Guides to the Identification of the Guides to the Identification of the Microinvertebrates

Microinvertebrates of the of the Continental Waters of the World

Continental Waters of the World Coordinating editor: H.J.F. Dumont ISSN 0928-2440

Coordinating editor: Henri J. Dumont

State University of Gent, Belgium
ROTIFERA
Volume 3: Notommatidae

by
ROTIFERA Thomas Nogrady
Queen's University
Kingston, Ont. Canada

and
Editor
Thomas Nogrady Roger Pourriot
Queen's University, Kingston, Ont. Canada Universite 6
Paris, France

Editorial Committee
w. Koste, QuakenbrOck, Germany
R. Shiel, Albury, Australia
Scaridiidae

by

Hendrik Segers
Gent University
Gent, Belgium

SPB Academic Publishing bv 1995 SPB Academic Publishing bv 1995

 Table of Contents

CIP-DATA KONINKLlJKE BIBLlOTHEEK, DEN HAAG

Rotifera. INTRODUCTION
Rotifera I red. Thomas Nogrady]. - Amsterdam:
SPB Academic Publishing ACKNOWLEDGEMENTS 2
Vol. 3: Notommatidae I by Thomas Nogrady and Roger
Pourriot, Scaridiiae I by Hendrik Segers. - 111.-
(Guides to the identification of the microinvertebrates
FAMILY NOTOMMATIDAE 3
of the continental waters of the world, ISSN 0928-2440; 8) Subfamily TETRASIPHONINAE 3
With index, ref. Subfamily NOTOMMATINAE 5
ISBN 90-51 03-1 03-3
Subject headings: Rotifera.
FAMILY SCARIDIIDAE 230

GLOSSARY 238

REFERENCES 239

INDEX 245
ISBN 90-5103-103-3

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KEY TO FAMILIES AND SUBFAMILIES

1. Unci pointing inwards Fam. Notommatidae, page 3

Unci pointing outwards Fam. Scaridiidae, page 230

Family NOTOMMATIDAE

INTRODUCTION

Notommatidae is probably the most difficult family among all monogonont

rotifers, and also comprises some of the genera possessing the largest number of spe-
cies (e.g., about 190 species of Cephalodella are reviewed here). A confusing superfi-
cial phenotypic similarity exists among and within genera, thus misidentification is a
very real danger. To minimize this, live observation of animals is imperative, first fol-
lowing free-swimming specimens then restraining them by gentle compression to ob-
tain precise measurements and detailed observations. Morphometric studies on a sig-
nificant number of animals is useful as intraspecific variability of body measurements
is small due to eutely (constant number of cells) in a species. Narcotization lnay fol-
low using carbonated water, procaine, bupivacaine, or tricaine (see Nogrady & Rowe
1993), but the shape of the animal is likely to change. Preserved material is of very
limited value without narcotization, although trophi alone may be sufficient for iden-
tification in some cases. If used, preserved specimens should be as fresh as possible,
as pigments of eyespots etc. deteriorate. Thorough study of the sclerotized part of the
mastax, the trophi (sing. trophus) is mandatory in all cases using the NaOCI (bleach)
technique; it is becoming increasingly obvious that modem taxonomy requires scan-
ning electron microscopic (SEM) study oftrophi. For techniques oftrophi preparation
for SEM see Kleinow et al. 1990, and Koste & Shiel 1991 and references therein.
There are many species which have never been observed since their description,
such as many of the North American acidophilic species reported by Harring and
Myers, and by Myers. This does not mean that they are not valid taxa, it only indi-
cates the paucity of rotiferologists in that area. On the other hand, anthropogenic deg-
radation of habitats may have caused the extinction of localized rare species. Whene-
ver possible, attention has been drawn to the fact of a unique observation: those spe-
cies are marked with an asterisk in the keys, followed by a remark in the species de-
scription; lack of cited literature also testifies to single observation. If such a species
is encountered, a most painstaking re-examination and re-description is mandatory
and of great scientific value if accompanied by biometric, ecological and behavioural
observations usually lacking in the older literature. Observations should be docu-
mented by drawings and photomicrographs (which do not obviate but supplant each
other; SEM pictures are most desirable, as are permanent slides). Photos ofholotypes
of these rare species have been provided wherever possible. There are a number of
"species inquirendae", taxa which need careful reexamination. Their original desc-
riptions do not satisfy present-day requirements, and do not allow a decision regard-
ing their validity. These species are listed at the end of their genera. New rotifer spe-
cies should never be described from a single specimen and, for illoricates, only with
great caution if only preserved material is available. Holotypes and co-types must
be deposited at several locations; experience has taught us that even the best slides
2 3
deteriorate and the best museums lose material. Non-specialists are advised to seek
the counsel of a specialist in all doubtful cases; the literature is full of errors and
misidentifications, easily avoidable in our era of instant global communication. One
of us (T.N.) has inspected the type specimen slides of Myers available at the Ameri-
can Museum of Natural History. Unfortunately, most of them are of little use as gross
appearance is of limited value in this group; in addition many of the 60-70 year old
slides deteriorated. Nevertheless, Boyko (1994) published a list of all the slides to fa-
cilitate accessibility through the lending program of the Museum.
The identification keys provided follow classical patterns, but synonymy and spe-
cies relationships have been examined with methods of numerical taxonomy, using
the NTSYS-pc package of computer programs written by FJ. Rohlf (available from
Exeter Software, 100 North Country Road, Building B., Setauket, NY 11733,
U.S.A.). Unfortunately, a multiple entry key proved to grow unmanageably cumber-
some in some species-rich taxa (e.g., Cephalodella). Here only a less desirable
dichotomous key has been provided, even though it increases the danger of
misidentification caused by even a single error in observation. In most cases both a
multiple entry key and a dichotomous key have been supplied. We also provided a
code at the end of each species description (except in small genera) comprised of a
series of descriptors, defined before the key to that genus. The codes were used in
key construction as well as in numerical methods. Significant and crucial codes are
printed in bold. Such codes might eventually lend themselves to the development of
a computer-based identification system. Since species descriptions are arranged in
the sequence in which they "key out", it is relatively easy to compare the codes of
related species and uncover potential errors in identification. In species-rich genera,
. however, we deviated from this rule as it seems that an alphabetical listing is easier to
manage.
Proper identification requires perusal of the original literature, which contains
more detail than can, by necessity, be provided in a taxonomy text. Therefore, we en-
deavoured to call attention to the most impOliant additional sources pertaining to a
species. This list is selective, rarely complete; old sources, obsolete or available only
with difficulty are often omitted. To avoid redundancy, we also omitted constant cita-
tion of the indispensable text of Koste (1978); almost all species described before
1977 (with the exception of some North American taxa) are described and figured in
that work, which can still be consulted profitably. It also provides complete syn-
onymy and references, but the latter are not error-free.
A final counsel to the adventurous taxonomist by an anonymous sage: "When you
hear hoofbeats, think horses - not zebras".
The genus Notommata was re-examined by R. Pourriot and bears his authorship;
the family Scaridiidae has been reinstated and reviewed by H. Segers. The rest of the
book (and overall editing) is the work ofT. Nogrady with consulting by R. Pourriot.
ACKNOWLEDGEMENTS
Our warm thanks are due to our friends Rama Chengalath, Walter Koste, and Russel
Shiel for their invaluable help and consultation in solving long-standing problems
and for editorial assistance. We are grateful to C.R. Boyko and the AMNH for the
loan of type material. R.P. thanks Claude Rougier and Andre-Jean Francez for help in
sample collection and manuscript preparation.
March 1995 T.N. and R.P.
FAMILY NOTOMMATIDAE RE MANE 1933
A taxonomically unsatisfactory assemblage of diverse taxa, in need of revision by
modem methodology. The subfamily of Proalinae was separated from the original
family of Remane by Bartos (1959), even though resemblance of body shape makes
separation difficult on occasion, and given family rank.
The family was then further divided into two subfamilies by Bartos (1959), based
on the presence of a wreath of bulbous glands between stomach and intestine
(Tetrasiphoninae) or its absence (Notommatinae). The former contains two g~rera,
the latter 19 'C{i(~'.2~M .
In an attempt to mitigate the "catch-all" nature of the family, the genus" fUlra has
now been transfered as a S'l±BfamlJ;-.~, which is to be published as a
separate volume of these Guides. While this may not be entirely satisfactory, it is
based on trophi structure which - in agreement with views of Myers, Remane and
others - puts Itura closer to Dicranophorids than Notommatids. The genus Scaridium
has been restored to family status on the recommendation ofH. Segers (Segers 1995),
who reviewed the family at the end of this volume (see p. 230). The erection of a sub-
family Cephalodellinae seemed feasible but would not have achieved much, as the
genus constitutes almost half of Notommatinae anyway.
In his new, radically different system Markevich (1990; see Nogrady et al., 1993,
Vol. 1 of this series) established a superfamily Notommatina within the order Septira-
mida, based entirely on trophi ultrastructure. Eosphoridae, Notommatidae, and
Trichocercidae belong to this superfamily, but Markevich's ideas still require addi-
tional justification and examination.
KEY TO SUBFAMILIES
L Stomach and intestine separated by a wreath of bulbous glands " .
.............................................................................................. Tetrasiphoninae
Glands absent Notommatinae page 5
Subfamily Tetrasiphoninae
Genus Tetrasiphon Ehrenberg
Figs. 1-8
Monotype: Tetrasiphon hydrocora Ehrenberg 1840 p.2l9.
Type locality: Berlin.
Holotype: Not designated
Description: Body soft, illoricate, but integument stiff; head and trunk not separated.
Foot short with two segments; toes elongated, acute. Corona oblique, extending to a
"chin". Dorsal paired antennae on long stalk, lateral antennae towards posterior end
of trunk, also on very long stalk with long sensory bristles. Single cerebral eyespot,
subcerebral glands very long. Connection between stomach and intestine a narrow
tube surrounded by a wreath of about six round glands. Vitellarium is a long narrow
4 5

GENUS ET SPECIES INQUIRENDA, INCERTAE SEDIS

Genus Repauliana Berzil}S

Figs. 9-11

Monotype: Repauliana dicerea BerziJ}s 1960, p.I-3, Figs. 1-3.

Type locality: Laguna Lanirago, Madagascar.
Holotype: Not designated

Description: Unsatisfactorily described from a few preserved specimens, not neces-

sarily belonging to this subfamily.
Fusifonn hyaline body; flexible integument surrounded by a gelatinous sheath.
Trochus is an unbroken wreath of cilia. Toes large, ventrally recurved. Lateral anten-
nae somewhat below mid-point on trunk, on a long knobbed stalk with long sensory
setae. Trophus submalleate, not figured: rami wide, somewhat asymmetric and bent,
with sharp alulae; manubria lightly bent, unci thin and strongly bent.
Length of sheath to 400 !lm, total length 200-220 !lm (preserved), toes 25-34 !lm,
trophus 30 !lill, lateral antenna 11-14 !lill, sensory bristles 10-12 !lm.
Ecology: Apparently pelagic.
Literature: Not seen since description.

Subfamily Notommatinae

DESCRIPTORS OF NOTOMMATINAE GENERA

01 CORONA
Figs 1-8. Tetrasiphon hydrocora. 1,2: dorsal view;
g=wreath of glands; 3:1ateral view; 4,5: trophi
dorsally; 6: trophus laterally; 7: oral plates; 8: resting 01 on cylindrical snout
egg. 02 anterior
Figs 9-11. Repaulania dicera. 9: ventral view; 10: 03 oblique
lateral view with jelly case; 11: antenna. (1,5,6,7: 04 ventral
Nogrady 1979; 2,3,4,8: Koste 1978; 9-11: Berzi:Qs
05 auricles present
9 10 1960).
06 palps present

02 EYES

band with about 20-30 nuclei arranged in a row, wound around the gastrointestinal 01 absent
tract. Unci with single tooth, rami lyrate, curved dorsally, with alulae; manubria com- 02 frontal eyes
plex, with appendages, connected to unci and rami. Adults usually covered in a yel- 03 frontal eyes on stalk
lowish mucus encasement. 04 frontal and cervical eyes
Total length 450-1000 !lm, toes 60-80 !lm, male to 300 !lm, parthenogenetic egg 05 cervical
140-155xllO-l15 !lill, resting egg 200x155 !lill. 06 variable
Ecology: Acidophil, cosmopolitan in Sphagnum pools; feeds on desillids.
Literature: Barring & Myers 1922, Pourriot 1965, Koste 1968, Nogrady 1980, Koste 03 TRUNK
& Shiel 1991.
01 loricate
02 illoricate
6 7

03 with deep dorsal folds, plicated NUMERICAL KEY TO GENERA OF NOTOl\1MATINAE

04 lorica plates smooth
05 lorica plates granular, thick 11,21,32,42,52,53,61,64,71,81,92 Drilophaga p. 8
06 distal end of trunk with spine 12,15,24,32,42,45,52,53,61,64,72,81,91 (Itura t)
07 cuticle with rows of spines 12,23,31,42,45,52,53,61,64,71,82,83,91 Sphyrias p. 14
12,21,32,36,42,44,52,53,61,64,71,81,91 Dorystoma p. 32
04 FOOT 12,21,32,42,44,52,53,66,71,81,91 Pseudoharringia p. 13
12,25,31,41,52,54,61,64,71,81,91 Monommata p. 14
01 foot and toes longer than body 12,25,32,42,44,51,53,62,71,81,91 Resticula p. 210
02 foot and toes shorter than body 12,25,32,42,45,51,61,66,71,81,91 Tylotrocha p. 29
03 end of foot with spine 12,25,32,42,45,52,53,61,64,71,81,91 Pleurotrocha p. 217
04 segmented 12,25,32,42,45,52,53,61,64,71,81,91 Eosphora p. 158
05 unsegmented, pseudosegmented, or variable 13,16,25,31,34,42,45,52,61,64,71,81,91 Paracephalodella p. 147
13,21,31,34,41,52,53,61,64,71,81,91 Scaridiidae p. 230
05 TOES 13,22,32,44,52,53,61,64,71,82,84,91 Enteroplea p. 11
13,24,32,42,44,52,53,63,71,81,91 Eothinia p. 153
01 single 13,25,31,37,42,44,52,53,61,64,71,81,91 Pleurotrocha p. 217
02 double 13,25,32,42,43,52,53,61,64,71,81,91 Rousselettia p. 31
03 equal length 13,26,31,34,42,45,52,53,61,64,71,81,91 Cephalodella p. 33
04 unequal length 13,27,33,35,44,52,53,61,64,71,81,91 Metadiaschiza p. 33
14,15,25,32,42,45,52,53,61,64,71,81,91 Notommata p. 164
06 SALIVARY GLANDS 14,15,25,33,42,45,52,53,61,64,71,81,91 Taphrocampa p. 150

01 symmetrical N.B. The first number is the group descriptor, the second one the character within the group. The zeros of
02 aSyl1lll1etrica1 or rudimentary the descriptor list (necessary for ordination purposes) are omitted in the numerical key (e.g., 0102 be-
comes 12, etc.).
03 single .
04 double t see footnote on next page.
05 triple
06 absent
DICHOTOMOUS KEY TO GENERA OF NOTOMMATINAE
07 STOMACH
1. Corona on cylindrical snout, mouth deeply sunk, annular adhesive organ
01 without appendices present Drilophaga p. 8
02 with appendices Corona and mouth otherwise; no adhesive organ 2.
2 (1). Vitellarium band or ribbon-shaped , 3.
08 VITELLARIUM Vitellarium oval or kidney-shaped 5.
3 (2). Vitellarium nuclei linearly arranged 4.
01 oval or kidney-shaped Vitellarium nuclei randomly arranged Enteroplea p. 11
02 ribbon-shaped 4 (3). Eyeless; foot has two or three segments Pseudoharringia p. 13
03 nuclei linearly arranged Two frontal eyes on stalks; foot unsegmented, wrinkled Sphyrias p. 14
04 nuclei randomly arranged 5 (2). Foot and toes longer than body 6.
Foot and toes shorter than body 7.
09 ECOLOGY 6 (5). Toes (with one exception) of unequal length; foot short .. Monommata p. 14
Toes equal; foot very long Scaridiidae t p. 230
o1 free-swimming 7 (5). Foot with single toe; body orange-red Tylotrocha p. 29
02 parasitic Foot with two toes 8.
8 (7). Distal end of trunk or foot with spine 9.
8 9
Without spine 10. scribe all three species but refrain from providing a key, as the literature data are con-
9 (8). Distal end of trunk with spine Dorystoma p. 32 fusing, contradictory and the figures rife with artefacts. Further comparative inves-
End of foot with spine Rousseletia p. 31 tigations are necessary.
10 (8). Trunk loricate, 3-5 plates 11.
Trunk illoricate 12.
11 (10). Plates thick, granulated Metadiaschiza p. 33 Drilophaga bucephalus Vejdovsky 1883
- Plates smooth, occasionally indistinct Cephalodella p. 33 Figs. 12-15
_ Same, palps on corona Paracephalodella p. 147
12 (10). Cuticle with rows of small spines Pleurotrochopsis p. 147 Drilophaga bucephalus Vejdovsky 1883 p. 390, Figs. 1: 1-8
Cuticle without spines 13.
13 (12). Trunk deeply plicated dorsally Taphrocampa p. 150 Type locality: Unknown.
Trunk not plicated 14. Holotype: Not designated
14 (13). Two frontal eyes and one cervical eye 15.
- No frontal eyes, cervical eye absent or present 17. Description: As described in genus. The corona forms a tucked-in hood over the
15 (14). Stomach with appendices or sacs Iturinae t mouth on sessile specimens; this hood is dependent on the degree of contraction, that
- Stomach without appendices 16. might account for differences between D. bucephalus and D. delagei figures (see
16 (15). Mastax with one salivary gland Eothinia p. 153 Figs. 12-19). Fulcrum straight, without tenninal knob; straight epipharyngeal rods at-
- Mastax with two salivary glands Eosphora p. 158 tached to tip of rami; alulae pointed, manubrium with strong crutch. Large gastric
17 (14). Corona ventral, ciliated auricles present Notommata p. 164 glands.
Corona anterior, no auricles 18. Total length 110-355 !-tm, toe 6-11 !-tm, trophus 20-32 !-tm.
18 (17). Salivary glands symmetrical...................................................................... 19. Ecology: Ectoparasitic on the integument of oligochaetes (exclusively?), but can
- Salivary gland rodimentry or asymmetrical Resticula p. 210 leave the host and swim (Koste 1972c). In Gennany (ibid.) at 8-10°C, pH 6.6-7.0 in
19 (18). Rami triangular, robust Pleurotrocha p. 217 decaying Sphagnum and Typha. Germany, Poland, Roumania.
- Rami lyrate, thin and slender Pleurata p. 223 Literature: Paw10wski 1934, Koste 1972c.
t The genus Itura is included here for historical reasons only. As discussed in the Introduction, it was
transferred as Subfamily Iturinae to the Family Dicranophoridae, to be published as a separate volume of
these Guides. Scaridium has been promoted to the rank of a family, see p. 230.
Drilophaga judayi Harring & Myers 1922
Figs. 17-19

Genus Drilophaga Vejdovsky D.judayi Harring & Myers 1922, p. 612-614, Fig. 52:6-8

Drilophaga Vejdovsky 1883, p. 390
Type: Drilophaga bucephalus Vejdovsky 1883, p. 390, Figs. 1:1-8.
Description: Body slender, fusifonn, faintly segmented head cylindrical, narrow,
with only a circumapical ciliary band, combined with a circular suction ring. Small
tail, toes small, conical, footglands with reservoir; three salivary glands. Large RCO,
eyespot absent; dorsal and lateral antennae present. Trophus virgate, incus anchor-
shaped, rami curved with large alulae; unci thick, bidentate, resembling a crab's claw;
manubria terminally crotched. Parasitic on oligochaetes and leeches, except D. judayi
which was found free (see below).
While Koste (1972c and 1978) synonymized all three known species, Koste &
Shiel 1991 state that this decision requires re-examination, due to habitat and trophi
differences. Harring & Myers 1922 found D. judayi free-swimming in decaying
Sphagnum, albeit apparently in a moribund state. A free-swimming individual of a
Drilophaga sp. was also reported by Koste & Shie1 (1991) from Australia. We de-
Type locality: Ditch near Mamie Lake, Vilas County, WI, U.S.A.
Holotype: Not designated
Description: Body spindle-shaped, soft, transparent. Corona apical with feebl.e
circumapical ciliation only; movement (according to Harring & Myers 1922) restn-
cted to bending, not swimming, but they may have dealt with moribund specimens.
Small tail over the unsegmented foot, toes minute. No Reo (?) or eyespot. Trophus
virgate, fulcrum slender rod; rami strongly curved with large rounded alulae, no teeth
and no rods; manubria slender, S-shaped, crutched. Unci small, two oval plates, re-
sembling a crab's claw.
Tota11ength 240-275 !-tm, toes 8 Ilm, trophus 20 Ilm.
Ecology: Rare among decaying Sphagnum, free-swimming. Harring & Myers' de-
scription of feeble motion indicates moribund specimens.
Found in Russia, Czechoslovakia, Sweden (as quoted in Kutikova 1970; original
source unknown).
Literature: Kutikova 1970.
 10
11

Drilophaga delagei de Beauchamp 1904

Fig. 16

D. delagei de Beauchamp 1904, p. 159, Fig.C.

Type locality: Paris, Vaux de Cernay, on Helpobdella octooculata

Holotype: Not designated

Description: Similar to D. buceplzalus but shown in all figures in the literature with
extended corona with a small hood or without hood, unlike D. bucephalus. Thus, the
difference shown in the corona may be a perpetuated artefact. Main diagnostic differ-
ence may be the large, rounded unci, trapezoidal rami, fulcrum with terminal knob,
13 alulae with small terminal projection, manubria with small crutch; small gastric
glands; no body pseudosegmentation (although Pawlowski 1934 questions the reality
of the deep segmentation in the original description and figure of Vejdowsky 1883).
Specimens attached to the host are immobile, including the mastax.
Total length 148-277 [lm, toes 6 [lm, trophus 23 ~tm.
Ecology: Ectoparasitic on Herpobdella spp. (exclusively?), but juveniles have been
encountered free-swimming (Koste 1972c). France, Poland, Roumania
Literature: Pawlowski 1934, Kutikova 1970, Koste 1972c.

