytilus 502
Stylonychia pulchra, Gastrostyla 818, 824 Stylonychnia histria 650
Stylonychia pusilla 555, 594 Stylonychnia histrio 650
Stylonychiapusilla, Oxytricha 555 Stylonychnia mytilus 502
Stylonychia pusilus 555 Stylonychnia pustulata 502
Stylonychia pussila 549, 555 Stylonychnia pustulatus 566
Stylonychia pustalata 566 Stylonyehia sp. 502
Stylonychia pustulata 6, 20, 35, 43, 49, 59, 63, 64, Styloychia mytilus 502
68, 69, 75, 76, 77, 78, 87, 205, 245, 250, 259, Stynolychia monostylus 502
296, 403, 446, 502, 503, 517, 521, 530, 534T, Stynolychia virgula 548, 597
553,554,557,560,561,564,565,586,587,588, Styxsp.738
589,590,591,593,598,602,603,604,667,681, subcylindrica, Oxytricha 253
685,723,731,778,781,833,835,929 subrotondata, Kerona 840
Stylonychia pustulata, Oxytricha 565, 566 subrotundata, Kerona 840
Stylonychia pustulatus 566 suctorians 81, 517, 584
Stylonychia pustullata 603 sylurus, St. 511
Stylonychia pustutata 566 sylurus, Stylonychia 511
Stylonychia pusulata 566 syngen 1, Stylonychia mytilus 506
Stylonychia putrina 35, 87, 502, 503, 505, 506, syngen 2, Stylonychia mytilus 510
517,546,553,555,565,568,571,573,584,586,
591,592,593,604 Tachiosoma pellionella 432
Stylonychia putrina, Oxytricha 588 tachyblastum, Trachelophyllum 631
Stylonychia putrinum 588 Tachyosoma pellionella 432
Stylonychia quadrinucleata 677, 678, 679 Tachysoma 1, 14, 17,20,56,61,66,67,68,73,74,
Stylonychia regularis 600 76, 77, 78, 79, 80, 81, 103, 106, 112, 116, 166,
Stylonychia rostrata 599 236,250,253,259,282,375,395,431,435,456,
Stylonychia silurus 504, 517 457,458,893
Stylonychia similis 665 Tachysoma Actinotricha saltans 236
Sty[onychia sp. ? 504 Tachysoma agile 434, 444
Stylonychia sp. 28, 86, 503, 505, 604, 666 Tachysoma agilis 434
Stylonychia sp. n. sp. 605 Tachysoma balatonica 433, 454, 456, 457
Stylonychia spec. 505, 517 Tachysoma bicirratum 4, 86, 87, 173, 433, 452,
Stylonychia sphaerica 600 454,456,457
Stylonychia spp. 495 Tachysoma chilensis 432, 451
Stylonychia steini 605 Tachysoma dragescoi 11,432,433, 442T, 463
Stylonychia steinii 506,517,605 Tachysoma dubia 281
Stylonychia stylomuscorum 4, 86, 87, 503, 594, Tachysoma echinata 434
595,604 Tachysoma echinata, Oxytricha 435
Stylonychia sylurus 511 Tachysomafurcata 173,457
Stylonychia trochiformis 599 Tachysomafurcata, Oxytricha 248, 457
Stylonychia virgula 548 Tachysomafurcatum 457
1076 SYSTEMATIC INDEX
Tachysomafusiformis 175, 178
Tachysoma granulifera 459
Tachysomagranuliferum 11,432,433, 442T, 459
Tachysoma humicola 4, 253, 432, 433, 452, 456,
458
Tachysoma humicola humicola 433, 442T, 452,
454,455,456
Tachysoma humicola longisetum 433, 442T, 452,
455,456
Tachysoma hyalina 432
Tachysoma longa 432
Tachysoma mirabile 434, 444
Tachysoma mirabilis 434
Tachysoma mirabilis, Oxytricha 435
Tachysoma monstrosa 250
Tachysoma ovata 4, 11,432,433, 442T, 464
Tachysoma Oxytricha pellionella 434, 468