Genus Enteroplea Ehrenberg

12 Figs. 20-25

S'(I/I. ~T(rfe/&{s

~
b Enteroplea Ehrenberg 1830, p.46.

tf%
i...... "J
,.c~-"
l,I
Type (Monotype): Enteroplea lacustris Ehrenberg 1830.
Type locality: Berlin.
18 Holotype: Not designated
16

fl
a Description: Body saccate, head wide, rectangular; foot well separated, shifted ven-
trally, 3-segmented. Toes short, footg1and long. Corona slightly oblique, consisting of
a ciliary wreath. The dorsal margin forms a pseudotrochus incorporating four oblique
membranelles. Adjacent to the mouth on either side another row of membranelles is
19 placed on a protuberance, mouth surrounded by another ciliary wreath. Several sen-
17 sory membranelles on the corona, two frontal eyespots on papillae. Dorsal antenna on
a small wart, lateral antennae on distal dorsal end of trunk. Two small salivary glands,
F~'gs 12-15. prilophaga buc~phalus. 12: dorsal view; 13,14: trophi orsally and laterally; 15: lateral view. stomach cellular, intestine conical, funnel-shaped with longitudinal furrows (not
F'.lg. 16. ?nlophaga delagel. Ca): de Beauchamp's (1904) original figure (b): de Beauchamp's later
fIgure wIth trophi. ' "rope-shaped" (strangf6rmig) as stated in Koste 1978 and Koste & Shiel 1991). Gas-
Figs 17-19. Dril0p,hagajudayi. 17: lateral view; 18,19: trophi. (12-14: Koste 1972c' 15 16: de Beauchamp tric glands band-shaped with forked ends; on the dorsal side of the stomach a pair of
1904; 17-19: Harrmg & Myers 1922). ' ,
additional long and narrow appendages. Trophus powerful: rami bent at right angle,
extensible through mouth; on a finely denticulate margin one powerful tooth on each
side; the ventral end of rami plier-shaped. Alulae large, one small papilla on each
ramus. Unci with one main and one ancillary tooth; fulcrum board-like.
Total length 500-600 [lm, toes 30-35 [lm, trophus 70 [lm, male 306 [lm, parthenoge-
netic egg appr. 160x120 [lm.
 12 .
13

Ecology: Predator in cultures (Pourriot 1965). In shallow or ephemeral ponds in

spring, cosmopolitan but not frequent.
Literature: Koste 1986.

Genus Pseudoharringia Fadeew

Pseudoharringia Fadeew 1925, p. 73, Figs. 1:3-7

25 Type: Pseudoharringia similis Fadeew 1925
Type locality: ?

Description: Unsatisfactorily described. Body plump, saccate, gibbous, narrowing

towards foot, head not widened. Body and foot segmented or pseudosegmented; foot
conical, toes tiny, blunt. Oesophagus very long and narrow dilating into a pro-
ventriculus and a wide stomach; intestine thin, rope-like, Ovovitellarium a ventrally
placed band with six linearly arranged nuclei. No eyespots. Trophus not well de-
scribed: rami with small alulae, unci "apparently" consisting of five platelets (?), ful-
crum terminally spatulate or spoon-shaped.

KEY TO PSEUDOHARRINGIA

1. Oesophagus long with proventriculus, manubria shorter than fulcrum .........

....................................................................................... ....................... similis
Oesophagus short, no proventriculus, manubria longer than fulcrum ..
....................................................................................................... romanica*

Pseudoharringia similis Fadeew 1925

Fig. 26

P. similis Fadeew 1925, p.73, Figs. 1:3-7.

Type locality: Russia?

Holotype: Not designated

ffi Description: As described in genus. Trunk with distinct wrinkles.

Total length 300-400 ~m, foot 60-70 ~m, toes 8-14 ~m.
Ecology: In plankton and submerged plants in lakes and running water. East- and
27 southeastern Europe, China, Russia.
Literature: Koste 1978

SPECIES INQUIRENDA
!"igs 20-~5. Enteroplea. lacustris. 20:laterally; 21: dorsally; 22: trophus dorsally; 23: trophus laterally; 24:
mcus oblIque frontal VIew; 25.: tr?phus oblique frontal view (a: right ramus teeth; b: epipharynx). Bar 50 Pseudoharringia romanica Rodewald 1937
~L~ for body, 10 ~m for trophl. FIg. 26. Pseudoharringia similis. Fig. 27. Pseudoharringia romanica. Fig. 27
Fl~S 28-30. Sphy~zas lofitana. 28: lateral and dorsal view; 29: trophus, ventral and lateral view; 30: trophus
oblIque frontal VIew. (20-24: Harring & Myers 1924; 25: Koste 1986; 26,27: Kutikova 1970 ; 28-30: P. romanica Rodewald 1937, p. 236, Figs. 2a-c.
Harrmg & Myers 1924).
 14 15

more folds; cuticle thin, finn, laterally and dorsally with longitudinal striae; foot h\'o-
Type locality: Lake Tabacarei, Roumania. or three-jointed, sometimes indistinctly; toes up to twice of body length, right longer
Holotype: Not designated than left (with exception of M. aequalis) and filled with striated muscle, used for skip-
ping locomotion. The corona is slightly oblique, with a marginal whorl of cilia and
Description: Trunk smooth, rami with distinct alulae, unci consist of three plates (?). lateral auricle-like tufts of longer cilia for swimming; apical field unciliated, buccal
Described from single specimen ? field ciliated. Mastax variable, from simple virgate to intermediate between virgate
Total length 280 p,m, foot 70 ~lm, toes 9 ~m. and forcipate type; in fonner type (Figs. 31-38) rami are lyrate or triangular without
Ecology: Benthic, pH 8.4 inner teeth, manubria simple rods, unci with one weak tooth or reduced to thin lamel-
Literature: Not found since description. lar plates (Myers 1930). In the trophus of the intennediate type (Figs. 40-46), rami
lyrate with one or more teeth on inner margin, manubria broad and lamellar at base,
unci with three unequal, long, slender clubbed teeth; dorsal antennae single or paired,
Genus Sphyrias Harring 1913 sometimes on papillae; number of antennae uncertain in some species and requires
Figs. 28-30 revision. Lateral antennae normal, cerebral eye at posterior end of brain (absent in M
caeca). Variations from generic characters are detailed by Koste (1978) and de-
Sphyrias Harring 1913, p. 96. scribed in the species diagnoses below. Observation of live animals and very gentle
preparation of trophus is mandatory for proper identification, which is based almost
Type (monotype): Sphyrias lofuana (Rousselet 1910) = Notops lofuana Rousselet entirely on trophus morphology. Seventeen species are described, which include three
1910, p. 795, Figs. 75:1-3. that are synonymized. The North American species were all collected from acid
Type locality: River Lofu, at Lake Tanganyika, Kenya. waters.
Holotype: Not designated

Description: Body pI ump, saccate, head separated from trunk, wider than body, trian- MONOMMATA DESCRIPTORS
gular in stationary animal, trapezoidal in swimming specimens. Foot long, segmented,
toes wide, dagger-shaped. Corona reduced, sensory papillae and cirri on apical field 01 TOES
as well as widely separated eyespots also on projections. Dorsal antenna on a stalk.
Salivary glands huge, gastric glands round, stomach cellular, not separated from in- 01 equal length
testine. Ovovitellarium band-shaped with linearly arranged nuclei. During feeding/ 02 unequal, shorter than body
sucking the mouth-opening becomes conical and a snout-like antenna covered with 03 unequal, longer than body, right/left toe ratio <1.2
sensory cirri appears. Trophus robust, prehensile, rami with huge alulae, bent at 90°, 04 unequal, longer than body, right/left toe ratio> 1.2
the bent part covered with numerous fine teeth. Unci with a single clubbed stout tooth, 05 right toe/body ratio <1.2
manubria bent, sometin1es with a small bump at the middle. Fulcrum plank-shaped, 06 right toe/body ratio> 1.2
fanned in dorsal view. Epipharynx consists of two pleural rods expanded tenninally
into triangular plates. Parthenogenetic egg dark, hairy. Closely related to Eosphora. 02 RED SPOT AT LATERAL ANTENNAE
Total length 204-320 ~tm, toes 25-35 ~m, trophus 60-65~m, epipharyngeal rods 20
~m, egg 48x65 ~m. 01 present
Ecology: Predator, apparently cosmopolitan (exc. AustralialN.Z.), pH 6.5, warm 02 absent
stenotherm. According to Koste 1978, very frequent in warm-water aquaria.
Literature: Harring & Myers 1924. 03 DORSAL ANTENNA

01 single
Genus MOl'lommata Bartsch 02 double

Monommata Bartsch, 1870, p. 344. 04 STOMACH SACS

Type: Vorticella longiseta Miiller, 1786 = Monommata longiseta (Miiller, 1786). 01 present
Type locality: Copenhagen. 02 absent

Description: Body cylindrical or fusifonn; head separated from abdomen by one or

 16
17
05 BODY LENGTH 14, 15,21,327,42,51,61, 72, 81,86,91,94 actices
14,15,22,31,42,51,63,74,81,86,91,94 pseudophoxa
01 <200 llm 14, 15, 22, 31,42, 51, 63, 74, 83, 84, 93, 95 dentata
02 >200 llm 14, 15,22,31,42,52,61,71,82,83,92,95 hyalina
14,16,21,31,42,52,64,73,81,84,92,95 grandis
06 RAMI TEETH 14,16,21,32,42,51,61,71,75.81,86,91 (amdti)
14, 16, 22, 31, 42, 51, 61, 72, 81, 86, 91, 95 caeca
01 missing 14,16,22,31,42,51,61,74,81,86,93 enedra
02 1/1 or 1/2 14,16,22,31,42,51,61,77,83,86,91,94 phoxa
03 2-3 14,16,22,31,42,51,62,73,81,86,93,95 longiseta
04 5 and tooth-series 14,16,22,32,41,51,61,74,81,84,91,95 viridis
05 4+3+ 12-13

07 UNCI DICHOTOMOUS KEY TO MONOMMATA

01 with single tooth 1. Toes equal aequalLs' p. 18
02 with 2 teeth Toes unequal 2.
03 with 2-3 teeth 2(1). Stomach with sacs viridis p. 29
04 thin lamella without teeth or ribs Stomach without sacs 3.
05 thin lamella with ribs or thin edge
g~~~:~ ::~:::: ~i~~~~ ::}:#q~~::~~~~~~~~:.~~?~f!}f[:!/~.~· . ~.~~.~~.
3(2).
08 FULCRUM 4(3). Rami without teeth 5.
Rami with teeth 12.
01 frontalIy a thin rod 5(4). Basal apophysis present 6.
02 bulging middle Basal apophysis absent 9.
03 terminally widened 6(5). With large caudal appendage 7.
04 basal apophysis present No caudal appendage 8.
05 basal apophysis double 7(6). Rami with large alulae appendiculata p. 20
06 basal apophysis absent Rami without alulae (caudata)# p. 22
8(6). Manubrium with terminal hook and median bump aeschyna p. 20
09 MANUBRIUM Manubrium slightly bent, no bump astia p. 21
9(5). Fulcrum fusiform, striated; manubria hooked hyalina * p. 25
01 with terminal crutch Fulcrum a thin rod 10.
02 with terminal hook 10(9). Eyeless _ caeca p. 22
03 terminally straight With eyespot 11.
04 base with single lamella 11(10). Uncus a thin lamella without teeth enedra p. 24
05 base with double lamella Uncus with one tooth, fulcrum spatulate diaphora p. 24
Uncus with a bundle of 5 teeth phoxa p. 28
12(4). Ramus with a single tooth or projection 13.
NUMERICAL KEY TO MONOMMATA Ramus with 2 or more teeth 14.
13(12). Long single tooth longiseta p. 27
11,15,21,31,42,52,61,74,81,86,93,95 aequalis Tooth is a dull projection covered by uncus tooth #(caudata) p. 22
12, 15,14,22,31,42,51,61,74,81,84,93 astia 14(12). Ramus with 2-3 teeth 15.
12, 15, 14,22,31,42,52,61,72, 75(?), 81, 84, 93, 95 appendiculata Ramus with multiple teeth 16.
13,15(16),22,31,42,51,62,71,75,81,84,93,95 caudata 15(14). 3 + 3 teeth, one pair terminal dentata p. 24
13,15,22,31,42,51,61,71,81,84,93 aeschyna 2 + 3 teeth, none terminal pseudophoxa p. 28
13,16,21,31,42,52,65,73,81,85,92,94 maculata 16(14). Row of 25 teeth; basal apophysis simple grandis p. 25
13,16,22,31,42,52,61,71,81,86,91 diaphora 4 + 3 + 12 teeth; basal apophysis double maculata p. 27
 18 19

Mononunata actices Myers 1930

Fig. 31

Monommata actices Myers, 1930, pp. 394-5, Figs. 26:4-7.

r I j If . /I , ) " .
V/I- Ml/l~ I 1'(::3'1.',,0;,1 e It.
() J .
,.J~tr JI , -I
'1
I), ." (' 7
rlO/TX u)";;
Type locality: Atlantic County, NJ, D.S.A.
Holotype: Not designated

Description: Body slender, cylindrical, tapering to foot; head well separated frgm
trunk by dorsal wrinkles and folds of integument, which, as usual, is striated. Th~. !/1.<J
!,

g1e"dorsa1 antenna is tubular, retractile, and thus may escape observation. Lateral an- patted,
teImae with two associated round reddish or clear areas in lumbar region; only M.
grandis, maculata, arndti, and Cephalodella subsecunda carry such spots. Corona ob- 31
lique, trophus small, simple virgate; fulcrum long, rodlike with small apophysis; rami 32
slender, lyrate, curved downward in the middle, rodlike alu1ae, unci with two short
teeth; pair of thin lamellar plates between unci teeth and manubria. No constriction
between stomach and intestine; foot-glands, bladder and retrocerebra1 sac small; eye-
spot at posterior end of ganglion; egg with thorns. Described containing zoochlorellae
in the stomach, but this has not beenconTirmedsubsequently.
Body length 150-195 ~m, right toe 200-21 0 ~m, left toe 150-170 ~m, trophus 22-24
~m, parthenogenetic egg 48x5 8 ~m, spinu1es 15 ~m.
Ecology: Europe, N. America, in bog pools and lightly acid waters. In Australia and
Tasmania 23.0-25.0°C, pH 5.7-6.2, 48 ~S cm-I, DO 3.7 mg L-l.
Literature: Wu1fert 1960a, Donner 1978; Koste et al. 1988; Koste & Shiel 1991.
Comment: The single dorsal antenna was considered doubtful by Wu1fert (1960a);
Koste (1978) suggested that paired antennae*were Qy_erlooked.,byM:Y~rs, and that the
appearance of M. actices is identical with M. arndti Remane. The dimensions given in
Koste (1978) are ambiguous. There are also differences in the trophus as figured by
the two authors, although not very significant ones. We go along with Koste & Shie1
(1991), and Shiel (personal communication) and refrain from synonymizing M. arndti
with M. actices until further study.
Code:,14,)5, 21, 3~?, 42, 51, 61,, ' 72,81,8;;;6 91,94. P
*"
" ',., I I .' , ' ,~I
d~~U:.() Vdi0 Lt i Vl /"7DU prfl'hd lad] -)(IJ:JS
/,J.i" , ,'"I'

Monommata aequalis (Ehrenberg) 1832 37

Fig. 32

Notommata longiseta aequalis Ehrenberg, 1832, p. 134.

Type locality:, Berlin.

Holotype: Not designated
Description: Long, slender, hyaline body indistinctly separated from long and broad
head; single dorsal antenna. Rami 1yrate, without teeth on inner margin; when seen
laterally, rami form right angles to fulcrum; unci uniformly lamellate, fluted or with
marginal denticles; fulcrum rodlike; manubria bilaterally lamellate; subcerebra1
glands absent; toes of equal or similar length; red or yellow-orange vesicles beside
intestine. Trophus similar to that of M. caudata.
Fig. 31. Monommata actices laterally, with various views oftrop?i. .
Fig. 32. Monommata aequalis laterally, with lateral and frontal Vl~WS of troplu..
Fig. 33. Monommata aeschyna laterally, with lateral and ventral Vlews o~ trophl.. .
Figs 34-35. Monommata appendiculata laterally, with lateral and ventral Vlew oftrophl: 35: postenor end
of body, dorsally. . .
Fig. 36. Monommata arndti laterally, with ventral and lateral VIews of t!ophl.
Fig. 37. Monommata astia laterally, with frontal and lateral views oftrophi. (31: Wulfert 1960a; 32: Myers
1937; 33-35,37: Myers 1930; 36: Koste 1972).
20
Total length 200-227 ~Lm, toes 110-120 ~m.
Ecology: Rare, among macrophytes, Europe, S. America, Australia. In Tasmanian
collections 9-11 DC, pH 7.6-7.8, 9.0-13.4 ~S cm-I.
Literature: Myers 1937, Koste et al. 1988.
Code 11, 15,21,31, 42, 52,61,74,81,86,93,95.
Monommata aeschyna Myers 1930
Fig. 33
Monommata aeschyna Myers, 1930, p. 387, Fig. 24:4-6.
Type locality: Cordoy Creek, Atlantic County, NJ, U.S.A.
Holotype: Not designated
Description: Body elongate, fusiform; head demarcated by slight constriction; body
tapers gradually to indistinctly three-segmented foot. Single dorsal antenna on low pa-
pilla; trophus virgate, small, simple; rami triangular, without teeth or denticles; alulae
prominent; manubria simple rods, distally curved, with median blunt tooth-like proc-
ess dorsally; unci single-toothed; fulcrum staight rod with basal apophysis. Gastric
glands, retrocerebral sac small; eyespot on ventral side of ganglion.
Body length 130-150 ~m, right toe 150-165 ~m, left toe 120-145 ~m, trophus 25-35
~m.
Ecology: Sphagnum, ice-water pools, psammon; cosmopolitan but rare. In Australia
13-27°C, pH 5.4-6.9, 25-292 ~S cm-I, TDS 16.2-21.1 mg L-I, 1.1-7.3 NTU.
Literature: Berzi1)s 1949a, Ruttner-Kolisko 1953, Koste & Shiel 1991.
Code: 13,15,22,31,42,51,61,71,81,84,93.
Monommata appendiculata Stenroos 1898
Figs. 34-35
M. appendiculata Stenroos 1898, p. 135, Figs. 33, 34.
Type locality: Finland.
Holotype: Not designated
Description: Body stout and cylindrical, prominent tail overhangs two-jointed foot;
toes shorter than in other species. The tail remains visible in contracted animals.
Trophus small, rami triangular ventrally, tip bent at right angle, alulae large and diver-
gent. Sharp basal apophysis. Unci with two teeth that fonn the edges of a thin plate.
Manubria Sh011 with two lamellae. Ganglion and RCO are large, body calor character-
istically orange to brown.
Body length 225-250 flm, right toe 235-250 ~m, left toe 165-185 ~m, trophus
25 ~m.
Ecology: Acidophil, pH ::: 6.6
Literature: Myers 1930.
Code: 12,15,14,22,31,42,52,61,72, 75(?), 81, 84, 93, 95.
21
Alo110mmata arndti Remane 1933
Fig.36_
!vfonommata arndti Remane, 1933, p. 567-568 (nomen nudum).
Type locality: Moor pool near Kiel, Germany.
Holotype: Not designated
Description: Close to, if not identical with AI. actices. Paired dorsal antennae also on
retractile papillae; lateral antennae tubular; mastax with two paired and one single
salivary gland; retrocerebral sac with with two excretory ducts; two tubular sub-
cerebral glands; gut contents generally yellow-gold. Koste (1978) notes that paired
red lumbar bodies develop in response to food intake, and possibly are excretory de-
posits; ratio of body/toe length changes during development; pe:tIthenogenetic egg
spiny, male egg smooth-shelled. Male undescribed. Trophus: sickle-shaped rami slim
wIthout teeth, with extremely long hanging alulae, unci lamellar with single tooth (?),
fulcrum thin rod, no basal apophysis; manubrium bent, with median thickening, no
lamellae.
Body length 150-21 0 ~m, right toe 220-250 ~m, left toe 160-21 0 ~m, parthenoge-
netic egg 85 x 75 ~m, male egg 48x40 ~m.
Ecology: Eats phytoflagellates, by sucking cell contents. Acidophil. Europe, Aus-
tralia, 20.0 c C, pH 7.2, DO 9.1 mg L-'.
Literature: Koste 1972b; Koste & Shiel 1980, 1991.
Code: 14,16,21,32,42,51,61,71,75,81,86,91.
Remark: This species is probably a synonym of M actices. See remarks there.
Monommata astia Myers 1930
Fig. 37
M. astia Myers 1930, p. 390-391, Figs. 25:5-7.
Type locality: Not specified.
Holotype: Not designated
Description: Very slender according to original description, more robust in others
(Wulfert 1961). Toes relatively short, foot two-jointed, tail an.d zoochlorell~e s?me-
times present. Trophus very delicate and simple, rami lyrate WIth alul~e, unCI thm la-
mellar plates, manubria simple rods thickened in the middle. GanglIon long, RCO
black, eye may be missing.
Body length 120-170 ~m, right toe 75-80 ~m, left toe 60-65 ~m, trophus 15 ~m.
Ecology: Acidophil, not uncommon. Sluggish. Cosmopolitan?
Literature: Wulfert 1961. -?1'j1. Cl sho. ~ l~ (J. B~
Code: 06 12, 15,14,22,31,42,51,61,74,81,84,93.
22 23

Monommata caeca Myers 1930

Fig. 38

M. caeca Myers 1930, p. 391-392, Figs. 25:8-10.

Type locality: Bargaintown, NJ, U.S.A.

Holotype: Not designated

Description: Body cylindrical, slender, head separated by dorsal folds. Toes long,
ReO large, round, and clear, the only species of the genus without an eyespot. Rami
triangular, base of divergent alulae at right angle to slender fulcrum. Unci with two ~.,\.
linear teeth, between them and rami thin lamellae. Manubria sigmoidal, on dorsal la-
mella (cell) a small lobe or knob.
Body length 170 !-tm, right toe 210 !-tm, left toe 170 !-tm, trophus 25 !-tm.
V\ B
Ecology: Rare, moderately acidophilic (pH 6.4); North America (New Jersey,
Maine), Sweden, Russia, New Zealand. 38
Literature: Berziq.s 1949a.
A
39
Code: 14,16,22,31,42,51,61,72,81,86,91,95.

Monommata caudata Myers 1930

Fig. 39

M. caudata Myers 1930, p. 392-393, Figs. 26:1-3.

M. sphagnicola BerziIfs 1949a ?

Type locality: Mount Desert Island, !VIE, U.S.A.

Holotype: Not designated

Description: Body fusifonn (in the American description) or cylindrical and even
gibbous and coloured in the Gennan specimens. Integument flexible, dorsal striation
V-shaped, meeting in the dorsal middle line. Bifid caudal projection that does not
reach the end of foot like inM appendiculata. (N.B., the bifid shape is not shown in
Myers' figure, no tail at all is shown in Wulfert's figures). Only the trophi are unmis-
takeably identical in all specimens described, including M. sphagnicola. Laterally the
trophi resemble those of M. aequalis: manubria Y-shaped with a long, pointed, basal
apophysis. Rami lyrate with an inner projection, more slender and sickle-shaped with-
out a projection in Wulfert's figure (Fig. 39; this could be misleading in identifica-
tion). The unci are really the border of a wide lamella. ReO is small, black, eye small.
Body length 112-180 !-tm, right toe 144-210jJ.m, left toe 126-180~lm, trophus 18-22 40 41 42
jJ.m.
Ecology: Acidophilic, North America, Sweden, Gennany.
Literature: BerziI).s 1949a, Wulfert 1960a. Fig. 38. Monommata caeca laterally, with d?rsa~ and l~teral vi~ws of trophi. .
Code: 13,15(16),22,31,42,51,62,71,75,81,84,93,95. Fig. 39. Monommata caudata Group A: SWll111111ng ammals wIth frontal, lateral, and dorsal views of
trophi; Group B: Myers' figures. . '
Fig. 40. Monommata dentata laterally with dorsal and lateral vle:ws of troplu ..
Fig. 41. Monommata diaphora laterally with dorsal and lateral VIews o~ trophl. .
Fig. 42. Monommata enedra laterally with oblique frontal and lateral views of trophl. (38,41,42:
Myers 1930; 39: A: Wulfert 1960a,B: Myers 1930; 40: Wulfert 1940).
 24
25
Monommata dentata Wulfert 1940
from trunk. ReO and sack are absent, eyespot large, with lens. Rami triangular, lyrate,
Fig. 40
without teeth, bent at rightangle; alulae large. Manubria simple rods, bent at tenninal
third. Rami and manubria connected by a thin lamella.
M. dentata Wulfert 1940, p. 578-579, Fig. 22.
~ Body length 150 ~tm, right toe 210 !lm, left toe 155 ~tm, trophus 25 ~tm.
Ecology: Rare, pH 6.4, D.S.A, Sweden.
Type locality: Birkhorster Moor, Germany.
Literature: BerziQ.s 1949. ';--
Holotype: Not designated
Code: 14, 16,22,31,42,51,61,74,81,86,93.
Description: Body stout, head wider than trunk, separated by a fold. Longitudinal
striation dense. On the ganglion rc. sack filled with bacteroids. Rami with denticulated Monommata grandis Tessin 1890
frontal appendage and symmetrical or asymmetrical teeth, alulae horizontal. Large Fig. 43
basal apophysis, pleural rod occasionally present. Variable species.
Body length 100-200 [.tm, right toe 115-200 [.tm, left toe 89-160 [.tm, trophus 16-22 M. grandis Tessin 1890, p. 151, Figs. 1: 11-12.
[.tm. M. robusta BerzilJs 1949a, p.l, Figs. 1-3
Ecology: In acidic and alkaline pH, eurytherm. Europe, Australia, Tasmania.
,
Literature: Donner 1943, 1964. Koste & Shiel 1991. Type locality: Rostock, Germany. I
,';1
.'
Code: 14, 15,22,31,42,51,63,74,83,84,93,95. Holotype: Not designated I' (I//it) ,.:::(L)

Description: Body elongate, fusiform; characteristic red pigment spots beneath lat-
Monommata diaphora Myers 1930 eral antennae; foot indistinctly two-jointed; mastax of intermediate type between
Fig. 41 forcipate and virgate types: fulcrum about equal length to rami, broad at base, taper-
ing. Rami with thin lamellae medially; large basal apophysis; large alulae; inner mar-
M. diaphora Myers 1930, p. 388, Figs. 24:7-9. gins of rami with 25 or more comb-like ventral dentic1es and two pairs of four long,
slender opposing oral teeth; each uncus with plate-shaped ventral tooth, ending in five
Type locality: NJ, ME, D.S.A. tooth-like projections at tip, and distal rodlike tooth; manubria hooked distally, lamel-
Holotype: Not designated late proximally; retrocerebral sac small, clearly ducted to corona surface; no sub-
cerebral glands; mastax has confluent salivary glands; eyespot ventral at posterior end
Description: Body long, cylindrical, with characteristic dorsal swelling at stomach of ganglion. Related to M. maculata.
height. Eye with lens on ventral side of ganglion. Mastax with two large confluent Length of body 190-240 [.tm, right toe 210-470 [.tm, left toe 150-336 [.tm, trophus 35
salivary glands. Fulcrum without basal apophysis; rami triangular in ventral view, lat- [.tm.
erally right-angled, without teeth or denticles; alulae prominent; manubria reduced to Ecology: Cosmopolitan, rare, oligosaprob indicator; pH 4.5-7.2, 16-27°C. Europe,
simple rods, crooked terminally, attached to rami by thin lamellar plates; unci single D.S.A., Australia, Tasmania.
toothed. Literature: Koste & Shiel 1991.
Body length 225 [.tIn, right toe 260 [.tm, left toe 225 [.tm, trophus 25 [.tm. Code: 14,16,21,31,42,52,64,73,81,84,92,95.
Ecology: pH 6.2-6.4, D.S.A, Roumania, Australia.
Literature: Koste & Shiel 1991.
Code: 13,16,22,31,42,52,61,71,81,86,91. Monommata hyalina Myers 1930
Fig. 44

Monommata enedra Myers 1930 M. hyalina Myers 1930, p.382-383, Figs. 23:1-3
Fig. 42
Type locality: Mount Desert Island, ME, D.S.A.
M. enedra Myers 1930, p. 385-386, Figs. 24:1-3. Holotype: Not designated

Type locality: Mount Desert Island, ME, D.S.A. Description: Body stout, largest of all Monommata, flexible; corona very oblique,
Holotype: Not designated ending in a chin; dorsal antenna single. Strong constriction between clear int.estine
and stomach. Trophus very large, modified virgate; fulcrum long, stout, fusIform~
Description: Long cylindrical body with distinct tail separated by several wrinkles swollen in the middle, longitudinally striated, distal end recurved and serrated. RamI
 26
27

triangular with huge downward pointed blunt alulae; on right ramus at the articulation
a comb-like structure with 8-10 teeth, otherwise no teeth on rami. Unci \:vith one long
sigmoidal clubbed tooth. Basal part of manubria is almost circular (including the la-
mellae) and flat, distal end a pointed hook. Slender (pleural ?) rods embedded in
mastax below rami.
Body length 300 /-lm, right toe 350 /-lm, left toe 250 /-lm, trophus 70 ~lm.
Ecology: Acidophilic (pH 6.4-6.8) fairly common,Maine, New Jersey, D.S.A.
Literature: Not seen since discovery, not included in Koste 1978 (only figure sup-
plied).
Code: 14,15,22,31,42,52,61,71,82,83,92,95.

Monommata longiseta (Muller) 1786

Fig. 45

Vorticella longiseta Muller 1786, p. 285, Figs. 42:9-10

Monommata longiseta Bartsch [1870], p. 344.
45
Type locality: Copenhagen.
43 Holotype: Not designated
44
Description: Body slender, elongate, head short. Transparent integument marked
with closely spaced striae as in M. dentata. It differs from that species by trophus dif-
ferences: rami bent at right angle near mid-length, long slender tooth at angle on each
ramus; right uncus has three long slender teeth, left uncus two; fulcrum lacks basal
apophysis; manubria proximally broad, lamellar, distally rodlike, outward curving;
length of toes variable.
.,. Body length 86-115 /-lm, right toe 155 ~m, left toe 120 ~m, trophus 15-16 ~m .
III Ecology: Presumably abundant everywhere, but confusion with other species must be
III suspected.
W
Literature: Harring & Myers 1924, Koste & Shiel 1991.
:tl]'
i ~' a Code: 14,16,22,31,42,51,62,73,81,86,93,95.
[ )1
J93 0
Monommata maculata Harring & Myers jJ24'.....
Fig. 46
46 48 M. grandis Tessin 1890, Harring & Myers 1924, p. 538-540, Figs. 43:6-10. This was a
misindentification, corrected as M. maculata Harring & Myers 1924, in Myers 1930,
p.385.

Type locality: Not specified, "common".

Holotype: Not designated

Description: Body slender, fusiform, integument stiff, striated; foot two-segmented;

F~g. 43. Monommata gran~is laterally with dorsal, lateral, and oblique frontal views of trophi.
F~g. 44. Monommata hyalma laterally with oblique frontal and lateral views of trophi. toes variable, but always unequal. At the lateral antennae two rou~:R.9!s.
F~g. 45. Monommata longiseta laterally with dorsal and lateral views of trophi. Mastax intermediate between virgate and forcipate; fulcruffifnmtally short, dagger-
F~g. 46. Monommata maculata laterall~ with oblique frontal, ventral, and lateral views of trophi. like, laterally flat, conical with large, curved basal apophysis; inner margin of rami
F~g. 47. Monommata phoxa laterally WIth ventral and lateral views of trophi.
Fzg. 48. Mo~ommatapseudophoxa with various views oftrophi; (a): pleural plate? (43-45: Myers 1930'
46-47: Harnng & Myers 1924; 48: Wulfert 1960a). '
 28
29
w~th unique, compl~x denticulati.on. There are three ~oups of teeth: a ventral group
Monommata viridis Myers 1937
~Ith 12-.14 comb-lIke teeth; mIddle oral group wIth four large curved, pointed
Fig. 49
mterlockmg teeth; and a dorsal group with three long needle-like teeth. Unci with
three long clubbed teeth; manubrium terminally crooked with ventral strong lamella;
retrocerebral sac present. Related to M grandis. M. viridis Myers 1937, p.10-11, Figs. 12, 19,21.
Body length 210 !lm, right toe 340-470 !lm, left toe 270-410 !-tm, trophus 26-34 !lm.
E~ology: Common, cosmopolitan; in ~ustralia 19-27°C, pH 5.4-6.9, 25-33 I-lS cm-I.
Type locality: Adirondack Mountains, NY, U.S.A. ("common")
LIterature: Koste & Shiel 1991. O~lJ:'tl\ Holotype: Not designated
Code: 13, 16,21,31,42,52,65,73,81,85,92,94.
Description: Body slim, flexible, no neck constriction behind head; dorsal antenna a
double papillose projection; stomach ends in four blind sacs; gut contents yellow-
Monommata phoxa Myers 1930 green globules, gastric glands absent (according to Koste 1978 and Koste & Shiel
Fig. 47 1991, present; in original description expressly designated absent and not shown in
figure either). Mastax modified virgate; fulcrum short, in lateral view curved, taper-
M. phoxa Myers 1930, p. 395-396, Figs. 26:8-10. ing; rami slender, lyrate, bent dorsally near mid-length at approximate right angle;
dorsal portion has small tooth on inner margin; unci reduced to thin lamellar plates,
Type locality: Pond near English Creek, Atlantic County, NJ, D.S.A. posterior edges thickened to resemble slightly clubbed teeth; manubria curve dorsally,
Holotype: Not designated have small digitiform process near mid length; retrocerebral sac characteristic: round,
clear, ductless.
Description: Body shape variable, usually long, cylindrical; single minute dorsal an- Body length 166 !-tIn, right toe 2l61lm, left toe 150 f.1m, parthenogenetic egg 65x52
t~nna; troph~s virgate, with long, distally spatulate fulcrum; rami lyrate in ventral !lm.
VIew; unCI WIth 5-6 linear teeth arising from a common point; two rods run from bases Ecology: Acidophilic, common, cosmopolitan.
of teet~ to tips o~ ran:i, mark.ing the li.mits of two thin lamellar plates resting on sides Literature: Koste 1972b, Koste & Shiel 1991.
of ramI; manubna WIth medIan lobehke projection, distally slightly crutched. Retro- Code: 14, 16,22,32,41,51,61,74,81,84,91,95.
cerebral sac small.
Body leng~h 15~ !lm; right toe 190 !lm; left toe 140 !lm; trophus 35 !-till.
Ecology: ACIdophIl, cosmopolitan but rare. SPECIES INQUIRENDA
Literature: Hauer 1937/1938; Koste & Shiel 1991; Koste & B6ttger 1992.
Code: 14, 16,22,31,42,51, 61, 77, 83, 86,91,94. Monommata dissimile Berzilfs 1949
Fig. 50

Monommata pseudophoxa Wulfert 1960 M dissimile Berzilfs 1949, pA, Figs. 4-8.
Fig. 48 N.B. In a reprint of this paper Berzi.Qs corrected the name by hand to M. dissimilis.

M pseudophoxa Wulfert 1960a, p.289, Fig. 29. Type locality: Lakes Fiolen and Allgunnen, Sweden.
Holotype: Not designated
Type locality: Wildenhainer Bruch, Germany.
Holotype: Not designated Description: Poorly described species. Considered nomen nudum.

Description: Slender, va~able shape with several neck-folds. Trophus resembles that
of M fh~xa. The n:anubna hang on very delicate rods going from the base of the rami Genus Tylotrocha Harring & Myers
to theIr tIp and whIch, pres.umably, are the unci. The five unci teeth of M phoxa are Fig. 51
absent. However, on both SIdes of the trophus there are two roughly triangular striated
plates that ill?Ve a~ound and are perhaps pleural plates. The rami carry three small Tylotrocha Barring & Myers 1922, p.555.
teeth on the nght SIde, two teeth on the left.
Body len~h 14~ p-m, right toe 144 !lm, left toe 105 !lm, trophus 30-32 f.1m. Monotype: Notommata monopus Jennings 1894 = Tylotrocha monopus (Jennings)
E~ology: ACIdophIhc, Germany (not rare at type locality), Russia. 1894, p.14, Figs. 5,6.
LIterature: Kutikova 1970. Type locality: Lake Michigan, D.S.A.
Code: 14, 15,22,31,42,51,63,74,81,86, 91, 94. Holotype: Not designated
30 31

I.1 51

49

Fig. 49. Monommata viridis laterally with frontal and lateral views of trophi.
Fig. 50. Monommata dissimile laterally with ventral, lateral, and oblique frontal views of trophi. (49:
Myers 1937; 50: Berzi.Qs 1949).
53 54

Description: Body soft, fusifonn with two lateral retractable humps; always reddish- Fig. 51. Tylotrocha monopus dorsally and laterally, ~ith ventral and lateral. views of trophi.
purple or rust-coloured. Foot long, conical, deeply wrinkled; conical single toe but Fig. 52. Rousseletia comiculata dorsal and lateral VIews ~f body and trophI
Fig. 53. Dorystoma caudata swimming animals and trophl frontally a~d laterally. .
two footglands indicating that the toe was fonned by the fusion of two toes. Two dor- Fig. 54. Dorystoma caudata "form" furcata swimming animals, trophl ventrally and laterally. (51,52.
sal antennae on papillae near the neck. On the corona two small movable and retract- Harring & Myers 1924; 53,54: Wulfert 1960b).
able palps or auricles carrying tufts of cilia. Gastric glands contain refractile globules;
such globules float in the body cavity and also in the lateral humps. Ganglion large
and circular with a lenticular red body at the posterior end, assumed to be the rudi- Genus Rousseletia Harring
mentary ReO and eyespot. Trophus specialized virgate, all element fused into a
dome-shaped structure. The elongate triangular rami and slender fulcrum are joined;
Rousseletia Harring 1914, p.393
unci and manubria also fonn a unit and show a more or less oval window; the ventral
margin is fused to the rami. The operation oftrophus has not been observed; it may be
a piston as in Lindia. Rousseletia corniculata Harring 1914
Total length 175-250 IlID, toes 18-24, trophus l81lID. Fig. 52
Ecology: Fairly common in circumneutral waters; prefers cool temperatures? North
America, Eastern Europe. Rousseletia corniculata Barring 1914, p. 393, Figs. 37:1-3.
Literature: Barring & Myers 1922, Nogrady 1976.
 32
Type locality: Kenilworth, DC, U.S.A.
Holotype: Not designated
Description: Body stout, somewhat saccate, longitudinally striped; neck well
marked, tail collar-like; foot long with small dorsal seta (may be broken off), toes
short, conical. Corona terminal with two ciliated papillae. RCO with Y-shaped ducts,
cerebral eyespot. Stomach large with blind sacks near mastax; no intestine. Trophus
very large, specialized virgate. Rami large, domed, fenestrated, without denticulation
small thorn-like alulae. Manubria curved rods with ventral branch; fulcrum strongl;
spatulate, spoon-shaped; unci absent. Epipharynx rod-shaped.
Total length 130 !-lm, toes 10 ~m, trophus 40 ~m.
Ecology: Widely distributed but rare, northeastern U.S.A, Australia?
Literature: Harring & Myers 1924, Koste & Shiel 1991.
Genus Dorystoma Harring & Myers
Dorystoma Harring & Myers 1922 p. 555
Dorystoma caudata (Bilfinger) 1894
Fig. 53/54
Proales caudata Bilfinger 1894 p. 46, Figs. 2:3-4.
Type locality: Wurttemberg, Germany.
Holotype: Not designated
Description: Body spindle-shaped with longitudinal striation; transparent, but gut
may be coloured. It swims on its back and is then slender. Neck well marked foot
long, seemingly two-jointed, toes long, sharp. At base of foot over the anus a' con-
spicuous bulb with a thof? ~s long as the toes. Red cerebral eyespot may be 9ccasion-
ally .ab~ent.. Mastax speCIalIzed virgate with long pharyngeal tube. Rami thin thorns
contmumg m a long, thin, thornlike epipharynx that serves to pierce algae. Manubria
ovally curved with a side-arm (Wulfert 1960b and Koste 1978 call them "cheek-
shaped"). Fulcrum cnltched, unci absent.
Total length 130-260 ~m, toes 16-22 ~m, spine 10-22 ~m, trophus 18 ~m pharyn-
geal tube 22· ~m, parthenogenetic egg 56-44 ~m, smooth. ' Description: More or less cylindrical body varying from spindly to stubby or conical.
E~ology: In periphyton, also Sphagnum, rare to frequent; cosmopolitan. Eats algae.
LIterature: Wulfert 1960b, 1961; Koste & Shiel 1991.
Incertae sedis: D. furcata Wulfert 1939b p. 72, Fig. 4
D. caudata " form" furcata (Wulfert) 1939 in Koste 1978 p. 308.
This taxon di~fers from the nominate species by articulated toes, a very long spine
and somewhat dIfferent trophus, smaller size, a segmented foot and lack of eyespot.
suffi~ient criteria for specific status. However, Koste (1978) ob~erved that narcotized
speCImens of D. caudata developed similar toe-shape to furcata and the corona also
changed. He therefore designated D. fucata a "form". Obviously, further observations
are necessary.
33
Total length 140-150 ~m, toes 15-18 ~m, spine 20 ~lm, trophus 14-16 p.m.
Ecology: In acidic ponds.
Literature: Wulfert 1960b, Koste 1978.
Genus Metadiaschiza (Rousselet)
Metadiaschiza trigona (Rousselet) 1895
Fig. 55
Diplois trigona Rousselet 1895 p. 199, Fig. 6:2 = fvfetadiaschiza trigona (Rousselet
1895) according to Fadeyew 1925
M. sculpturata (Daday 1897)
Cephalodella leptodactyla (Hauer 1922)
Type locality: A ditch, England.
Holotype: Not designated
Description: Very closely related to Cephalodella, this taxon differs substantially in
the structure of the lorica which is heavier and granulated. The general shape of the
animal is of Cephalodella character, with a square head, conical foot, long toes ending
in a bristle. The head has seven (?) smooth longitudinal plates; there are two
dorsolateral plates accompanied by two dorsal keels, and one ventral plate. Lateral
sulci wide. Two widely spaced frontal eyespots, no retrocerebral organ. Trophus type
B (Wulfert system), rami without teeth, manubria terminally fanned.
Total length 190-260 f-lm, toes 58-95 f-lm, trophus 23 f-lm.
Ecology: In small ephemeral bodies of water, early spring. Europe, Eastern Asia.
Literature: Wulfert 1937, Kutikova 1970.
Genus Cephalodella Bory de St. Vincent
Cephalodella Bory de St. Vincent, 1826, p. 43.
Type: Cercaria catellina Muller 1786, p.130 = Cephalodella catellina (Muller)
Type locality: Copenhagen.
Head separated from trunk unlike the foot, which bears two toes of varying length and
varying shape. Ratio of total length to toe .length is taxonomically signific~n~. O~ca­
sionally i1loricate, but majority of species has three to five more or less dIstmgUl~h­
able lorica plates. A dorsal and two lateral sulci (furrows, spaces) s~parate the lor!ca
plates. A tailor a cuticular appendage is often present, partly or entlrely o~erlapPI?g
the foot. The corona is usually oblique (tilted ventrally) and convex, sometImes WIth
protruding "lips" or "beaks"; in raptorial species the extensible epipha~nx m~~ pro-
trude from the mouth and look like a beak. The corona bears long margmal CIlIa and
two lateral tufts of swimming cilia, but no auricles ("ears"). Retrocerebra1 organ (ab-
breviated as RCO) absent or present; red or black pigmented "eyespot(s)" can be ab-
34 35