Tachysoma Oxytricha rigescens 461
Tachysoma parvistyla 259,466,468
Tachysoma parvistyla, Oxytricha 466
Tachysoma parvistylum 466
Tachysoma parvulum 432
Tachysoma pelionella 435
Tachysoma pelionella, Oxytricha 435
Tachysoma pelionellum 435
Tachysoma pellinnella 446
Tachysoma pellionella 435, 469
Tachysoma pellionella, Oxytricha 435
Tachysoma pellionellum 2, 4, 5, 35, 57, 68, 69, 73,
74, 86, 87, 106, 116, 125, 135, 143, 184, 187,
222,238,245,252,253,282,369,375,403,431,
432,433, 442T,458,459, 467,469, 737
Tachysoma perisincirra 432
Tachysoma petlionella 435
Tachysoma raptans 432
Tachysoma rigescens 433, 461
Tachysoma rigescens, Oxytricha 461
Tachysoma saltans 236, 253
Tachysoma siseris 187
Tachysoma sp. 469
Tachysoma spec. 432
Tachysoma terricola 35, 68, 73, 74,432, 433, 442T,
454,458
Tachysoma, Oxytricha 79, 116,432
Taxysoma Oxytricha pellionella 432
tchadensis, Pleurotricha 700, 701, 709
tchadiensis, Pleurotricha 700
tekirghiolica, Leptocirra 492
tekirghiolica, Parurosoma 492
tenella, Oxytricha 10, 120, 129, 131 T, 191, 221
tennel/a, Oxytricha 191
tericola, Opisthotricha 245
terrestris, Opisthotricha 683, 684
terrestris, Opistotricha 683, 684, 689
terrestris, Oxytricha 683
terrestris, Pseudouroleptus 888, 889
Terricirra 79, 885, 894
terricola, Opistotricha 245
terricola, Paraurostyla 843
terricola, Sterkiella 4,34,669, 674T, 682, 690, 693
terricola, Tachysoma 35, 68, 73, 74, 432, 433,
442T,454,458
Territricha 15, 18, 32, 56, 74, 78, 103, 104, 470,
777,884
Territricha stramenticola 11, 111, 112, 676, 694,
720, 769, 773,884,887T
terruginea, Oxytricha 487
tetracirrata, Cyrtohymena 280, 281, 291T, 308,
317,320,643
tetracirrata, Oxytricha 317
tetracirrata, Steinia 317
Tetrahymena pyriformiS complex 510
tetraculeatus, Tricholeptus aculeatus 894
tetranucieata, Oxytricha 253
thompsoni, Sterkiella 4, 113, 245, 668, 669, 674T,
679
tihanyensis, Pleurotricha 782
tihanyiensis, Apoamphisiella 11, 781, 782, 786T,
787, 788
tihanyiensis, Onychodromopsis 476,781,782,843
tihanyiensis, Pleurotricha 262
torrenticola, Cyrtohymena 280,281,295,307
torrenticola, Steinia 307
Trachelochaeta 79, 242, 368, 894
Trachelochaeta bryophila 894
Trachelochaeta gonostomoida 392
Trachelochaeta sphagni 894
Trachelophyl/um tachyblastum 631
Trachelostyla 79, 80, 81,103,242,368,894
Trachelostyla affine 370
Trachelostyla bryonicolum 370
Trachelostyla canadensis 370
Trachelostyla ciliophorum 370
Trachelostyla geleii 370
Trachelostyla macrostoma 370
Trachelostyla pediculiformis 894
Trachelostyla spirotrichoides 370
Trachelostylidae 81, 103
transfuga, Oxitricha 116
transfuga, Trichoda 116
transfuga, Uronychia 116, 580
tremula, Kerona 841
 SYSTEMATIC INDEX
triangularis, Kerona 841
triccirratus, Histriculus 682
Trichoda acarus 514, 566
Trichoda ambigua 116,246
Trichoda bulla 116
Trichoda calvitium 566, 700
Trichoda cicada 254
Trichoda cimex 501