1c 1d
2a 2b

55
[
d4c
[
k

Fig. 55. Metadiaschiza trigona dorsally, ventrally and laterally; trophi dorsally and laterally; optical cross
section. g-k: Rousselet's original figures. (55: Wulfert 1937).
4b 2c 2d
4a

sent, cervical (on central ganglion), or single or double frontal, sometimes with a re-
fractive "lens" or capsule. The virgate trophi, as in all notommatids, are of paramount
-importance for identification, although not always species-specific. They were classi-
fied into six types by Wulfert (1 938a); it is sometimes enough to determine the type
for identification, but in most cases a detailed study of the mastax is necessary; for
new species a scanning electron micrograph is highly desirable. The six types of
trophi are shown in Fig. 56 and are as follows (cf. also Koste & Shiel 1991):
Type A. Rami simple, without teeth on inner margin, with or without alulae (wing-
like extensions); unci simple; manubria thin curved rods without basal lamellae 5a 3a 3b
(plates on basal = proximal end of manubrium) and distal expansions. Fulcrum
Gb
spatulate at distal end both in dorsal and lateral view.
Type B. Rami complex, ribbed or fenestrated (having "windows"), inner margin
toothed or ribbed, sometimes with alulae. Unci simple; manubria with single or dou-
ble basal lamella, distal end T-shaped or crutched, occasionally spatulate or fanned. 56
Fulcrum distally spatulate, sometimes with a basal apophysis, a thornlike apically
Fig. 56. Trophi types of Cephalodella s p e c i e s . . " .
pointed continuation of the proximal end of the fulcrum (or rami, where it meets the 1: Type A, (a) ventral; (b) lateral; (c,d) variant. 2: Type B, (a) ventral; (b) lateral; (c,d) vanant. 3. Type C, (a)
fulcrum). ventral; (b) lateral. 4: Type D, (a) ventral; (b) lateral; (c) uncus, lateral. 5: Type E, (a) ventral; (?) lateral.
Type C. A variant of type B: trophus asymmetric, and the distal crutch of manubria 6: Type F, (a) ventral; (b) lateral. Scale bars 10 I-lm. (after Wulfert 1937, 1938a and Koste & Shlel 1991).
is closed to a loop. On occasion it is difficult to see whether the loop is closed or not.
Loops with small breaks still belong to this type.
Type D. Very complex, with unique parts. Rami with terminal teeth, inner margin Type E. Known only in C. megalocephala, the rami are bent at right angle
with numerous comb-like teeth. Unci consist of a dorsal plate with a single tooth; (dorsally, when seen laterally); manubri.a S-sh~pe~ with lamellae that are separa~e.
subunci, fanned broomlike structures, are found behind the manubria. Manubria Type F. Known only in C. mira, ramI look h~e mner basal lamellae of the ~ehcate
distally abruptly curved, with wide basal lamellae. Fulcrum spatulate, proximal end rod-like manubria and are attached to the proxImal top of the equally rod-hke ful-
often widened, basal apophysis common. In some species (forficula, gigantea, tenui- crum. The whole fragile structure is presumably only a pump.
seta) a thin denticulate plate occurs above the rami.
36 37

Cephalodella is probably the most difficult genus among all rotifers because of the 05 with asymmetrical alulae
great similarity of body shapes, and also because of the large number of species (about 06 no teeth on inner edge
190 taxa are discussed in the present compilation). Thus, misidentification is a very 07 with teeth on inner edge
real danger. As mentioned in the Introduction to this volume, due to the historical pau- 08 left ramus with terminal tooth/teeth
city of workers outside Europe a large number of species (e.g., those of Myers) have 09 frontal pegs
not been seen since description, compounded by the fact that even Myers failed to
provide figures of trophi in many cases. All the precautions outlined in the Introduc- 05 MANUBRIA
tion to this volume are thus especially pertinent to this genus, as Cephalodella species
never occur in as large numbers as planktonic taxa do, and the usually sparse study 01 dorsally bent
material is therefore very valuable. Their biology, ecology, variability, and biometrics 02 ventrally bent
are not well known and deserve special attention and further study (cf. Pejler & 03 rod-like, no terminal widening
Berzit;ls 1993). 04 end crotched
05 end forked
06 end fan-like
CEPHALODELLA DESCRIPTORS 07 end with median loop (hole)
08 end with terminal loop
01 TOTAL LENGTH/TOE LENGTH RATIO 09 asymmetric
10 bent in plane, semicircular
01 <3 11 other
02 3-5
03 >5 06 FULCRUM
02 EYE 01 with basal apophysis
02 without basal apophysis
01 absent
02 single 07 PLEURAL ROD
03 double
04 cervical 01 present
05 frontal 02 U, Y or V shaped pleural apparatus
06 colourless
07 red 08 TROPHUS SIZE
08 with lens
01 <30 ~Lm
03 TROPHUS TYPE of Wulfert 02 30-70!-Lm
03 >70!-Lm
01 A
02 B 09 BODY SIZE
03 C
04 D 01 <200!-Lm
05 E 02 200-300!-Lm
06 F 03 >300!-Lm

04 RAMI 10 BODY SHAPE

01 symmetrical 01 stout
02 asymmetrical 02 slender
03 without alulae 03 elongate
04 with sylnmetrical alulae 04 conical
 38
39
05 faintly gibbous
06 gibbous 04 forked
05 with claw
07 laterally compressed
08 bulbous 06 bristle-like
09 fusiform 07 blunt
08 with septum, or bendable
11 TAIL
16 CORONA
01 short
02 as long as foot
01 with lips
03 longer than foot
02 with rostrum
04 with double keel
03 with styli or palps
05 with short upright hook
04 dorsally wider than body
06 caudal antenna with bristles
05 moderately oblique
06 oblique
12 FOOT
07 convex
08 other
01 in body axis
02 ventrally displaced
17 VITELLARIUM
03 segmented
01 4-6 nuclei
13 TOES
02 8 nuclei
03 12 nuclei
01 slim
02 stout
18 GLANDS
03 base bulbous
04 middle wide
01 salivary glands absent
05 with segmentation or bendable
02 salivary glands conspicuous
06 straight
03 retrocerebral organ absent
07 dorsally bent
04 retrocerebral organ present
08 ventrally bent
05 retrocerebral organ brown
09 S-shaped
10 conical 06 subcerebral glands present
07 gastric glands red/brown
1 I divergent
08 gastric glands conspicuous
12 w~th dorsal appendage
13 WIth dorsal denticles
19 MISCELLANEOUS
14 with ventral denticles
01 zooch10rellae present
14 TOE SIZE
02 parasite
03 acidophilic
01 <20!lm
04 inhabits moss/terrestrial
02 20-60 (.tm
05 rheic
03 >60!lm
06 saltwater, brackish, marine
07 psaIDlTIophilic
15 TIP OF TOES
N.B. DUE TO THE VERY LARGE NUMBER OF CEPHALODELLA SPECIES THEIR DESCRIPTIONS
01 tapered evenly, gradually ARE, EXCEPTIONALLY, LISTED ALPHABETICALLY INSTEAD OF IN THE ORDER IN WHICH
02 tapered abruptly (German: "abgesetzt") THEY KEY OUT.
03 recurved
40 41

I~r~ ~i:.~#t:~:~: : ~: : : : : ;: : ~: : : : :~: : : : : ~ : : ;: : : : ;: : ~: : : ~: : : :IJ .

KEY TO CEPHALODELLA

N.B., SPECIES MARKED WITH AN ASTERISK HA YE NOT BEEN SEEN SINCE THE ORIGINAL DE-
SCRIPTION.
3(2 ).
LONG TOES, totallengthltoe length <3 page 40 Toes (viewed dorsally) form a rigid lyre pseudeva *
MEDIUM TOES, total length/toe length 3-5 ~ page 40 4(3). Tip of toes sickle-shaped, dorsally curved apocolea
SHORT TOES, totallengthltoe length >5 page 43 Tip of toes lanceolate, straight astricta
Tip of toes ventrally curved 5.
5(4). Curvatures just a blip; manubria asymmetrical rostrum *
LONG TOES, totallength/toe length <3 Curvature pronounced; manubria symmetrical 6.
6(5). Body slender but not conspicuously long rzcznza
.. *
1. Eyeless 2. Body extremely elongate, lorica rigid praelon~a*
Single cevical eyespot 5. 7(3). Lips present, gastric glands reddish lepzda
2(1). Trophus type A 3. Lips absent, gastric glands colourless oxyda.ctyla
Trophus type B 4. 8(3). Body conical; tip of toes with septum spe~·LOsa*
3(2). Toes bendable, reo. and V-shaped pleural rod present.. gobio Body gibbous; lateral antennae on conical tubules , , papzllosa *
None of the above, toes saber-like maerodaetyla Body fusiform 9.
4(2). Abdomen with hooked long tail; toes with single tip mucronata 9(8). Corona with prominent lips; base of toes normal. eurynota
Tail absent, toes with double tip biungulata Corona without lips; base of toes bulbous melia*
Toe with bulbous base, single tip planera * 10(2). One or both gastric glands red-brown 11.
5(1). Trophus type A 6. Gastric glands colourless : 13.
Trophus type D; giant form, toes thin, undulating gigantea 11 (10). Manubria asymmetric, ending in a fan for/icata
6(5). Toes straight 7. Manubria symmetric and crotched : : ~2.
Toes dorsally bent 9. 12(11). Toes ending in claw; inhabits spherical casing znq~zlzna
Toes ventrally bent 10. Tip of toe with septum; rami with teeth zntuta
Toes S-shaped, ; hyaline area at lateral antennae subsecunda * Toes ending in bristle coliactea~
Toes S-shaped, no hyaline area; foot with long sensory bristle 13(10). Toes S-shaped, blunt tip : paxz
................................................................................................ psammophila * Toes almost S-shaped; long, thin, ventr~. l~ ~urved t~~i.1A ~ eva
7(6). Toes with slender abruptly upturned tips euknema Toes straight or ventrally bent r•• D;f..()~tJ.f!:A. cki.~:~~ :. ~4 ..
Tips otherwise 8. 14(13). Toes thick, compressed dorsoventrally, abru'Ptly tapered : gzslen~
8(7). Body conical; tip of toes uniformly tapered belone Toes with four spinules at ventral distal end lzndamaya

9(6).

10(6).
Body gibbous; tip of toes abruptly tapered
Body laterally compressed
Head small and narrow, body compressed
Head large, body compressed
Body stout, prominent beak on corona
dorseyi*
compressa *
tantilia
sabulosa
tachyphora
IS( 14).
16(15). With prominent beak; four salivary glands boettgerz
t
~~~: ~~~e:s:~::::::: : : : : : : : : : : : : : : : : : : : : : : : : : : : : :: : : : : : : : : : : : : : :
No beak; number of salivary glands unknown vaeuna* p.146
Body slender, no beak elegans 17(15). Rami without teeth on inner edge 18.
Left ramus with teeth; caudal antenna present asta
Both rami with teeth 1.9.
MEDIUM TOES, totallength/toe length 3-5 18(17). Rami with long symmetrical alulae bry~phzla
Alulae absent lzmosa
1. Eyeless 2. 19(17). V-shaped pleural rod present; manubria crutched ·..· 20.
Cervical eyespot(s) 24. Pleural rod absent : 21.
Frontal eyespot(s) 36. 20(19). Distal end of ganglion with unpigmented crystal; manubna fanned .,
2(1). Trophus type A 3. (spatulate) r.lgz~a
Trophus type B 10. Crystal absent; manubria crutched Ine.z a
Trophus type C 22. 21 (19). Gastric glands with stalk; toes usually with septum hyahna
 42
43
Gastric gland normal; toes abruptly tapered; rami with alulae, manubria 41(40). Manubria with bump in middle; gastric glands filled with shiny fat globules
asymmetric , dentata ........................................................................................................ gibboide.'.;
22(2). Toes with claws, manubria with terminal ring paggia* Manubria smooth 42.
23(2). Toes slender, long, bulbous at base tenuiseta 42(41 ). Manubria asymmetric, rami with pseudoalulae graciosa
Toes thick, decurved, with a dull tooth dorsally stenroosi Manubria symmetric, rami without alulae carina*
24(1). Trophus type A 25. 43(37). Very slender cylindrical body, corona not oblique elongata
Trophus type B 35. Body normal or stout 44.
25(24). Toes more or less ventrally curved 26. 44(43). Body very stout, foot ventrally bulged, tail long; crutch on manubria
Toes straight or dorsally curved 33. characteristic, fulcrum spatulate reimanni
Toes S-shaped, conspicuous caudal antenna nana Stout, tail short, fulcrum not expanded innesi*
26(25). Body elongate, pleural rods present hoodi Body normal, tail short; manubria long and slender mineri*
Body slim, narrowing distally, pleural rods absent exigua 45(36). Toes straight 46.
Body bulbous, stout; pleural rods absent 27. Toes dorsally bent 47.
27(26). With zoochlorellae; sulci very deep plieata 46(45). Toes long, slender; manubria with closed rings, without lamellae; five lorica
Without zoochlorellae 28. plates nzisgurnus
28(27). Vitellarium with 12 or more nuclei licina Toes more conical with septum or claw; manubria with open rings and
Vitellarium with fewer (usually 8) nuclei 29. lamellae tecta
29(28). Rami with huge alul~el. ) ..I glypha* 47(45). Both rami with teeth; manubria end in a spoon hol101,vdayi
Alulae smaller .. Or. @~:;' ~ 30. Only right ramus with teeth; asymmetrical manubria end in open rings .
30(29). Manubria with median hole 31. ............................................................................................................... obvia
Manubria without median hole 32. Rami with frontal pegs; symmetric manubria end in closed rings paehyodon
31(30). Tail hides ventrally shifted foot mus 48(36). Dorsal surface of short toes with a row of spicules forficula
Foot not ventrally shifted; toes with ventral cusp .. Dorsal side of long toe occasionally with a single tooth; tail very short ........
.......................................................................... physalis (European ecotype) . panarista
32(30). Toe with ventral cusp physalis (N. American ecotype) Toe without tooth; large keeled tail covers foot. tinea
Toe smooth; one dorsal and one ventral cerebral eyespot zeteta Toe without tooth; trophus different from tinea tinea/ormis
33(25). Toes uniformly straight, long, blade-shaped; head same width as body
.......... ~ ··.· mucosa*
Toes with bulbous base, somewhat ventrally curved. Head in dorsal view SHORT TOES, total length/toe length >5
pentagonal, wider than body labiosa
Toes dorsally curved, with septa 34. 1. Eyeless 2.
34(33). Very gibbous, laterally compressed tantilloides Cervical eyespot(s) 20.
Less gibbous, but stout derbyi p. 145 Frontal eyespot(s) 30.
35(24). Toes long, 1/3 total length, dorsally curved hiulea
i~~~~~~ ~: ~ ::::::::::::::::::::::::::::::::::::::::::::::::::::::::::::::::::::::::::::::::::::::::::"1 ~:
2(1).
Toes short, 1/4 total length, straight viteZZa
36(1). Trophus type B 37. Trophus type C; free living jluviatilis
Trophus type C 45. Trophus type C; parasitic in Uroglena .:................................................ edax
Trophus type D 48. Trophus type E : ~ 8.
37(36). Toes straight or only slightly curved 38. 3(2). Parasitic on oligochaetes parasltlea
Toes dorsally bent
Toes ventrally bent
38(37). Laterally strongly compressed, base of toe bulbous............................. laisi *
40.
43. 4(3). ~~~t::~~~~~
Toes ventrally curved
.: : : : : : : : : : : : : : : : : : :: : : : : : :: : : : : : :: : : : : : :: : :: : : : : : : : : : ~:
: 7.
Not compressed, rami withalulae 39. 5(4). Body extremely squat (length/depth ratio 1.6/1) lzpara
39(38). Both rami with alulae, long tail with appendage paehydaetyla Body normal or slim : 6.
Large alula on left ralTIUS only, five lorica plates pentaplax 6(5). Body very elongate; corona not oblique, but continues on ventral sIde. of
Small alula on left ramus, three pleural rods sterea head' toes conical asarCla *
40(37). Total length >250 Ilm gibba Bod; normal, corona oblique, toes blade-shaped dora
Total length <250 Ilm 41. Toes pencil-shaped, appressed, appearing as a single toe dixon-nuttalli*
 45
44

Corona extremely oblique pheloma Dorsal plates cover ventrally displaced foot and toes tenuis*
7(4).
Corona nOlwaIly or only slightly oblique 8. 24(23). Head wider than rump; conspicuous salivary glands and caudal antenna.
Base of toe bulbous 9. Common auriculata
8(7).
Toes of uniform width, tapering abruptly to bristle-like points ablusa* Head not as wide, no caudal antenna; manubria very thin, curved, with swol-
Toes short, ventrally curved; gastric glands size of mastax dorystoma len tips. Acidophil doryphora
Toes slender, tapering uniformly nelitis* 25(22). Eyespot colorless, gastric glands the size of the mastax dorystoma
9(8).
Toes constrict past bulbous base and expand again, to form sharp tips gradu- Eyespot pigmented, gastric glands normal ·.. ·.. ·· 26.
ally celeris 26(25). Tail longer than foot, with double keel arcuata
Tail shorter than foot 27.
1'0(2). Terrestrial, in moss, manure, and leaf-litter 11.
Aquatic only 12. 27(26). Rami with large alulae delicata
11 (l0). Manubria asymmetric, Y-shaped pleural rod dara Rami without alulae. Baikal endemity vittata*
Manubria symmetric, no pleural rod; crystal at end of ganglion rotunda 28(20). Toes ventrally curved, tapering uniformly ·· calosa
Toes curved, abruptly tapered 29 .
Trophus identical to rotunda; crystal absent rot. bryophila
29(28). Manubria crutched ............................................................................... •y:enica
12(10). Parasitic species 13.
Manubria not crutched, with median and terminal loop transition between
Free-living species 14.
trophus type Band D theodora
13(12). On gills of decapods; toes blunt, tapered abruptly crassipes
In the colony of the alga Coelosphaerium kiitzingianum; gastric glands with
vacuole; toes conical abstrusa
30(1).
i~~~~~:~~: ~ : : :: : : : : : : :: : : :: : : : : : : : :: : : : : : : : :: : : : :: : : : : : : : : : : : ;~:
Trophus type C. Foot shifted ventrally, tail much longer than foot; loops on
14(12). Manubria asymmetric; brown granular mass between the large gastric glands
manubria can be open. Variable species ., catellina
.................................................................................................... glandulosa *
Trophus type F. Cylindrical body, stomach with green granules mira
Manubria symmetric 15.
15(14). Manubrium with simple crutch 16. 31 (30). Marine or brackish habitat; toe slightly bent ventrally or straight ..
.. (epitedia) rnarina *
e, - Manubrium with a fan (spatulate) jorficata Freshwater species 32.
Manubrium with a combination of crutch and half-loop evabroedi*
32(31). Body bulbous , head narrower than body, . corona not convex compacta* *
16(15). ~~mi with teeth, U-shaped pleural rod; toe with septum; 1-2 crystals in gan-
Body more elongate, toes dorsally curved, total length <100 ~m angu~ta *
g Ion tenuior Body elongate, toes straight, total length >21 0 ~m paxzlla
Toes without septum, no crystals 17. Vitellarium with 4 (sometimes 6) nuclei. Very variable species, common ...
17(16). Rami without teeth, V-shaped pleural rod; frontal edge of head-lorica thick- 33(30)
............................................................................................................ gracilis
ened to a bead..................................................................................... wrighti Vitellarium with 8 nuclei 34.
Trophus ~ot fig~red; long cylindrical body with conspicuously bulging Body cylindrical, head tilted at an almost 90 0
angle tempesta
corona wIthout lIps montana * 34(33).
Body stout or fusiform 35:
Trophus not figured; short and stout body, bulging corona with lips . Body stout, head large, edge of head-Iorica thickened; pseudoalulae on ramI
35(34).
B~d~·~ii;;;::::::::::::::::::::::::::::: . ::::::::::::::::::..:: :::::::::::::..:::::::::::: :..::: ~~o.~~~
.......................................................................................................... poitera*
18(2). ~ody bUlbo.u.s, foot ventrally shifted, small straight toes, manubria with open
nngs (tranSItIon between types B and C) jluviatilis
36(35). Rami asymmetric, small triangular alula on left sterea
19(2). Foot 2-segmented, toes bendable but septa not always observable . Rami sylnmetric 37 .
............ n1egalocephala Large thornlike alulae; two large touching eyes in common capsule .
Foot and toe 2-segmented; toe consists of a cylindrical part and a tenninal 37(36).
. cyclops
needle-shaped part minora Rami with blunt alulae; eyes separated; toe with vacuole jorceps
20(1). i~~~~~~ ~~: ~ ~::::::::::~:::::::::::::::::~:::~:::~:~::::~:::~~::~::::::::~:~:::::::::::~::::::::::::::;~: Not included in key: C. conjuncta Myers 1940; C. eu~oma Myer~ 19~0; C. jriebei
21 (20). Toes straight, small thorn-like alulae on rami euderbyi Koste et al. 1984; C. retusa Myers 1924. With the exceptIOn of C. jrzebez, (p. 77) these
Toes ventrally curved 22.
are described among species inquirendae on p. 144.
22(21). Body stout, short 23.
Body more elongated· 25. N.B. DUE TO THE VERY LARGE NUMBER OF CEPHALODELLA SPECIES THEY ARE, EXCEPTION-
23(22). Ta~l covers foot; beak and epipharynx present; single cervical eyespot 24. ALLY, LISTED ALPBABETICALLY, NOT IN THE ORDER IN WHICH THEY KEY OUT.
Talll.onger than foot; foot usually ventrally displaced; no epipharynx, double
cervIcal eye ventripes
46 47

Cephalodella ablusa Myers, 1934

Fig. 57

Cephalodella ablusa Myers, 1934, p. 8-9, Fig. 8

Type locality: Round Pond, Mount Desert Island, ME, D.S.A.

Holotype: Not designated

Description: Body extremely elongate, very slender, laterally compressed. Head

relatively short, neck indistinct. Lorica soft, indistinct, foot tapering gradually. Toes
tapering abruptly to bristle-like tips. Foot-glands large, elongate. Corona oblique,
strongly convex, without lips; salivary glands present; large teardrop-shaped gastric
glands. Ganglion nonnal, no eyespots. Trophus type A: ?(not figured by Myers), stout
fulcrum enlarged posteriorly, slender recurved manubria, slightly swollen at the tip.
Resembles C. elongata Myers 1924, but differs by absence of eyespots, presence of
salivary glands and shape of toes.
Total length 148 Ilm, toes 25 Ilm. 58
Ecology: Among Fontinalis in the summer, pH about 6.4.
Literature: Not seen since description. 57
Code: 0103, 0201, 0301, 0401, 0501, 0901, 1002, 1003, 1007, 1101, 1201, 1308,
1402, 1506, 1605, 1607, 1802, 1808, 1903.

Cephalodella abstrusa Myers, 1934

Fig. 58

Cephalodella abstrusa Myers, 1934, p. 13, Fig. 14.

Type locality: Upper Hadlock Lake, Mount Desert Island, NIE, D.S.A.
Holotype: Not designated

Description: Body stout, cylindrical, dorsally arched, ventrally straight. Head small;
[
neck distinctly constricted. Lorica soft, but plates well defined; foot and tail short.
Toes short, straight, conical, ending in blunt tips. Corona oblique, strongly convex, 61
lips not visible. Gastric glands large, oval, containing a refringent vacuole in every 62
individual; ovary large. No eyes. Trophus type A: ? (B ? according to Wulfert 1938;
not figured by Myers) fulcrum long and staight, manubria longer than fulcrum, ex- 59
panded posteriorly.
Total length 104 Ilm, toes 12 Ilm.
Ecology: Parasitic in the planktonic alga Coelosphaerium kiitzingianum Naegeli. Was Fig. 57. C. ablusa.
abundant at type locality. Fig. 58. C. abstrusa.
Literature: Wulfert 1938. Fig. 59. C. akrobeles.
Fig. 60. C. anebodica laterally, toes, and trophus.
Code: 0103,0201,0302,0401,0901, 1001, 1101, 1201, 1306, 1310, 1401, 1501, Fig. 61. C. angusta.
1605, 1607, 1808, 1902. Fig. 62. C. apocolea laterally, dorsally, (c): trophus laterally; (d): fulcrum laterally; and (e): toes.
(57,58,59 Myers 1934; 60: B-erzinGs 1976; 61: Barring & Myers 1924; 62: Wulfert 1940).
 48
Cephalodella akrobeles Myers, 1934
Fig. 59
Cephalodella aA.Tobeles Myers, 1934, p.11-12, Fig. 12
Type locality: Barcelona Creek, Mount Desert Island, NIE, D.S.A.
Holotype: Not designated. Cotype AMNH 684
Description: Body elongate, slender, slightly arched. Head loricated, neck well
marked; 10rica plates well defined, lateral sulcus wide. Foot and tail short. Toes
widely apart at base, taper gradually to fine sharp tips. Gastric glands large, pyriform;
retrocerebral organ [RCO] present; no eyespot. Trophus type A: ? (not figured by
Myers), manubria slender, abruptly curved ventrally at end; fulcrum straight, ex-
panded posteriorly. According to Myers (1934), forms a group with C. praelonga
Myers and C. strepta Myers (=macrodactyla) with stiff loricae and head-shields,
elongate body, no eyespots, bacillar (rod-shaped) manubria. The toes of macro-
dactyla are very different.
Total length 120 !-tm, toes 20 !-tm.
Ecology: Among decaying Utricularia at type locality, also found in Atlantic County,
NJ.
Literature: Wulfert 1938a.
Code: 0103, 0201, 0301, 0401, 0502, 0901, 1002, 1003, 1101, 1201, 1307, 1402,
1501,1605,1804,1903.
Cephalodella anebodica (Berzivs), 1976
Fig. 60
Cephalodella anebodica Berzivs, 1976, p.ll, Figs. 18-20.
Type locality: Lake Stnlken, Sweden.
HoIotype: Not designated
Description: Body stout, laterally somewhat compressed, slightly gibbous. Head
large, neck indistinct. Foot short, toes long, slender, dorsally curved v.rith distal swell-
ing and sharp tips. Corona not oblique, convex. RCO present, single frontal eyespot.
Trophus type B: symmetrical, fulcrum stout, with a deep hollow channel on both
sides; rami consisting of two parts, no alulae; manubria carry lamellae on both sides,
crutche?; ~he dorsal end of the crutch carries a sharp hook. The proximal end of
manubna IS flat, smooth. Resembles C. gibba, but has shorter toes and different
trophus. Listed also under C. gibba anebodica.
Total length 250-270 !-tm, toes 64 !-tm, mastax 75 !-tm, fulcrum 45 !-tm , manubrium
42!-tm.
Ecology: Rare in the summers of 1952, 1958 and 1963. Lake Straken is an oligo-dys-
trophic lake (see Naumann 1932, p.116.)
Literature: Koste 1978.
Code: 0102,0202,0205,0302,0401,0403,0504,0803,0902, 1001, 1005,
1201,1301,1307,1403,1503,1701,1804.
49
Cephalodella angusta Myers, 1924
Fig. 61
Cephalodella angusta Myers, Harring & Myers, 1924, p. 467, Fig. 27.2
Type locality: Oceanville, NJ, D.S.A.
Holotype: Not designated
Description: Body small, stout, gibbous, head very large, neck indistinct. Lorica flex-
ible but plates distinct, sulci diverge distally. Foot small, conical, tail very small; toes
1/6 of total length, short, conical, slightly recurved dorsally, tapering to a sharp tip.
Corona strongly oblique, convex, no lips. Gastric glands small, ganglion very long,
pyriform, RCO absent. Double frontal eyespots very close together. Trophus type A:
fulcrum slightly expanded, manubria strongly decurved but not crutched (not illus-
trated).
Total lengths 90-95 !-tm, toes 15-18 !-tIll.
Ecology: Rare; among Riccia and floating Sphagnum in an acid pond.
Literature: Not seen since discovery.
Code: 0103, 0203, 0205, 0301, 0501, 0801, 0901, 1001, 1101, 1201, 1308, 1310,
1401,1501,1503,1606,1607,1803,1903.
Cephalodella apocolea Myers, 1924
Fig. 62
Cephalodella apocolea Myers, Harring & Myers, 1924, p.509, Figs. 33:1,2.
Type locality: Not specified, "common".
Holotype: Not designated
Description: Body very transparent, elongated, cylindrical, slightly compressed lat-
erally. Head short, large; lorica thin but well marked, with lateral sulci. Foot robust
with small tail; toes well separated at base, ending in dorsally recurved sickle-shaped
tips; toes often crossed in swimming animals. Footglands large, pyriform. Corona
wide, ~bliqu~, without lips.. RCO and e)fjes ab~ent. T~~~." hus modified type A:
manubna delIcate, fulcrum slIghtly bent. ~O'l'! flMM-f '-v tu/~.'
Total length 125-185 !-tm, toes 32-58 !-tm, trophus 29-33 !-tm .
Ecology: Common in acid-neutral waters among plants, periphyton, psammon, cos-
mopolitan.
Literature: Harring & Myers 1924, Wulfert 1940, Kutikova 1970, Koste 1976, Koste
& Shiel 1991.
Code: 0102, 0201, 0301, 0802, 0901, 1002, 1007, 1101, 1201, 1306, 1402, 1503,
1506, 1605, 1607, 1803.
50 51

Cephalodella arcuata Wulfert, 1937

Fig. 63

Cephalodella arcuata Wulfert, 1937, p.601-603, Fig. 12.

Type locality: Not specified, Germany.

Holotype: Not designated

Description: Body slender, tapering distally, curved in lateral view. Head wide,
short; neck indistinct; lorica well marked. Foot tapered; very large tail longer than
foot, has double keel viewed dorsally. Toes curved ventrally and tapering to fine tips.
Corona straight, slightly convex, two lips. Salivary glands present; gastric glands very
large. Double cervical eyespots. Trophus type A, delicate.
Total length 160-220 !lm, toes 32-34 !lm, trophus 27 !lm. 63
Ecology: In ditches, presumably alkaline pH. 64 65
Literature: Kutikova 1970, Koste 1978.
Code: 0103,0203,0204,0301,0501,0503,0801,0901, 1006, 1103, 1104, 1201,
1308, 1402, 1501, 1601, 1808.

Cephalodella asarcia Myers, 1942

Fig. 64

Cephalodella asarcia Myers, 1942, p. 27-276, Fig. 5. Ovl ~:ede:

iu ~'y~rs If(t); p fl'-1/'~ / i {

'Type locality: Twin!:-_~kesrPA, V.S.A. np$ -/1 - (Q.l/l/tOqUO w:.<e •

Holotype: Academy of Natural Sciences of Philadelphia. Not shown in recent listing.

Description: Body very elongate and parallel sided. Head long, convex anteriorly;
lorica thin, flexible; plates ill-defined. Foot robust, conical. Toes straight, diminishing
gradually to fine tips; foot glands large. Corona frontal anteriorly but extending from 68
there down the ventral side one quarter of the entire length of body. Gastric glands
small, oval; ganglion extremely long, saccate; no eyespots. Trophus type A: manubria
slender, straight. Resembles C. elongata, but lacks the double eyespot, and is distin-
guished by the unique corona.
Total length 95 !lm, toes 15 /-lm, trophus 15 /-lm.
Ecology: Among floating Myriophyllum humile in a dammed pond, Pocono Plateau,
in the summer, pH 4.
Literature: Not seen since discovery.
Code: 0103, 02eN, 0301,0801,0901, 1002,1003, 1201, 1310, 1401, 1501, 1608,
1903.
c d
Fig. 63. C. arcuata laterally, dorsally, and trophus.
Fig. 64. C. asarcia.
Cephalodella asta Donner, 1970 Fig. 65. C. asta laterally, trophus dorsally, and manubrium laterally.
Fig. 65 Fig. 66. C. astricta.
Fig. 67. C. auriculata; for trophi see Fig. 56: 1. Scale bar 50 !lID.
Fig. 68. C. belone. h)
Cephalodella asta Donner, 1970, p.207-208, Figs. 6 a-c. Fig. 69. C. biungulata (a): laterally; (b): tip of toe; (c and d): trophus ventral and lateral; (e): uncus; (f,g, :
ramus from different directions; (i): toes ventrally. (63, 67: Wulfert 1937; 64: Myers 1942; 65: Donner
1970; 66: Myers 1934; 68: Myers 1924; 69: Wu1fert 1937).
52
Type locality: River Salzach, Austria.
Holotype: Not designated
Description: Body slender, parallel-sided, straight. Lorica distinct. Tail covers foot
entirely. Toes long, straight, ending in a bristle, caudal antenna with tactile bristles
above toes. Corona not oblique, convex, no lips. No salivary glands; stomach sepa-
rated from intestine; vitellarium with eight nuclei. Ganglion long, foamy; no eyespot.
Trophus type B, asymmetrical: left ramus has strong terminal teeth, no alulae;
manubria crutched with lamellae on both sides. Differs from C. intuta and vacuna by
the long bristles at the end of staight toes; C. intuta has a reddish left gastric gland. The
toes of C. eva also end in bristles, but the toes are tapered and the animal is much
larger.
Total length 140-149 ~m, toes 40 ~m, trophus 20-21 !lm.
Ecology: In submerged Hygrohypnum moss in the River Salzach, pH 7.5 in SUlnmer.
Rare.
Literature: Koste 1978. Not seen since discovery.
Code: 0102, 0201, 0302, 0402, 0408, 0504, 0801, 0901, 1002, 1102, 1106, 1201,
1306, 1402, 1506, 1702, 1801.
Cephalodella astricta Myers, 1934
Fig. 66
Cephalodella astricta Myers, 1934, p.9-10, Fig. 9.