Trichoda cyclidium 566
Trichoda cypris 514
Trichoda erosa 514
Trichoda jelis 116
Trichodafimbriata 248,514
Trichoda gallina 116
Trichoda gibba 116, 842
Trichoda histrio 647
Trichoda lepus 115
Trichoda ludio 514
Trichoda mytilus 501,504,549
Trichoda pellione/la 115,431,433
Trichodapraeceps 193,246
Trichoda pulex 115, 246, 566
Trichoda pu/laster 116
Trichoda rostrata 515
Trichoda sannio 515
Trichoda silurus 504, 566
Trichoda transfuga 116
Trichodina pediculus 837
trichogaster multipes, Urostyla 873
Tricholeptus 80, 397, 894
Tricholeptus aculeatus 397, 894
Tricholeptus aculeatus tetraculeatus 894
Trichotaxis 243, 244
Trichotaxis caudata 398
Trichotaxis crassa, Holosticha 243
Trichotaxisjossicola 848, 874, 875
Trichotaxisjossicola, Holosticha 876
Trichotaxis Oxytricha crassa 243
Trichotaxis, Holosticha 116, 243, 876
Trichototaxis 243,244,398,401,874,876
tricirrata, Oxytricha 682, 693
tricirrata, Sterkiella 668, 669, 674T, 693
tricirratus, Histriculus 646, 682
tricirratus, Histriculus similis 36, 646, 682
tricornis, Oxytricha 119,241,602
tricornis, Oxytricha Oxytricha 242
tricorn is, Stylonichia 242, 602
trija/lax, Oxytricha 254, 683, 684, 685
tristyla, Dipleurostyla 242, 893
trochijormis, Stylonichia 599
trochijormis, Stylonychia 599
1077
truncata dilatata, Oxytricha 254
truncata pirijorme, Oxytricha 254
truncata, Kerona 841
truncata, Oxytricha 10, 120,254
truncata, Stylonichia 599
tubicola, Oxytricha 244
tubicola, Stichotricha 244
turrita, Aspidisca 723
ultricirrata, Steinia 300
unguicularis, Kerona 841
unguiculus, Stylonichia 599
Uroleptoides 894
Uroleptus 243,248,347,348,367,401,430,893
Uroleptus agilis 345, 346, 347, 348, 367
Uroleptus caudatus 23
Uroleptus ellipticus 787
Uroleptus gallina 116
Uroleptus humicola 888
Uroleptus kahli 421,892
Uroleptus mobilis 893
Uroleptus piscis 116,450
Uroleptus retractilis 244
Uronychia transfuga 116,580
Urosoma = Parastrombidium planctonicum 494
Urosoma 1, 11, 15, 17, 20, 32, 56, 57, 62, 66, 67,
68, 69, 73, 74, 75, 76, 77, 78, 80, 81, 103, 112,
114, 118, 163,230,246,307,346,347,369,375,
395,396,407,414,419,424,432,476,491,492,
493,600,679,893
Urosoma acuminata 4, 11, 396, 397, 401, 405T,
414,495
Urosoma acuminata, Oxytricha 414
Urosoma acuta 404,407,409
Urosoma ambigua 404, 407, 409
Urosoma caudata 11, 106, 396, 398, 405T, 411,
414,691
Urosoma caudata, Oxytricha 400
Urosoma caudatum 400
Urosoma cienckowski 401
Urosoma cienkowski 398,431
Urosoma cienkowskii 396, 398, 400, 401, 403
Urosoma cienkowskii, Oxytricha 400
Urosoma cienkowsky 401
Urosoma cienkowskyi 398
Urosoma cinekowski 398
Urosoma emarginata 35, 163, 369, 396, 397, 402,
405T,407,409,414
Urosoma fOr Oxytricha gigantea 425
Urosoma fOr Parastrombidium planctonicum 494
Urosoma gigantea 11,35, 164,396,398, 405T, 425
1078 SYSTEMATIC INDEX

Urosoma gigantea, Oxytricha 425 Urostyla grandis 23, 243, 487,841,848,870