Type locality: Half Moon Pond, Mount Desert Island, ME, D.S.A.
Holotype: Not designated
Description: Body elongate, slightly gibbous, much compressed dorsoventrally i.e.,
nearly twice as broad as deep. Head relatively small; lorica thin but well defined. Foot
very short, stout; toes long, slender, lanceolate; if viewed dorsally, tips slightly
outcurved. Corona not oblique, strongly convex, no lips. Ganglion long; no eyespots.
Trophus small, manubria slender, slightly recurved (type A ?). Moves very fast. The
dorsoventrally compressed body and lanceolate toes distinguish this species.
Total length 155 ~m, toes 35 flm.
Ecology: pH 6.6-6.8
Literature: Not seen since discovery.
Code: 0102, 0201, 0301, 0501, 0802, 0901, 1002, 1003, 1101, 1201, 1306, 1402,
1501, 1607, 1803.
Cephalodella auriculata (O.F. Miiller), 1773
Fig. 67
Vorticella auriculata Miiller, 1773, p. 111.
for synonymy see Harring & Myers 1924, Koste 1978
Type locality: Copenhagen.
53
Holotype: Not designated
Description: Short and stout; dorsally viewed, head wider than trunk, with small ros-
trum. Lorica rigid, plates distinct. Foot and toes short; tail and caudal setae long. Sali-
vary glands large, round. Ganglion large; one cervical eye. Trophus type A: slender
recurved manubria, long fulcrum. Moves rapidly, sometimes oscillates attached to a
filament. Resembles ventripes in habitus which, however, has double cervical eye-
spot.
Total length 120-160 !-tm, toes 22-28 ~lm, trophus 36 ~tm. Male 95 ~tm.
Ecology: Cosmopolitan and frequent in the littoral and psammon. pH 6.4-7.3, temp.
16-22°C, 57-274 !lmS cm-I.
Literature: Harring & Myers 1924, Kutikova 1970, Koste 1978, Koste & Shiel 1991.
Code: 0103, 0202, 0204, 0301, 0501, 0802, 0901, 1001, 1102, 1106, 1201, 1302,
1308,1311,1402,1501,1602,1604,1607,1802.
Cephalodella belone Myers, 1924
Fig. 68
Cephalodella belone Myers, Harring & Myers 1924, p.490-491, Fig. 31:2
Type locality: WI and NJ, D.S.A.
Holotype: Not designated
Description: Body small, conical; head very large, one third length of entire body,
somewhat wider than abdomen; neck well marked; abdomen tapers rapidly. Lorica
fairly rigid, plates wellmarked; lateral clefts very narrow, parallel-sided. Foot moder-
ately long, conical; tail small; toes extremely long, straight and slender, slightly en-
larged basally, tapering very gradually to minutely rounded tips. Corona somewhat
oblique, convex, with prominent beaklike lips; gastric glands small; ganglion long;
eyespot cervical; no RCO; mastax type A: fulcrum relatively stout, slightly expanded
posteriorly; manubria slender, rodlike, slightly recurved, but not crutched. When
swimming, the toes are never separated, resembles a diminutive Trichocerca,' when
death occurs, the toes are thrown out sideways, thus resembling a small Monommata.
Resembles nana but the toes are different.
Total length 120-125 ~m, toes 45-50 flm.
Ecology: Among Fontinalis .and. in a bog pool at ~argaintown, near Atlantic City, I
New Jersey. u1l- 6~l,flV1Q~tf ' I[~;r IL{eiL\i,~§,
u ....,,-
(Ieef ({-\.VSW .loo; I (.,v1 f0utl ~ 0 f r..Z' O-I(1.-v/;t'" . ,.~.,'71 ~
.• . .
.-..J liP...,,,, tJ'1
Literature: Berzins 196 . .- v p • ":v
Code: 0101,0204: 0301,0901, 1001, 1004,1101, 1201,1306, 1507, 1601,1903.
Cephalodella biungulata Wulfert, 1937
Fig. 69
Cephalodella biungulata Wu1fert, 1937, p. 617-618, Fig. 26.
Type locality: Germany.
 54 55
Holotype: Not designated

Description: Body hyaline, oval, dorsally arched; head short, slightly oblique; abdo-
men projects over short foot, both covered by dorsal plates; toes long, flexible with
distinctive bifurcate tips due to presence of spinule at distal end. Mastax lacks salivary
glands; trophus type B: symmetrical rami denticulate on inner margin; manubria
crutched, with bilateral proximal lamellae; shaft of uncus with semicircular lamella.
Eyes absent. Close to C. gibba, but distinguished from it by the lack of eyes, charac-
teristic longer bifurcate toes. b
Total length 250-313 !lm; toes 88-112 !-tm, trophus 50-54 !-tm.
Ecology: Rare in littoral/moss of pools and streams in Europe South America, Aus-
tralia.
Literature: Wulfert 1943, Kutikova 1970, Koste 1986, Koste & Shiel 1991.
Code: 0101, 0201, 0302, 0407, 0504, 0802, 0902, 1005, 1102, 1201, 1306, 1403,
1504, 1605, 1801.

Cephalodella boettgeri Koste, 1988

Fig. 70 70 f
Cephalodella boettgeri Koste 1988a, p.313-3l4, Figs. 2 a-c, 3 a-g. e

Type locality: Lagoon at the Panguana biological station, Peru.

Holotype: SMG GP 7366, Senkenbergmuseum, Frankfurt am Main, Gennany.

Description: Body fairly stout, slightly gibbous, reminiscent of C. gibba. Lorica soft,
plates not discernible; foot short; toes tapering unifonnly, slightly bent dorsally. Co-
rona not oblique, with lips. Footglands with reservoir; four salivary glands, ganglion
with a sack-like appendage (RCO ?) but no ducts; no eyespot. Trophus type B: ful-
crum with anterior thickening and basal apophysis; rami with five tenninal teeth each;
manubria widen into a sickle-shaped ventrally twisted part that ends in knob. Close to
oxydactyla.
Total length 250-270 !-tm, toes 74-81 !-tm, trophus 48 !lm.
Ecology: In the plankton and littoral vegetation in a lagoon, Peru.
Literature: Not seen since discovery. la,);;. (~;~l:(i) t J!.:i."Q.;,.",-,., tJ..OIJ/ 1
Code: 0102, 0201, 0302,0407,0601,0803,0902,1005,1201,1307,1311, 1403,
1501, 1601, 1802.
c tr ptto.
(J OcJJ' toL, ,fa
Cept~lodella calosa Wulfert, 1956
Fig. 71

C. calosa Wulfert 1956, pA63, FigA.

72
Type locality: Teufelssee, Berlin, Gennany.
Holotype; Not designated Fig. 70. C. boettgeri laterally and ventrally; (a): trophus, (b): ramus laterally, (d): manubrium, (e): incus
laterally, (t): uncus.
Fig. 71. C. calosa.
Description: Body cylindrical, not compressed laterally. Head short; lorica well de- Fig. 72. C. carina dorsally, laterally, and trophus.
(70. Koste 1988; 71: Wulfert 1956; 72: Wulfert 1959).
 56 57
lineated. Foot almost entirely covered by tail; toes slightly bent ventrally, acute. Co-
rona oblique, convex, with lips; ganglion long, one cervical eyespot. Vitellarium with
eight nuclei. Trophus type B (not shown); rami without alulae (unlike in C. delicata)
manubria crutched. Resembles C. delicata, but toes less delicate.
Total length 126 ~m, toes 19 ~lm.
Ecology: In a forestpond (pH 7.2-7.7) in a boggy area (pH 4.5-5.0).
Literature: Kutikova 1970.
Code: 0103, 0202, 0204, 0302, 0403, 0504, 0901, 1003, 1102, 1201, 1302, 1401,
1501, 1601.

Cephalodella carina Wulfert, 1959

Fig. 72

C. carina Wulfert 1959, p.60-61, Fig.11.

Type locality: Rheocrene in the Siebengebirge mountains, Germany.

Holotype: Not designated

Description: Stout, hyaline, delicate, but lorica well delineated. Head short; neck dis-
tinct. Foot well separated, short tail. Toes slender, dorsally decurved, always spread.
Corona oblique, convex with small lips. Gastric and salivary glands small; vitellarium
with six nuclei; one very small frontaleyespot. Trophus type B: manubria crotched
with double lamellae, pleural rod (dissolves in bleach), fulcrum with terminal thicken-
ing. Related to C. gracilis, but sturdier, six instead of four vitellarium nuclei. Also re-
sembling C. graciosa which has longer toes.
Total length 105-115 ~m, toes 25 ~m.
Ecology: Among Fontinalis in. ~ ~u,nning spring, feeds. on diatoms.
Literature: Kutikova 1970. k'rfihUovOJ 1~13 ((ZVD< 14%)
Code: 0102, 0202, 0205, 0302, 0502, 0504, 0701, 0801, 0901, 1001, 1005, 1101,
1201,1301,1307,1402,1501,1601,1605,1607,1701,1905.

(". /) ,_./J ; ., • Cf}Phalod.e{l.a.c.at.ellina catellina (Muller), 1786

Gc ,IUL<£.l Uv. iJL C i)StJf!f~Sf ye StM(}f~ l'ft/ (uo.! Fig.73
114/;.(. /1iJ,;W/S,f?<1 Ol. C·CO.J(l(IL\",o.- (ji"-'fre !
Cercaria catellina Miiller, 1786, p. 130, Figs. 20: 12, 13.
Cephalodella catellina Bory de St Vincent, 1826, p. 43.
See Harring & Myers 1924, p.465-466, and Koste 1978, p. 371 for extensive syn-
onymy: myersi, armata, botezati . f..)c,j·k...cc .SY1'IO"lYtl-de ~( . ..J;eJwrd
Type locality: Copenhagen.
Holotype: Not designated
Description: Body short, stout, bulbous distally; abdomen laterally compressed;
10rica flexible with wide lateral sulci separating indistinct lorica plates; foot short,
ventral, with overhanging tail; toes short, straight, thin, conical; two separate red fron-
tal eyespots. Mastax with ventral salivary glands (may be absent in small specimens);
Y75
Fig. 73. C. catellina catellina (a,b): different indi-
viduals laterally; (c): male; (d): trophus laterally; (e)
'u trophus ventrally; (t): distal end of manubrium; (g):
photomicrograph.
Fig. 74. C. catellina major (a): laterally; (b): toe lat-
erally; (c): trophus dorsally; (d): trophus laterally
without manubrium; (e): uncus; (t): manubrium.
Fig. 75. C. catellina natans laterally, dorsallyand
trophi. (73: Koste & Shiel 1991, photo Koste; 74:
Donner 1972; 75: BerziI,ls 1976).
58
59
trophus type C: right ramus with a tooth; long fulcrum slightly expanded distally;
Cephalodella catellina volvocicola (Zavadovsky), 1916
manubria rod-shaped, decurved, ending in semicircular dorsal expansion. Hooked
Fig. 76
manubria ends may result from hypochlorite digestion (Koste 1978).
Total length 80-160 !lm; toes 9-14 !lm (18-20 !lm in Harring & Myers 1924); Diglena volvocicola Zavadovsky 1916 ?
trophus 27 !lm (45 !lm in Harring & Myers 1924); male 140 ~tm, toes 17 !lm. C volvocicola Kutikova 1970, Fig.123. See also Koste 1978 p. 372.
Ecology: Cosmopolitan in fresh to brackish water, occasionally reported in coastal
(marine) waters. Temp. 12-24°C, pH 4.4-6.8, 69-600 !lS cm-I. Type locality: Unknown.
Literature: Harring & Myers 1924, Kutikova 1970, Koste & Shiel 1991. Holotype: Not designated
Code: 0103,0203,0205,0303,0407,0508,0801,0901,1008, 1103, 1202, 1310
1401,1501,1607,1702. Description: Slender toes, rami symmetrical, manubria loops not fully closed (see
Cc.maior).
16 Total length 100-140 !lm, toes 19 !lm, male 100-11 0 !lm, resting egg 54-60/38-45
Cephalodella catellina major (Zavadowsky), 19J6 !-Lm .
:: C• I

1/t..~L:t1or
Fig. 74 l.n(- jJ7,S j
:2.. ~
Q Jg
t
0

"HVI \111
)
. illS
Ecology; Parasitic in Volvox colonies, Europe, N.America.
Literature: Kutikova 1970.
Cephalodella cateIlinaf major Zavadovsky 1916 p.? ;:'000{ J'erc,'t S Code: see C c. malor.
Cephalodella armata Rudescu 1960 p,.738, Fig.599.
~ C· wis7.J,I lew.s?1Ji Ed"u(?v1tJ(sOv 1 l: H:;(c1li<,soVl rq34 --7 Spec (Off!../.? rlJeSvt(2'/ 'l"q
f

Type locality: Russia. Cephalodella celerls Myers, 1937

Holotype: Not designated Fig. 77

Description: Body round, not flattened, dorsally flared. No salivary glands. Trophus C celeris Myers 1937, p.13, Fig.14.
type C: rami asymmetrical, with flaring tips, fulcrum strongly bent dorsally, manubria
with large loops that are not fully closed. Unci coarse with a lamella under the teeth. Type locality: Adirondack mountains, NY, D.S.A.
Total length 140-190 !lm, toes 21-26 !lm, trophus 43 !lm. Holotype: AMNH # 700 (lost)
Ecology: Common in saprobic areas including saline waters.
Literature: Donner 1972, Koste 1978. Description: Body elongate, nearly cylindrical, abdomen gradually inceases in depth
Code: 0103, 0203, 0205, 0303, 0402, 0405, 0508, 0802, 0901, 1008, 1103, 1202, posteriorly then falls away rather abruptly. Sulci distinct, flaring distally. Foot small;
1310,1402,1501,1607,1801. small tail; toes very characteristic: the basal portion is slightly expanded, then sud-
denly constricted and again expanded, whence they diminish gradually to acute tips.
Foot glands are stout and pyriform; gastric glands very large and oval. Corona oblique
Cephalodella catellina natans Berzilfs, 1976 and convex, no lips. Ganglion long, saccate; no RCO or eyes. Mastax type A ? (not
Fig. 75 shown), manubria slender, decurved dorsally. The animal - as the name implies -
moves very rapidly.
C cateIlina natans Berzilfs, 1976. Total length 135 !lm, toes 20 !-Lm.
Ecology: Common, in acidic oligotrophic mountain lakes, pH 6.4-7.0.
Type locality: Lakes Allgunnen and Hillen, Sweden. Literature: Kutikova 1970, figure in Koste 1978, 126:9.
Holotype: Not designated Code: 0103,0201,0301,0501,0901, 1003, 1101, 1201, 1303,1304,1308, 1402,
1501, 1605, 1607, 1803, 1808.
Description: Colourless, transparent. Dorsally, on both sides of the head, a bump.
Tota11ength 135-147 !lm, toes 22-24 !lm, trophus 38-40 !-Lm.
Ecology: Planktonic. Cephalodella dara Wu1fert, 1944
Literature: Not seen since discovery. Fig. 78
Code: see Cc.catellina.
C dara Wu1fert 1944, p.385, Fig.5.

Type locality: In manure, Austria.

Holotype: Not designated
 61

Description: Transparent, colourless, body widens posteriorly into a triangular stout

foot, covered only partly by the tail. Very thin elastic lorica. Toes dorsally bent, first
half thicker, second half uniformly thin, ending in blunt tips. Corona oblique, no lips.
Small salivary glands; vitellarium with eight nuclei. No eyespots. Trophus type B:
rami without alulae or teeth; manubria asymmetric, terminally thickened, crotched,
and bent. V-shaped pleural apparatus. Resembles C. indIa, limosa, and tenuior.
Total length 140-180 Ilm, toes 25-42 Ilm, trophus 22-29 !lill, manubria 14 and 17
Ilm.
Ecology: In manure and humic soil; similar to C. rotunda bryophila, but not a syno-
nym.
Literature: Wulfert 1944, Kutikova 1970, Donner 1970.
Code: 0103, 0201, 0302, 0401, 0509, 0702, 0801, 0901, 1006, 1101, 1201, 1307,
1402,1507,1605,1607,1702,1904.
78
77 Cephalodella collactea Myers, 1924
Fig. 79

C. collactea Myers 1924, Harring & Myers 1924, p.501, Fig.34:5

Type locality: Loon Lake, WI, D.S.A.

Holotype: Not designated

Description: Body moderately elongate, nearly cylindrical; head very short, strongly
oblique anteriorly; neck well marked; lorica fairly rigid, plates well marked, with lat-

rr
eral clefts narrow anteriorly and widening gradually towards posterior end; toes long,
straight, slender, very slightly tapering, ending in a bristle-like tip. Gastric glands fre-
quently tinted red; ganglion moderately long, saccate; eyespot and RCO absent; co-
rona oblique, strongly convex without projecting lips. Trophus type B: fucrum
straight, slightly expanded posteriorly; manubria slender, crotched (no figure). Near-
79 80 est relatives are C. intuta (v/hiGh ba51 a cerebral e¥.e) and C. asta, from which it is read-
ily distinguished by the form of the toes. V/ I /J.. (I
Total length 130 Ilm, toes 32 Ilm. ItA Yu'tt:J. ~J t.t..o eye ..'
Ecology: Found in one lake only; acidophil?
F~g. 76. C. catel~ina volvocicola (a): laterally and trophus; (b): in a Volvox colony. Literature: Not seen since discovery.
Fig. 77. C. celens. Code: 0102, 0201, 0302, 0504, 0901, 1002, 1101, 1201, 1301, 1306, 1402, 1502,
F:"g. 78. C. clara young animal dorsally, adult laterally, trophi. 1605, 1607, 1807.
FIg. 79. C. collactea.
Fig. 80. C. compacta.
Fig. 81. C. compressa.
Fig. 82. C. crassipes laterally and toe. Cephalodella compacta Wiszniewski, 1934
Fig. 83. C. cyclops dorsall;y, laterally, trophi and eye frontally. (76: Kutikova 1970' 77: Myers 1937' 78' Fig. 80
Wulfert 1944; 79, 81: Hamng&Myers 1924; 80: Kutikova 1970; 82: Kutikova 1970; 83: Wulfert 1937):
C. compacta Wiszniewski 1934, p. 351, Figs. 58:11-12. (not seen).

Type locality: Psammon, Poland.

Holotype: Not designated

Description: Short, stout, bulbous, laterally slightly compressed; lorica indistinct.

 62
Head narrower than body; foot small, conical; tail just indicated; toes short, straight.
Corona not convex, no lips. Double frontal eyespots close together. Trophus type A,
simple, manubria straight.
Total length 100-110 !lm, toes 15-18 !lm, trophus 20 !lm.
Ecology: Psammon. Koste claims presence in North America (without a citation).
Literature: Not seen since discovery ?
Code: 0103,0203,0205,0301,0503,0801,0901, 1001,1006,1008, 1101, 1201,
1301,1306, 1401, 1501, 1606, 1907.
Cephalodella compressa Myers, 1924
Fig. 81
C. compressa Myers 1924, Harring & Myers 1924, p. 487, Fig. 30: 1.
Type locality: Not designated, "common in Wisconsin and New Jersey", D.S.A.
Holotype: Not designated
Description: Body moderately elongate, slightly gibbous dorsally, strongly com-
pressed laterally; head rather small, distinctly deflexed; lorica fairly rigid, plates dis-
tinct; foot broad at base, fairly stout; toes long, slender and cylindric, taper gradually
to very acute tips; the dorsal edge is straight throughout and the ventral edge bends
upwards to meet it. Corona oblique without projecting lips. Trophus type A: fulcrum
long, stout, slightly expanded at posterior end; manubria very slender, rodlike,
strongly recurved posteriorly (not shown). Gastric glands small, irregularly ovate.
Ganglion long and saccate; eyespot at posterior end, well towards ventral side. RCO is
absent.
Total length 140-145 !lm, toes 45-48 !lm.
Ecology: Acidophil, among Sphagnum.
Literature: Not seen since discovery.
Code: 0101,0202,0204,0301,0901, 1002,1007,1101,1201,1301,1306,1501,160-
5, 1607, 1803, 1903.
Cephalodella crassipes (Lord), 1903
Fig. 82
Diaschiza crassipes Lord 1903, p.78, Fig.3:3
C. donneri Hauer 1959? C. harringi Smimov 1927, C.jakubskii Wiszniewski ?
Type location: On gills of decapods (Astacus, Potamobius).
Holotype: Not designated
~escription: Orig~nally described from a single specimen. Body moderately slender,
glb~ous d~rsally; llltegument very flexible, with no trace of a lorica; foot large and
comcal; .taIl reaches somewhat beyond mid-length of foot; toes are blade-shaped,
short, shghtly decurved., Corona very slightly oblique, without projecting lips.
Trophus type B:manubna somewhat stouter than usual, bent at obtuse angle near
63
midlength. Oesophagus very long, convoluted. Ganglion large, saccate; no (or only
very faint) eyespot (the species is listed under the eyeless category); no retrocerebral
organ. While Koste (1978, p. 369) synonymizes C. xenica with crwl'sipe!·;. we agree
with Hauer (1959 p. 101) that xenica is a good species, distinguished by a cervical
eyespot. C. eupoda was also synonymized by Koste, but we retain it as a valid species.
Total length 130-250 ~!m, toes 22-30 !lm.
Ecology: While discovered as an ectoparasite on decapod gills, (also as donneri
Hauer) other cognate species (eupoda) live free.
Literature: Harring & Myers 1924, Hauer 1959, Kutikova 1970.
Code: 0103, 0201, 0302, 0404, 0501, 0802, 0902, 1006, 1101, 1201, 1302, 1402,
1502, 1607, 1803, 1902.
Cephalodella cyclops Wulfert, 1937
Fig. 83
c. cyclops Wulfert 1937 p. 604-605, Fig. 14
Type locality: Pond near the River Saale, Germany.
Holotype: Not designated
Description: Body stout, round; head wide, neck indistinct; lorica well defined,
sulcus wide. Foot tapered; tail very short. Toes dagger-like, curved ventrally. Corona
slightly oblique, very convex, lips present. Four salivary glands; RCO present; double
frontal eyes with lens; however, Koste (1976, Fig.6: 1a) shows an eyeless figure.
Trophus type B: rami with thornlike alulae; manubria long, crotched; fulcrum spatu-
late, two thin decurved pleural rods. Related to reimanni, but rami are asymmetrical
with small thornlike alulae, foot characteristically bulging ventrally.
Total length 115-125 !lm, toes 19-23 /lm.
Ecology: Rare, but was common in type locality; also in submerged moss.
Literature: Kutikova 1970, Koste 1976.
Code: 0103,0203,0205,0208,0302,0404,0504,0701,0801, 0901, 1001, 1101,
1201,1302,1401,1501,1601,1605,1607,1804,1802.
Cephalodella delicata Wu1fert, 1937
Fig. 84
C. delicata Wu1fert 1937, p. 603, Fig. 13.
Type locality: Alkaline waters near Halle and Merseburg, Gennany.
Holotype: Not designated
Description: Body moderately elongate, nearly cylindric and slightly gibbous
dorsally. Head relatively long and strongly oblique anteriorly, in dorsal view wider
than body. Neck marked by a shallow constriction; integument very flexible, plates
ill-defined; the lateral clefts are somewhat obscure apparently narrow and parallel-si-
ded throughout their length. Foot short and broadly conical; the tail is small, rounded,
 65
64

with a thickening near mid-length. Toes long, slender, slightly decurved, taper gradu-
ally to extremely acute, needle-like tips. Corona strongly oblique, decidedly convex
and with projecting lips. Trophus type A: manubria very slender, rami corners are
elongated like an alula. Ganglion elongate and saccate; no RCO; eyespot cervical.
Total length 100-125 ~m, toes 15-23 ~tm, trophus 23-25 ~tm.
Ecology: Benthic, widespread in Europe, incl. salt and acid water.
Literature: Wulfert 1943, Kutikova 1970.
Code: 0103, 0202, 0204, 0301, 0404, 0501, 0601, 0901, 1002, 1005, 1101, 1301,
1308, 1401, 1501, 1601, 1605, 1607, 1803.

Cephalodella delicata "form" eudelicata (Wulfert), 1961 84

Fig. 85

C. eudelicata Wulfert 1961, p. 82, Fig. 5.

Type locality: Two ponds in Diibener Moor, Germany.

Holotype: Not designated

Description: Differs from C.d.delicata in not being as slender; tail covers half the
foot, and the ends of manubria are not crotched but are S-shaped, the end curved in-
ward and dorsally. Has the same triangular alulae as C.d.d. and C. bryophila. Perhaps
an ecotype or contraction artefact of C. delicata.
Total length 94-1 04 ~m, toes 18-21 ~m, trophus 20 ~m.
Ecology: In a pond with many macrophytes, pH 6.0.
Literature: Kutikova 1970.

Cephalodela dentata Wulfert, 1937

Fig. 86
:I
C. dentata Wulfert 1937, p. 614-615, Fig. 24

Type locality: In a ditch, Germany.

Holotype : Not designated

Description: Slender, relatively straight body, resembles C. rotunda bryophila in

~\
habitus, dorsal sulcus forms a keel. Head sturdy, only slightly oblique and convex. 11
Lateral sulci flare· distally; foot stout, covered by tail; toes slender, straight, ending in 89 91
blunt tubular tips. Trophus type B, asymmetrical:~gth*and thorn-like alulae on rami,
1_manubria asymmetrical. No eyespot. Related to C. rotunda bryophila, pachydactyla, Fig. 84. C. delicata dorsally, laterally, toe laterally, and trophi.
and intuta, but the toes and the long alulae distinguish this taxon. Fig. 85. C. delicata "form" eudelicata.
Hr! Total length 150-170 ~m, toes 36-38 ~m, trophus 22 ~m. Fig. 86. C. dentata lateraly, dorsally, toes and trophi
Fig. 87. C. dixon-nuttalli laterally; toes dorsally.
) Ecology: In a ditch. Fig. 88. C. dora.
Literature: Kutikova 1970. Fig. 89. C. dorseyi.
Code: 0102, 0201, 0302, 0404, 0407, 0504, 0509, 0801, 0901, 1002, 1102, 1201, Fig. 90. C. doryphora.
1301,1306, 1402, 1507, 1607. Fig. 91. C. dorystoma. .
Fig. 92. C. elegans. (84, 86: Wulfert 1937; 85,88: Wulfert 1961; 87,89,92: Barnng & Myers 1924; 90,91:
Myers 1934).
66 67

Cephalodella dixon-nuttalli Myers, 1924 Type locality: Atlantic City, NJ, D.S.A.
Fig. 87 Holotype: AMNH ? lost?

C. dixon-nuttalli Myers 1924, Barring & Myers 1924, p. 498-499, Figs. 33:4, 5.
Type locality: Lake Cathan, WI, D.S.A.
Holotype: AMNH? not listed.
Description: Body elongate, fairly slender and laterally compressed; head rather
small. Body cylindrical, lorica very thin and flexible and the plates are difficult to
trace. Foot very stout; tail small and somewhat beyond mid-length. Toes short, chang-
ing abruptly to conical, acute tips; viewed dorsally, the toes are strongly appressed,
looking like a single toe. Corona oblique, slightly convex and without projecting lips.
Mastax rather small, trophus type A: fulcrum very slightly expanded posteriorly;
manubria very slender and not crutched. Gastric glands very small. Eyespot and RCO
absent.
Total length 160 ~m, toes 30 ~m.
Ecology: Acidophil
Literature: Not seen since discovery.
Code: 0103, 0201, 0301, 0801, 1002, 1101, 1201, 1306, 1401, 1502, 1606, 1607,
1903.
Description: Body relatively short, stout and slightly gibbous dorsally; head large.
Lorica finn, plates distinct; the lateral clefts are fairly wide. Foot short and stout; tail
fairly prominent but short. Toes very long and wide apart at the base, fairly stout and
end in acute, conical tips. Corona strongly oblique, distinctly convex without project-
ing lips. Trophus large, type A: fulcrum moderately long and slightly expanded at the
posterior end; manubria slender, rod-shaped, slightly clubbed and decurved poste-
riorly, but not crutched. Gastric glands small and ovate; ganglion long and saccate;
eyespot very large and saucer-shaped, at the posterior end of the ganglion. ReO ab-
sent.
Total length 145-150 ~m, toes 50-54 ~m.
Ecology: Acidophil, rare.
Literature: Not seen since discovery.
Code: 0101,0204, 0301,0510,0901, 1001, 1101, 1201, 1306, 1402, 1502, 1606,
1607,1803,1903.
Cephalodella doryphora Myers, 1934
Fig. 90

C. doryphora Myers 1934, p. 2, Fig. 1.

Cephalodella dora Wulfert, 1961
Fig. 88 Type locality: Mount Desert Island, ME and NJ, D.S.A.
Holotype: AMNH ? lost?
C. dora Wulfert 1961, p.81, Fig. 4.
Description: Body very short and stout; head large, slightly oblique. Neck constric-
Type locality: Acidic pond, Germany. tion sharp; lorica firm, distinct. Foot very short, minute tail; toes short, ventrally
Holotype: Not designated decurved, blade-shaped. Corona strongly convex with beak. Ganglion long, cervical
eyespot somewhat ventrally shifted. Mastax huge, trophus type A: fulcrum long,
Description: Body cylindrical, dorsally straight, colourless; head large, neck indis- stout, expanded; manubria semicircular, tips swollen. Epipharynx long, the species is
tinct. Lorica distinct, sulci well visible. Foot fairly large, half-covered by tail. Toes raptorial. Resembles C. physalis, but is smaller, has the prominent beak and different
stout, uniformly tapered. Vitellarium with eight nuclei; three salivary glands. Eyes trophus. Dorsally, there is considerable similarity with C. auriculata; trophi are also
absent. Trophus type A: unci straight and long, manubria gently curved, of unifonn very similar.
thickness. Similar to C. dorystoma (four nuclei in vitellarium), C. calosa (big tail, Total length 105 ~m, toes 15 ~m.
crutched manubria). Ecology: Acidophil, predator.
Total length 195 ~m, toes 30 ~m, trophus 24 ~m. Literature: Berzins 1976.
Ecology: In detritus, October, pH 6.0. Code: 0103, 0204, 0301, 0510, 0701, 0802, 0901, 1001, 1101, 1201, 1302, 1308,
Literature: Kutikova 1970. 1402, 1501, 1602, 1903.
Code: 0103, 0201, 0301, 0510, 0801, 0901, 1002, 1101, 1201, 1302, 1402, 1501,
1605, 1702, 1802, 1903.
Cephalodella dorystoma Myers, 1934
Fig. 91
Cephalodella dorseyi Myers, 1924
Fig. 89 C. dorystoma Myers 1934, p. 5-6, Fig. 5.

C. dorseyi Myers 1924, Harring & Myers 1924, p. 487-488, Fig. 30:7. Type locality: Mount Desert Island, ME, D.S.A.
68 69
Holotype: AMNH 362 (co-type)

Description: Body elongate, a tapering cylinder; head small, obliquely truncated.

].~'."~'."
Lorica finn, plates well defined; toes short, acute, decurved ventrally; footglands very
large, pyriform. Corona convex, no lips; ganglion long, carries a small refringent ;.
, 1
vacuole (also called "clear eyespot"; "crystal" according to Wulfert (1938a) near the ~~:" .•\.I.\
......•.,.,•.•. .•.•.••'.'.'.•.I

posterior end; keyed under both eyed and eyeless taxa. Gastric glands huge, nearly
equal to the mastax in size. Trophus type A: fulcrum slightly expanded posteriorly, cl~
manubria straight, slender rods, somewhat expanded at the end (not shown). Related
to C. glandulosa. ff?·:
Total legth 170 [1m, toes 20 [1m.
Ecology: Among detritus, pH 6.4; single occurrence. See also C. eunoma among spe-
cies inquirendae at end of genus.
Literature: Myers 1934, Wulfert 1938.
)
Code: 0103,0204, 0206, 0301, 0801, 0901, 1002, 1003, 1101, 1201, 1301, 1308,
1402,1501,1605, 1607, 1808, 1903. 95 94

Cephalodella edax Hollowday, 1993

Fig. 93

C. edax Hollowday 1993, p. 445-448, Fig. 1-3.

Type locality: Blelham Tarn, England.

Holotype: British Museum Nat. Hist. 1991.4093

Description: Body stout, bulbous, neck marked. Head large, not deflexed; lorica thin,
plates not visible on iconotype. Foot not well differentiated, small; toes small, base
wide narrowing suddenly into fine tip. Corona not oblique, no lips. Trophus symmet-
ric, type C; fulcrum straight rod without widening; manubria slightly bent with small
loop at end, no basal plates; unci short, blunt; rami with large alulae, no teeth. No eye-
spot, no information on glands or RCO.
Total length 136 [1m, toes 15-16 [1m, trophus 16 [1m.
Ecology: Parasitic in Uroglena volvox.
93
Literature: Not seen since discovery.
Code: 0103, 0201, 0303, 0401, 0404, 0406, 0508, 0602, 0801, 0901, 1001, 1008, Fig. 93. C. edax live specimens, trophus dorsally and end of manubrium.
1101,1201,0301, 1401, 1502, 1701,1902. Fig. 94. C. elongata.
Fig. 95. C. euderbyi Ca): laterally; Cb): toes; (c) trophi dorsally and laterally; (d): rami open, (e): epi-
pharynx; (t): male.
Fig. 96. C. euknema.
Cephalodella elegans Myers, 1924 Fig. 97. C. eupoda. (93: Hollowday 1993, photo H. Canter-Lund; 94,96,97: Harring & Myers 1924; 95:
Fig. 92 Wulfert 1940).

C. elegans Myers 1924, Harring & Myers 1924, p.489-490, Fig. 31: 8,
Type locality: New Jersey sphagnum bog, D.S.A.
Holotype: AMNH 418 (co-type)
Description: Body elongate, very slender, somewhat curved and slightly compressed
dorsoventrally. Head relatively large and very slightly deflexed. Lorica firm and the
plates distinct; lateral clefts increase gradually and considerably in width towards the
posterior end. Foot small, terminates in a short, cylindric section; tail very minute.
Toes very long and slightly decurved, fairly stout at the base and tapering rapidly to
slender, nearly cylindric, acute rods. Corona slightly oblique and moderately convex
70
without projecting lips. Trophus small, type A: fulcrum slightly e~pan.ded at the poste-
rior end; manubria distinctly recurved and increase some~hat In wIdth to:vards the
ends. Gastric glands small and oval. Ganglion long and pynfonn; eyespot faIrly large,
at the posterior end of the ganglion; no RCO.
Total length 135-140 [.till, toes 48-50 [.till.
Ecology: In a sphagnum bog, not common.
Literature: Wulfert 1938.
Code: 0101,0202,0204,0301,0501,0901, 1002, 1003, 1101, 1201,1301, 1303,
1308,1402,1501,1607,1803,1903.
Cephalodella elongata Myers, 1924
Fig. 94
C. elongata Myers 1924, Harring & Myers 1924 p. 471-472, Fig. 26:2
Type locality: Atlantic City, NJ, D.S.A.
Holotype: Not designated
Description: Body elongate, cylindric, and extremely slender. Head unusually long
and convex anteriorly, neck indistinct. Abdomen cylindric throughout its length; the
lorica is very thin, flexible, and the plates ill-defined; the lateral clefts are very ob-
scure, but apparently very narrow and parallel-sided. Foot short and conical; the small
tail is near the posterior end. Toes short, slightly decurved ventrally, taper gradually.
Footglands very small and pyriform. Corona only slightly oblique, but strongly con-
vex, without projecting lips. Mastax is moderately large, trophus type B: fulcrum slen-
der and slightly recurved at the posterior end; manubria crutched. Gastric glands elon-
gate pyriform, the obtuse end forward. Ganglion very long, saccate; RCO absent, eye-
spot frontal and double, the two pigment spheres fairly wide apart.
Total length 105-115 [.tm, toes 21-25 [.tm.
Ecology: Acidophil, not common.
Literature: Kutikova 1970.
Code: 0102, 0203, 0205, 0302, 0504, 0901, 1002, 1003, 1101, 1201, 1301, 1308,
1402, 1501, 1607, 1808, 1903.
Cephalodella euderbyi Wulfert, 1940
Fig. 95
C. euderbyi Wulfert 1940, p.564, Figs. 4 a-k.
Type locality: Birkhorst Moor (eastern Germany).
Holotype: Not designated
Description: Body stout, lightly arched dorsally; head approx. one-third body length;
corona with slightly protruding lips and epipharynx; posterior dorsal lorica com-
pressed, resembles keel; foot short, almost covered by pointed tail. Toes short,
straight, occasionally slightly recurved; mastax large, with distinct salivary glands;
71
vitellarium with eight nuclei. Trophus of type A: fulcrum spatulate distally; manubria
thin, rod-like, with single crook (not crutch) distally~ rami single. with very small
apical teeth; two small ruby-red cerebral eyespots.
Total length 150-168 [.tm; trophus 32 !lm; toes 28 !lm~ male 1] 0 Ilm; resting egg 53
[.tm.
Ecology: Described from a shallow moor in Germany. feeding on algae. Single
unverified record from Boar's Lagoon, Vie., Australia.
Literature: BerziJ)s 1982, Koste & Shiel 1991.
Code: 0103, 0203, 0204, 0301, 0407, 0501, 0802,0901, 1001, 1101, 1201, 1306,
1402,1502,1601,1607,1702,1903.
Cephalodella euknema Myers, 1934
Fig. 96
C. euknema Myers 1934, p. 5, Fig. 4.
Type locality: Mount Desert Island, ME, D.S.A.
Holotype: PANS 130 (paratype)
Description: Body elongate, slender, cylindrical; head fairly large, obliquely trun-
cate, neck well marked. Lorica flexible but well defined. Foot short, tail minute. Toes
very long and slender, slightly swollen at mid-length, whence they taper to abrupt,
upturned, very slender tips. Trophus (type A ?) large, fulcrum stout and spatulate,
manubria slender and recurved. Ganglion long, single cervical eyespot. Very close to
nana, but toe-tips characteristic.
Totallength 225 [.tm, toes 75 [.tm.
Ecology: Among decaying meadow grass, pH 6.4.
Literature: Wulfert 1938.
Code: 0101, 0202, 0204, 0301, 0502, 0902, 1002, 1003, 1101, 1201, 1301, 1304,
1403, 1502, 1503, 1605, 1903.
Cephalodella eupoda Myers, 1924
Fig. 97
C. eupoda Myers 1924, HalTing & Myers 1924, p. 512, Fig. 36.7.
Type locality: Washington DC, U.S.A.
Holotype: Not designated
Description: Body moderately slender, gibbous dorsally. Head large, neck well
marked. Integument very flexible, no trace of lorica. Foot large and conical; the
minute tail reaches somewhat beyond mid-length of foot. Toes blade-shaped, short
and slightly decurved with conical, acute tips. Corona very slightly oblique, without
projecting lips. Mastax type A: manubria somewhat stouter than usual and b.ent at an
obtuse angle near midlength. Oesophagus very long and convoluted. Ganghon large
and saccate; no eyespot and no trace of RCO.
72 73

Total length l05 ~tm, toes 22 ~tm.

C. eupoda has been synonymized with crassipes by Koste 1978; it is, however,
eyeless, while crassipes has a faintly red eyespot. It differs from the closely related
xenica by having a bent but not crutched manubrium. Therefore, we consider it a valid
species until further study.
Ecology: Unknown.
Literature: Not seen since discovery.
Code: 0102, 0201, 0301, 0510, 0801, 0901, 1002, 1006, 1302, 1310, 1402, 1507,
1803.

Cephalodella eurynota Myers, 1934

Fig. 98

C. eurynota Myers 1934, p. 8, Fig. 7.

Type locaity: Mount Desert Island, ME and NJ, U.S.A. 98 99

Holotype: Not designated

Description: Body cylindric, fusiform in second half, gibbous; head small, slightly
deflexed and truncated. Lorica rigid, plates well defined. Toes very long, slender, uni-
formly tapered and curved ventrally. Footglands large. Corona strongly convex with
lips, ganglion long, no eyespots. Trophus large, (type A ? not shown) fulcrum long c
and spatulate, manubria sightly decurved.
Total length 105, ~m, toes 30 ~tm.
Ecology: Among Sphagnum, not rare at type locality.
Literature: Not seen since discovery.
Code: 0102, 0201, 0301, 0501, 0901, 1001, 1009, 1101, 1201, 1301, 1308, 1402,
1501, 1601, 1903.

100
Cephalodella eva (Gosse), 1887
Fig. 99
101
Furcularia eva Gosse, l887b, p. 864, Fig. 14:9. Fig. 98. C. eurynota.
Cephalodella eva: Harring & Myers, 1924, p. 507. Fig. 34:8. Fig. 99. C. eva laterally with trophi.
Fig. 100. C. evabroedi (a,b): laterally; (c): toes.
Fig. 101. C. exigua scale bars 50 and 10 /lm.
Type locality: England. Fig. 102. C.jluviatilis. (d): trophus; (e) fulcrum laterally. (98: Myers 1934; 99,101: Koste & Shie11991,
Holotype: Not designated photo Koste); 100: de Smet 1988; 102: Kutikova 1970).

Description: Body slender, laterally compressed, dorsally gibbous; head short, broad,
neck c1~ar1y marke~; plates distinct; corona oblique without proj ecting lips; foot
Ecology: Cosmopolitan, acidophil, also in psammon. Grazes on diatoms.
large, WIth large pynform foot-glands, tail can cover the foot (see Donner 1964); toes
Literature: Harring & Myers 1924, Wulfert 1940, Kutikova 1970, Koste & Shiel
long~ approx. half t?tal length, very slender and generally curved ventrally. Trophus
modIfied type B WIth long, strongly fanned manubria (not crutched), rami with nu- 1991.
Code: 0102,0201,0302,0407,0506,0801,0902, 1002,1007, 1101, 1201,
merous teeth; no eyespot or RCO. The European form seems to be variable the North
American morphologically stable. ' 1301,1309,1402,1503,1506,1601,1605,1803,1903.
Total length 190-285 ~m; trophus 23-30 ~m, toes 50-85 ~m.
74
Cephalodella evabroedi de Smet, 1988
Fig. 100
C. evabroedi de Smet 1988, p. 14, Figs. 6: 1a-e.
Type locality; Krillvatnet, Bj0m0ya Island.
Holotype: Lab. Allg. Biologie, Rijksuniversiteit Antwerpen, Belgium.
Description: Body stout, moderately gibbous, head large. Lorica distinct, sulci nar-
row. Foot and tail small, toes short, stout, stippled, base swollen. Corona convex with
lips. No eyespQts, vitellarium with eight nuclei. Trophus type B, very large, almost
symmetrical: fulcrum spatulate, broad in sideview, rami with inner teeth. Manubria
strongly decurved inward with a crutched inner projection, so that the end of the
manubria also appears to be crutched. This could be considered as a transition to type
C, and shows similarity to the manubria of C. forceps. The manubria also carry alulae.
Pleural rods present.
Total length 201-221 /lm, toes 31-41 /lm, mastax 40-42 /lm.
Ecology: In a medium-hard water subarctic pond, pH 8.3.
Literature: Not seen since discovery. ./7.(.
Code: 0103, 0201, 0302, 0407, 0504, 0507, 0509, 0701, 0802, 0902, 1001, 1008,
1101,1201,1302,1306,1402,1502,1601,1702.
Cephalodella exigua (Gosse), 1886
Fig. 101
Diaschiza exigua Gosse, Hudson & Gosse 18862:78, Fig. 22: 13.
) 1967, Donner 1978.
C. exigua (Gosse), Harring & Myers 1924,481, Fig. 28:2.
Type locality: England, aquarium.
Holotype: Not designated
Description: Body short, stout, truncated at posterior end; lateral sulci between lorica
plates distinct; head large, neck clearly demarcated; foot small, tubular; toes short,
tapering gradually to acute tips; salivary glands may be absent; trophus modified type
B: fulcrum slightly expanded distally, rami with teeth, manubria strongly curved
distally. Two red cerebral eyespots; caudal antenna very distinct; male known; resting
egg with smooth dark brown shell.
Total length 90-125 /lm, toes 20-26 ~m, trophus 30 /lm; male 74 /lm, toes 19 flm.
Ecology: Cosmopolitan, pH 6.2-7.4
Literature: Harring & Myers 1924, Donner 1970, Kutikova 1970 Koste & Shie1
1991. ' with large crutches and alulae; unci thick and coarse, rami with alulae (double a1ulae
Code: 0102, 0203, 0204, 0302, 0407, 0501, 0802, 0901, 1001, 1101, 1201, 1301,
1402, 1501, 1605.
75
Cephalodella .lluviatilis Zawadowski. 1926
Fig. 102
Diglenafluviatilis Zawadowski 1926, p. 266, Figs. 8,9. (not seen)
C. elmenteita de Beauchamp 1932.
Type locality: Zwenigorodskoi, Russia.
Holotype: Not designated
Description: (Pr~pare? after. figure, no text seen). Body stout, short, cylindrical, head
equally stout; sWImmmg ammal can be sack-shaped. Lorica indistinct, lateral sulci
appear wi.de. Foot small, en~irely ventral, no tail; toes small, dagger-like, straight.
Corona slIghtly convex, no hps. Described as eyeless (Wulfert 1938, Donner 1978)
but Rudescu (1960) claims two frontal eyespots which, however, are not shown in his
figure. Trophus type B (or C, according to Wulfert 1938): rami asymmetrical, no teeth
but sharp alulae; manubria also asymmetrical, the left one is bigger, with open rings or
forks. Closely related to C. catellina. Donner (1978) describes a '''variety'' Cl minor
which differs in size (only 79-125 flm, (contracted), toes 8-18 lAm, trophus' 30 ~tm):
i.e., very close to Rudescu's measurements. It has no alula on the right ramus, 12 ovar-
ian nuclei.
Total length 210-230 /lrn. Rudescu: length 180-210 /lm, toes 18-20 ~m, trophus 15-
i:~~gy: In freshwater (o~~~e~)"b~;~~~? o~'~~~~~C:;;~a~~ id: Be~uChamp
J
1932, PourrlOt et a!. 1967)/Jejwll~ (1~{1~)
Literature: Zawadowski 1926, de Beauchamp 1932, Rudescu 1960, Pourriot et a!.
Code: 0103, 0201, 0302, 0303, 0402, 0405, 0505, 0801,0902, 1001, 1006, 1008,
1101,1202,1302,1207,1401,1501,1507,1703,1906.
Cephalodella forceps Donner, 1950
Fig. 103
C.forceps Donner 1950, p. 313, Fig. 10.
Type locality: Moravia.
Holotype: Not designated
Description: Body stout, round, back similar to C. gibba, head short, neck indistinct,
lorica thin. Foot covered by tail, toes dagger-shaped, short, straight, but are carried
pointed dorsally. The toes are "empty" (vacuolated 7) in the middle. Mastax with sali-
vary glands, gastric glands large. Trophus sturdy, type B: fulcrum spatulate, manubria
according to Donner 1978). Double frontal eyespots. Swims fast, restless.
Total length 168-191 ~m, toes 26-30 ~m, trophus 32 flm.
Ecology: Not rare among plants, Europe.
Literature: Donner 1950, 1978; Koste 1970, Kutikova 1970.
Code: 0103, 0203, 0205, 0302, 0404, 0504, 0802,0901, 1001, 1006, 1102, 1201,
1302, 1306, 1402, 1501, 1605, 1808.
76 77

Description: Body elongate, slender, slightly compressed laterally~ neck well-

marked; lorica plates distinct; sulci narrow, parallel-sided. Foot and tail small, toes
widely spaced at base, short, stout, taper to acute tips~ foot glands large, pyrifonn;
gastric glands large, red pigmented in older individuals. Trophus type B: manubria
asymmetrical, cro?~ed: with a ?ump in the middle, fanned in side view~ rami with
numerous teeth~e;otabse~t;:!In "variety" macrura \Viszniewsky 1936 the tail and
toes are longer, an eraiTIiteeth are absent (see Fig. 105). Included in both medium-
and short-toe key. Close to incila which, however, has a crotched manubrium and a U-
shaped pleural apparatus.
Total length 122-260 [lm; toes 36-58 [lm; trophus 16-26 ~lm.
Ecology: Cqsmopolitan. "var." macrura was found in psammon.
Literature: \Harring & Myers 1924, Donner 1964, Kutikova 1970, Koste & Shie1
1991. '\,S ivd~t eye?
103 Code: 0103, (O~ 0302, 0407, 0506, 0509, 0801, 0902, 1003, 1101, 1201, 1302,
1402, 1501, 18lrr.

Cephalodella forjicula (Ehrenberg), 1832

Fig. 106

Distemmaforficula Ehrenberg 1832, p.l39.

Cforficula, Harring & Myers 1924, p. 476-478, Figs. 34:1-3.

Type locality: Berlin.

Holotype: Not designated
(\

c d 9 Description: Elongate, spindle-shaped body, slight constriction at neck; head large.

Integument t1exible, without lorica plates; abdomen tapers to ill-defined foot; toes
short, stout, recurved dorsally and outwards, and have distinctive transverse spicule
105 row (2-4 spicules) on the dorsal median sUlface which terminates in a larger spine;
104 mastax with salivary glands; trophus type D: rami serrate at the apex, manubria di-
lated distally, but not crooked, with distinctive oval basal plate; single frontal eyespot;
Fig. 103. C. forceps laterally, trophi, toes. no RCO. Builds detritus tubes.
Fig. 104. C.forficata, two different animals; (b): trophi; (c): manubria; (d): fulcrum laterally; (e): uncus; Total length 160-425 [lm, toes 30-95 [lm, trophus 45-55 !-tm (see Koste 1978, p.359
(t): ramus; (g) fulcrum ventrally. for details).