Urosoma karini 11,396,398, 405T, 422 Urostyla hologama 845, 848, 855, 870
Urosoma longicirrata 163 Urostyla hologamma 845
Urosoma longicirrata, Oxytricha 163 Urostyla latissima 842
Urosoma macrostoma 370, 373 Urostyla lynchi 845, 848, 855, 870
Urosoma macrostya 404 Urostyla marina 842
Urosoma macrostyla 36, 125, 398, 404, 405T, 409, Urostyla multipes 873
931 Urostyla naumanni 842
Urosoma macrostyla, Oxytricha 404, 600 Urostyla Oxytricha multipes 873
Urosoma octonuc/eata 2,4,396,397, 405T, 417 Urostyla paragrandis 844, 848, 870
Urosoma Oxytricha acuminata 414 Urostyla polymicronucleata 845, 848, 870
Urosoma Oxytricha caudata 398 Urostyla rubra 842
Urosoma Oxytricha macrostyla 404, 409 Urostyla sp. nov? 844
Urosoma pelagica 494 Urostyla trichogaster multipes 873
Urosoma planctonica 494 Urostyla vernalis 844, 845, 846, 848
Urosoma planctonicum 494 Urostyla viridis 843, 870
Urosoma planctonisum 494 Urostyla weissei 841,844,848,878
Urosoma planktonicum 494 Urostyla weissei naine 845
Urosoma polynuc/eata 4, 11, 18,35,396,397,402, Urostyla weissi 845
405T,411,417,419,421 Urostyla weissii 844, 845, 878
Urosoma salmastra 396,398, 405T, 422, 424 Urostyla wessei 845
Urosoma similis 18,245,396,398, 405T, 419 urostyla, Kerona 897
Urosoma sp. 23, 431 urostyla, Oxytricha 244, 873
Urosoma stenocephala, Oxytricha 822, 933 urostyla, Pseudourostyla 244
Urosoma, Oxytricha 79, 116,396,414 Urostylen 261
Urosomas 396 Urostylidae 26,79,80,841,884
Urosomoida 1, 15, 17, 18,20,56,61,66,67,68,70, urostylids 263, 401,842,884
73, 74, 75, 76, 77, 78, 81, 112, 114, 118, 159, Urostyloidae 80
170,194,207,224,226,288,292,325,345,349, Urotricha pelagica 494
367,397,431,432,480,679 Urozoma cienkowekii 404
Urosomoida agiliformis 35, 36, 57, 64, 68, 158, Urozoma cienkowskii 396
159,170, 184, 186,345,347, 349, 352, 354T, 356 Urpleptus agilis 367
Urosomoida agilis 8, 11,35,64, 106,224,345,347, Utroleptus agilis 347
352, 354T,407, 409, 817
Urosomoida antarctica 121,346,347, 354T, 365 vannus, Euplotes 839
Urosomoida dorsiincisura 4,11,194,207,345,346, vannus, Kerona 839
354T,362 variabilis, Onychodromopsis 268
Urosomoida granulifera 11, 120, 346, 347, 352, variabilis, Oxitricha 116, 193,246
354T variabilis, Oxytricha 122, 170, 190,193,246
Urosomoida minima 35, 345, 346, 347, 351, 354T, variabilis, Pleurotricha 261, 262, 263, 265, 268,
366 476,699,782
Urosomoida perthensis 4, 345, 346, 354T, 362 Varietllt 1, Stylonychia mytilus 505,550
Urosomoida spp. 18, 159 Varietat 2, Stylonychia mytilus 505,549
Urosomoidea 346 velox, Oxytricha 244
Urosomoides 346 vernalis, Urostyla 844, 845, 846, 848
Urosomoides agiliformis 359 Viosoma planctonicum 494
Urostila we issei 845 virgula, Stylonichia 549
Urostyla 61, 250 virgula, Stylonychia 548
Urostyla coei 845, 846, 848, 855, 870 virgula, Stynolychia 548, 597
Urostyla dispar 842 viridis, Oxytricha 244, 254
Urostylaflavicans 844, 845, 848, 870 viridis, Paraurostyla 475, 842, 843
 SYSTEMATIC INDEX 1079

viridis, Perisincirra 894 Wallackia 74, 368, 369

viridis, Stylonichia 599 weisei, Paraurostyla 845
viridis, Urostyla 843, 870 weissei complex, Paraurostyla 111, 843,844, 868T
vitiphila, Pattersoniella 4, 111, 112, 500, 766, 776T, weissei naine, Urostyla 845
884 we issei, Paraurostula 841
volutator, Himantopus 116 weissei, Paraurostyla 10,23,25,26,39,49,54,87,
volutator, Oxitricha 116 244,263,268,487,843,844,846,845,848,852,
Volvox oniscus 566 854,855,856,857,858,859,862,863,864,865,
vorax, Clara 591 866,867, 868T, 870, 871, 873,874, 877
vorax, Gastrostyla 791, 792, 797 we issei, Parurostyla 841, 845
vorax, Histrichulus 646 weissei, Urostila 845
vorax, Histriculus 87,647,656,659,667 we issei, Urostyla 841, 844, 848, 878
vorax, Histriculux 646 weissi, Urostyla 845
vorax, Histrio 659 weissii, Urostyla 844, 845, 878
vorax, Oxytricha Histrio 591, 659 wessei, Urostyla 845
vorax, Oxytricha Stylonychia 591, 659 wrzesniowskii, Oxytricha 244
vorax, Stylonichia 591
vorax, Stylonychia 36, 76, 86, 87, 502, 503, 557,
562,565,568,571,588,591,593,604
Vorticella sp. 84
Table Index

Table! ............. page 4 Table 48 ............ page 802

Table 2 ............. page 9 Table 49 ............ page 868
Table3 ............. page 33 Table 50 ............ page 876
Table 4 ............. page 64 Table 50.1 ........... page 883
Table 5 ............. page 65 Table 50.2 ........... page 883
Table 6 ............. page 67 Table 51 ............ page 887
Table 7 ............. page 79 Table 52 ............ page 927
Table 8 ............. page 79 Table 53 ............ page 937
Table 9 ............. page 80
Table 10 ............ page 80
Table II ............ page 80
Table 12 ............ page 81
Table 13 ............ page 81
Table 14 ............ page 87
Table IS ............ page 131
Table 16 ............ page 185
Table 17 ............ page 186
Table 18 ............ page 205
Table 19 ............ page 267
Table 20 ............ page 291
Table 21 ..... . . . . . .. page 339
Table 22 ............ page 354
Table 23 ............ page 382
Table 24 ............ page 405
Table 25 ............ page 442
Table 26 ............ page 448
Table 27 ............ page 449
Table 28 ............ page 450
Table 29 ............ page 474
Table 30 ............ page 486
Table 31 ............ page 534
Table 32 ............ page 543
Table 33 ............ page 544
Table 34 ............ page 545
Table 35 ............ page 546
Table 36 ............ page 585
Table 37 ............ page 618
Table 38 ............ page 644
Table 39 ............ page 654
Table 40 ............ page 674
Table 41 ............ page 691
Table 42 ............ page 710
Table 43 ............ page 722
Table 44 ............ page 736
Table 45 ............ page 750
Table 46 ............ page 764
Table 47 ............ page 776
Table 47.1 ........... page 786