Fig. 105. C.forjicata ''var.'' macntra. (103: Donner 1978 and Koste 1970; 104: Harring & Myers 1924,
Donner 1954 & 1964; 105: Kutikova 1970).
Ecology: Cosmopolitan, prefers pH 4.8-6.8, t: 12-25°C.
Literature: Harring & Myers 1924, Kutikova 1970, Koste & Shiel1991.
Code: 0102,0202,0205,0304,0511,0802,0902,1003,1101, 1201, 1302,1312,
Cephalodella forficata (Ehrenberg), 1832 1313, 1402, 1501, 1803.
Fig. 104/1 05

Notommata forficata Ehrenberg 1832, p. 134. Cephalodellafriebei Koste, Robertson, and Hardy, 1984
Fig. 107
C forficata, Harring & Myers 1924, p. 499-500, Fig. 33:7

Type locality: Berlin. C friebei Koste, Robertson and Hardy 1984, p.564, Figs. 4a-g.
Holotype: Not designated
Type locality: Lago Camaleo, Amazonia, Brazil.
Holotype: Inst. Nacional Pesq. da Amazonia, Manaus, Brazil.
 78 79

Description: Described from contracted specimens. Body short and stout, trapezoidal
in cross section, lorica flexible, sulci wide. Ventral plate projects as a tail over the
small short foot, positioned ventrally. Toes short, blade-shaped. Salivary glands
present, gastric glands very large, large vitellarium with many (32 ?) nuclei. Eyes ab-
sent. Trophus type C, large, asymmetric: fulcrum slightly expanded terminally,
manubria of different size, crutched; rami with comb-like denticulate lamellae.
Total length (contracted) 110-135 lLll1, toes 19-20 ~lm, trophus 33 ~m.
Ecology: Common in a varzea lake on an island in the Amazon river, subjected to
huge (9-11 m) fluctuations of water level.
Literature: Not seen since discovery.
Code: Not applicable to contracted specimens. Trophus and multinucleate vitellarium
are diagnostic. Not included in identification key.

106 Cephalodella gibba (Ehrenberg), 1832

Fig. 108

m Furcularia gibba Ehrenberg 1832, p. 130, Fig.4: 16.

107 Cephalodella gibba, Harring & Myers 1924, p. 472-473, Figs. 30:4-6.

Type locality: Berlin.

Holotype: Not designated

Description: Body slightly elongated, compressed laterally; gibbous rump; lorica

firm with distinct plates; sulci widen slightly posteriorly; toes long, straight or
recurved, slender. Mastax very large; trophus type B: rami with denticulate lamellar
combs on inner ventral margins; manubria strongly crutched, shaft with lamella on
both sides. No retrocerebral organ; single frontal eyespot with lens.
A somewhat variable species with several ecotypes:
C. gibba anebodica Berzit;ls 1976 has trapezoidal swelling at the tip of toes, ful-
crum with apophysis, one end of the manubrium crutch is sharp. Also listed under C.
anebodica. ? l;yvt. tLA C-~. S.5/r. J rIpJ1Svf;,J:loop (MS.)
C. gibba microdactyla Koch-Althaus 1963 has shortet toes, and fan-lIke structure
on the right side of the rami. See also Koste & Shiel1991 p. 122.
Total length 250-450 ~m; toes 67-150 !lm; trophus 70-90 !lm.
Ecology: Cosmopolitan, the most common Cephalodella, including acid and brackish
108 waters.
109 Literature: Harring & Myers 1924, Kutikova 1970, Koste 1978, Koste & Shiel 1991.
Code: 0102, 0202, 0205, 0208, 0302, 0407, 0504, 0803, 0903, 1003, 1006, 1101,
Fig. 106. C. fO/jleula. Scale bars 50 and 10 f!m. 1201,1301, 1307, 1403, 1501,1607, 1803.
Fig. 107. C. jriebei contracted specimens dorsally, ventrally and laterally; trophus, with manubrium (m)
and end of fulcrum (t) laterally.
Fig. 108. C. gibba laterally; scale bars 50 and 10 f!m.
Fig. 109. C. gibboides laterally; p: pleural rod. (106: Barring & Myers 1924; 107: Koste 1984; 108: Cephalodella gibboides Wulfert, 1950
WuJfert 1937, photo Koste; 109: Wulfert 1950). Fig. 109

C. gibboides Wulfert 1950, p. 451-453, Figs. 3a-d.

Type locality: Sphagnum bogs, Glatzer Schneeberg, Gennany.

 80 81
Holotype: Not designated

Description: Similar to C. gibba, but smaller, the back is not gibbous; toes shorter and
not so thin. Lorica very sturdy, sulci well visible even on the head-Iorica. Corona ob-
lique but not very convex; tail covers the foot and has a longitudinal notch. Gastric
glands very large, filled with shiny fat-globules; vitellarium with eight nuclei. The
neck-antenna is hidden in a deep crevice; ganglion very large; double frontal eye-
spots. Trophus is also different: Fulcrum not spatulate, manubria lack lamellae but
possess a median bump, and there is aY-shaped pleural rod. The species resembles C.
panarista Myers, but the thick lorica and smaller size distinguishes C. gibboides.
Total length 184-205 !-Lm, toes 52-56 ~m, trophus 38 !-Lm.
Ecology: Acidophilic, pH 4.1-5.5.
Literature: Wulfert 1960b, Kutikova 1970.
Code: 0102, 0203, 0205,0302, 0407, 050~ 0702, 080~ 0901, 1002, 1102, 1201,
1307,1402,1501,1507,1605,1702,1808,1903.

Cephalodella gigantea Remane, 1933

Fig. 110

m
C. gigantea Remane 1933, p. 554 (nomen nudum)
·i..':
." ...... 1':--

Type locality: Pond sediment, Kiel, Germany.

Holotype: Not designated Ll

Description: Body very large and uniformly slender, lorica soft, sulci invisible. Foot

f\
triangular, not quite covered by tail. Toes very long, thin, undulating, foot-glands
small. Vitellarium with eight nuclei; gastric glands large; ganglion small; eyespot ab-
sent. Trophus type D: rami with numerous small teeth and one large one. There is an
additional plate as well as a tree-like sub-uncus diagonal to the plane of rami.
Manubria with double alulae, fulcrum with basal apophysis. The species is very close
to, if not identical to, C. tenuiseta (Burn) 1890, from which it differs only in size.
Total length 720-800 !-Lm, toes 240-270 ~m, trophus 78-88 !-LID.
Ecology: Benthic, rare, raptorial. Besides in eutrophic sediment, it was also found in
the oligotrophic Stechlinsee, Germany. Cosmopolitan? 111
Literature: Remane 1933, Wulfert 1937, Koch-Althaus 1963, Koste 1985, Koste
1988a.
Code: 0101,0201,0304,0404,0407,0601,0803, 0903, 1002, 1101, 1201, 1301, Fig. 110. C. gigantea laterally and dorsally. (a):
1309, 1403, 1502, 1702, 1808. trophus ventrally; (b): laterally; (c): uncus; (d): with
open unci dorsally; (e): ventrally; (f,g,h): Wulfert's
figures. bo: border of rami lamellae; fan: fanned tips
of subunci; fm: fulcmm; la: manubrium lamella; ma:
Cephalodella gisleni Berzil)s, 1953 manubrium; ra: rami; rt: rami teeth; su: unci teeth.
Fig. 111 Fig. 111. C. gisIeni.
Fig. 112. C. glandulosa, three different animals, and 112
trophi. (110: Koste 1985, Wulfert 1937; 111
c. gisleni Berzi1)s 1953 pA-6, Figs. 1-4. B-erzinGs 1953; 112: Koch-Althaus 1962).

Type locality: Porongorups, Western Australia.

Holotype: Not designated
82
Description: Described from a single contracted individual. Body squat, rounded,
with small bulge in dorsal posterior part; head very large, almost half body length,
somewhat narrower than body; dorsally a distinct longitudinal sulcus visible; foot
stout, distally extending slightly over base of toes; toes proximally thickened, com-
pressed dorsoventrally, tapering abruptly in distal one-third to sharp tips; in
dorsoventral view a very peculiar semicircular deflection of toes; in thickened part of
toes a relatively large duct; mastax large, with two salivary glands; trophus symmet-
ric, intemlediate between type A (manubria) and type B (rami); fulcrum slender, unu-
sually short for genus, slightly dilated distally; manubria slender, tenninally crooked;
rami wide, of simple construction, without teeth. No eyes or lenses. Koste & Shiel
(1991) regard C. gislbzi as a valid species on the basis of the distinctive trophus, toes,
and other characters described above, despite its description from a single specimen.
They maintain that this species may be more widely distributed across southern Aus-
tralia than the sparse records indicate.
Total length (contracted) 85 ~m, body (contracted) 44 ~m; head width 40 ~m;
trophus 20 ~m; toes 20 ~m; proximal toe thickness 5 ~m.
Ecology: From small alkaline waters or "Brackwasser". A single individual closely
resembling Berzil)s's description was recorded from Salt Creek, near Bern, S. Aust.
(l9.0°C, pH 7.8, 220 ~ cm-I, DO 9.2 mgl- l , TDS 132 mgl- I , 45 NTU) (Shiel & Koste
1979) and another in Ryan's 1 Billabong at Albury, Australia (Shiel unpublished).
Literature: Berzins 1953, Koste & Shiel 1991.
Code: 0102, 0201', 0301,0302,0801,0901,1001,1101,1201, 1302,1311,1402,
1502, 1906.
Cephalodella glandulosa Koch-Althaus, 1962
Fig. 112
C. glandulosa Koch-Althaus 1962 p. 74-76, Fig. 7.
Type locality: Stechlinsee, Gennany.
Holotype: Not designated
Description: Elongated, somewhat saccate body; foot conical, almost covered by tail,
toes dagger-like. Corona very convex; gastric glands huge, four salivary glands. A
brownish granulated mass between the gastric glands (eyespot ?). Trophus type A:
asymmetric, right ramus with two blunt tenninal teeth, longer than left ramus; on the
other hand, the left uncus is larger with two teeth, the right uncus is more delicate and
has one tooth only. Two pleural rods are attached to the base of the rami. Fulcrum
straight, not fanned but cleaved. Similar to dorystoma.
Total length 190-220 ~m, toes 29 ~m, trophus 27-31 ~m.
Ecology: Among Chara, May, August, October.
Literature: Not seen since discovery.
Code: 0103, 0201, 0301, 0402,0403,0408, 0503,0701,0902, 1003, 1301, 1402,
1501, 1606, 1808.
83
Cephalodella globata (Gosse), 1887
Fig. 113
Diaschiza globata Gosse 1887, p. 362, Fig. 8:4
C. globata, Harring & Myers 1924, p. 475-476, Fig. 31:3
Type locality: Dundee, Scotland.
Holotype: Not designated
Description: Body short, stout, round and gibbous dorsally. Head rather short, very
broad, slightly deflexed and oblique anteriorly, frontal edges puffed up, beaded.
Neck not very strongly marked. Foot very short and broad; the small tail reaches
nearly to the posterior end. Toes short, rather slender, slightly decurved ventrally, and
taper gradually to acute tips. Corona oblique and somewhat convex without projecting
lips. Trophus large and type B: fulcrum slightly expanded, manubria long, strongly
crutched. Ganglion very long and saccate; no RCO, conspicuous caudal antenna. Eye-
spot frontal, consisting of a somewhat diffuse cluster of very pale pigment granules.
Resembles C. physalis, which is, however, larger, has longer toes and a cervical eye-
spot; reimanni, which has sharp alulae on both rami, and cyclops which has a sym-
metrical trophus.
Total length 125-140 ~m, toes 22-25 ~m, trophus 20-22 ~m.
Ecology: Widespread, but not common. Raptorial? (Donner 1978, p. 167).
Literature: Harring & Myers 1924, Donner 1954, 1978, Kutikova 1970.
Code: 0103, 0202, 0205, 0302, 0402, 0504, 0601, 0701, 0801, 0901, 1001, 1006,
1101,1106, 1201,1302,1308,1402, 1501,1606,1607,1803.
Cephalodella glypha Wulfert, 1950
Fig. 114
C. gZvpha Wulfert 1950, p. 453-454, Figs. 4a-d.
Type locality: Peat-puddle, Glatzer Schneeberg, Gennany.
Holotype: Not designated
Description: Small, gibbous, distally narrowing to a keel, abdomen straight. Head
small, oblique, very deep neck. Foot short, covered by tail; toes robust, base bulbous,
tip sharp and separated by septum. Stomach green, gastric glands orange, vitallarium
dark, opaque, ganglion long, double cervical eyespot. Trophus modified type A: ful-
crum spatulate, manubria thin, curved; rami with large hanging alulae. Very similar to
C. ventripes, somewhat related to C. crassipes.
Total length 88-101 ~m, height 40 ~m, toes 20 ~m, trophus 21 ~m.
Ecology: Frequent in black sediment of peat puddle.
Literature: Kutikova 1970.
Code: 0102, 0203, 0204, 0301, 0404, 0510, 0801, 0901, 1001, 1102, 1202, 1302,
1303, 1402, 1501, 1508, 1606, 1808, 1903.
84 85

covered by tail; toes long, slightly curved dorsally, tapered evenly. Lorica soft, lateral
sulci very narrow, but start on the head, not only on the trunk, as usual. Corona
strongly convex, small lips. Ganglion large, no subcerebral glands, no eyespot.
Trophus type A: rami with strong teeth and small sharp alulae, manubria straight, bent
at the end and an indication of crutch, one basal lamella; two straight pleural rods.
Donner (1964) calls attention to some variability: the toes are bendable, manubria
cnltched or fanned, no alulae on rami.
Total length 140-145 ~m, toes 40-52 ~tm, trophus 18-20 ~m.
Ecology: Benthic, rare.

d~
Literature: Donner 1964.
e Code: 0101,0201,0301,0302,0404,0407,0504,0506,0601, 0701, 0801,0901,
1002, 1102, 1201, 1301, 1305, 1402, 1502, 1601, 1607, 1804.
113

Cephalodella graciosa Wulfert, 1956

Fig. 116

C. graciosa Wulfert 1956, p.464, Fig. 6.

Type locality: Teufelssee near Berlin, Germany.

Holotype: Not designated

Description: Very transparent, slender, lorica thin and pliable. Similar to C. gracilis,
but round, laterally not flattened, and with longer toes that are thin, abnlptly tapered.
Tail longer than foot. Corona oblique and very convex, vitellarium with six nuclei,
gastric glands elongated with numerous light globules. Ganglion very long, very small
single frontal eyespot. Trophus type B: rami with pseudoalulae, manubria asymmetric
with double crutch.
115 Total length 105~m, toes 30 ~m.
116 Ecology: In a forest pond situated in a bog, but pH 7.2-7.7.
Literature: Kutikova 1970.
Code: 0102,0205, 0302, 0404, 0504, 0509, 0802, 0901, 1002, 1103, 1201, 1301,
Fig. 113. C. globata dorsally and laterally; (d): manubrium; (e): rami and fulcrum; (f): trophus from left
without unci; (g): laterally after Harring & Myers.
1307,1402,1502,1606,1607,1701,1808.
Fig. 114. C. glypha laterally, head contracted, trophi.
Fig. 115. C. gobio laterally, trophi, and uncus.
Fig. 116. C. graciosa. (113: Wulfert 1937, (e & f) Donner 1954, (g) Harring & Myers 1924; 114: Wulfert Cephalodella gracilis gracilis (Ehrenberg), 1832
1950; 115: Wulfert 1937; 116: Wulfert 1956). Fig. 117

Furcularia gracilis Ehrenberg 1832, p. 130.

Cephalodella gobio Wulfert, 1937 C. gracilis, Harring & Myers 1924, p. 473-474, Fig. 27: 1.
Fig. 115 C. sagitta Wu1fert 1956, p. 466, Figs. 8a-c.
C. gobio Wu1fert 1937, p. 618-619, Fig. 27. Type locality: Berlin, Gennany.
Holotype: Not designated
Type locality: Floodplain pool, River Unstrut, Germany.
Holotype: Not designated Description: An extremely variable species (see Donner 1950, p. 312, and 1964, p.
271). Body rather short but slim, laterally compressed and slightly gibbous dorsal~y.
Description: Slender tapering body, widest at the head, slightly gibbous. Foot square, Head relatively short, broad and convex anteriorly, neck well marked. Abdomen In-
86 87

creases gradually in width for about two thirds of its length~ posterior third gently
rounded; lorica thin and flexible, but plates fairly distinct; foot conical and rather
short; very small tail reaches beyond midlength of the foot. Toes short, fairly slender,
very slightly recurved dorsally, tapering gradually and evenly to acute tips; however,
European forms can have straight dagger-shaped toes. Corona oblique and strongly
convex without projecting lips. Trophus fairly large, type>.A: fulcrum very slender and
slightly recurved posteriorly, but not expanded; manubria slender, rod-like, decurved
at the ends and crutched or fanned, sometimes with a bump or small alula in the mid-
dle. Rami may have denticles. The ganglion is elongate and saccate; RCO absent; eye-
spot frontal, sometimes unpigmented and difficult to see (er "var." lenticulata, be-
A
low). Donner (1950) described some deviant ecotypes. OlAty ~ VII.
Total length 125-150 !lm, toes 22-30 !lm, trophus 22-27 !lm, male 65-75 ~un.
Ecology: Cosmopolitan, common, also in saline waters.
Literature: Harring & Myers 1924, Wulfert 1956, Donner 1964, 1970, Kutikova
1970, Koste & Shie11991, Sanoamuang & Stout 1993.
Code: 0103,0202,0205,0301,0407,0408,0504,0511,0801, 0901,1001,1003.
1101,1201, 1301,1307,1402,1501,1606,1803.

I: The following "varieties" have been described

11

C. gracilis "var." lenticulata Wulfert, 1938b (Fig. 117.D). Differs from C. g.

11
gracilis in having a colourless eye with one crystal only, spatulate fulcrum. possessing
pleural rods and an epipharynx.
i\1
I1I
.!':, )
C. gracilis "var." sigmoides Wulfert, 1950 (Fig. 117.B). Differs from gracilis in the
J S-shaped toes with a bulbous base and ventrally bent tips, and a spatulate fulcrum.

C. gracilis keta, Reed 1978 (Fig. 117.E), was elevated to subspecific rank by Koste
(1978). It is closely related to C. rotunda (bryophila) ; it is apparently an ecotype
found in Sphagnum-ponds, in floating mosses and humic soil.

Cephalodella hiulca Myers, 1924

E
117
Fig. 117. C. gracilis group. Group A: C. graciliS gracilis laterally, trophus and toe (Wulfert 1956 as C.
sagitta). Group B: "var." sigmoides Wulfert, with trophi, toe and fulcrum laterally. Group C: variations
in rami. Group D: "var." lenticulata, L: lip, E: epipharynx. Group E: C. g. keta. Group F: C. gracilis
gracilis after Donner (a): trophi; (b): same ramus laterally; (d,e): other rami. (117 A: Wulfert 1956; B:
Wulfert 1950; C: Donner 1970; 0: Wulfert 1938; E: Koste 1978 p. 344; F: Donner 1964).
Fig. 118
C. piulca, Harring & Myers 1924, p. 488-489, Fig. 30:3
Also spelled C. hiulca by Harring & Myers (ibid.), Hauer(1935) and Koste (1978).
Piulea in the title of description as well as in the legend to the figure would have pri-
ority; however, this is most likely a misprint! Hiulca (Latin) means "split" or "open",
d
whereas piulca is meaningless. --;> Oo~ revied ljv d ?'J/u{cJ. if H &. j{;f
i) F' 423 !
Type locality: Oceanville and Bargaintown, NJ, D.S.A.
Holotype: AMNH 491 (co-type)
Description: Body elongate, slender, slightly curved and laterally compressed. Foot
fairly long and stout; small tail near the posterior end. Toes long, slender and slightly
recurved dorsally, moderately stout at the base and taper gradually to extremely slen-
der, acute tips. Corona oblique, strongly convex, no lips. Trophus type B: fulcrum
 88 89
eA
eye very slightly expanded posteriorly, manubria distinctly crotched. Ganglion long and
saccate; small eyspot at the posterior end, on the dorsal side. No ReO. Resembles C.
gibba.
Total length 130-135 (.lm, toes 38-40 (.lm.
Ecology: Acidophil, North America, Germany, Roumania; not rare.
Literature: Hauer 1935, Kutikova 1970.
Code: 0102, 0202, 0204, 0302, 0504, 0901, 1002, 1003, 1007, 1101, 120], 1301,
1307, 1402, 1501, 1606,1607,1803, 1903.

Cephalodella hollowdayi Koste, 1986

Fig. 119

C. hollowdayi Koste 1986, p. 147, Fig. 13.

Type material: Concepcion, Paraguay.

Holotype: Zoological Institute, University Kiel, Gennany.
m
Description: Ovoid body, short head well separated from body. Lorica thin, sulci in-
119 distinct. Foot short, covered by tail, toes long slender, dorsally curved. Salivary glands
present; no RCO; double frontal eyespot with lens. Trophus type C: rami symmetri-
cally bent with four tenninal teeth; unci with single tooth with a lamella and a little
side-corner. Manubria fonn terminal spoons that appear as crutches in dorsal view.
Reminiscent of C. gibba or paggia. The spoon-like ends of the manubria are similar to
those of misgurnus, pachyodon, tecta and obvia; but in all these species the end of the
manubrium is a ring, not a spoon.
b Total length 223 (.lm, toes 72 (.lm, trophus 40 (.lm.
Ecology: Floodplain of Rio Paraguay, pH 6.0-6.3, 65-300 (.lS. Peru.
Literature: Koste 1986, 1988a.

I c
Code: 0102, 0203, 0205, 0208,0303, 0407, 0508, 0511, 0802, 0902, 1001, 1006,
1102, 1201, 1301,1307,1403,1501, 1605,1803.

~ ~
Cephalodella hoodi (Gosse), 1886
a Fig. 120 illLf(Jor/aYt4 vhv..V''):jJ'
-, i)
Diaschiza hoodi, Hudson & Gossel886, p.2:79, Fig. 22: 15 y.
C. hoodi, Haring & Myers 1924, p.482, Fig. 28: 1
120 C. strigosa Myers 1924, Harring & Myers 1924, p. 485, Fig.29:7
C. remanei Wiszniewski 1934 p. 353, Figs. 59:17-21

Fig. 118. C. hiulca laterally, and photomicrogram of holotype.
Fig. 119. C. hollowdayi (a,e): laterally; (b): ventrally; (c,d): dorsally; (f,g,h): trophi; (i-I): a manubrium
from different sides; (m): right ramus.
Fig. 120. C. hoodi (c): rami; (e): posterior end and toes dorsally; (f,g): "var." grande. (118: Barring &
Myers 1924, photo Nogrady; 119: Koste 1986; 120: Barring & Myers 1924, Wulfert 1937).
Type locality: Loch near Dundee, Scotland.
Holotype: Not designated
Description: Fusiform body, gibbous dorsally; abdomen unusually elongate, with
dorsal plates; foot small, conical; tail prOlninent; toes short, stout, decurved, tapering
to acute tips; foot glands small, pyriform; corona oblique, with prominent beak-like
lips (rostrum); trophus type A: short, rodlike manubria, curved at end but not crooked;
90 91
two pleural rods present; rami sometimes with "pseudoalulae", with toothed inner
margin; caudal antenna with long setae; retrocerebral organ present; eyespot large, at
posterior end of cerebral ganglion.
Total length 110-195 !lm, toes 32-47 !lm, trophus 30-38 ~m, male 110-115 ~m.
A larger "var." grande (remanei var. grande) Wulfert 1937 (Figs. 120 f, g) has
been described, but it differs only in size (195 ~m), and the presence of two curved
epiphmyngeal rods.
Ecology: Cosmopolitan, rare. .
Literature: Harring & Myers 1924, Wulfert 1950, Koste 1970, Kutlkova 1970, de
Smet 1988, Koste & Shiel 1991.
Code: 0102, 0202, 0204, 0301, 0404, 0407, 0701, 0802, 0901, 1002, 1101, 1106,
1201, 1302, 1308, 1402, 1501, 1602.

Cephalodella hyalina Myers, 1924

Fig. 121
121 123
C. hyalina Myers, Harring & Myers 1924, p. 505, Fig. 32:4.
C. horni Wulfert 1960, p. 269-270, Fig. 4.

Type locality: Acid waters in Wisconsin and New Jersey.

Holotype: Not designated

Description: Body elongate, fairly slender and slightly compressed laterally; head
large, strongly oblique anteriorly; neck well marked. Abdomen very nearly parallel-
sided, rounded dorsally and ventrally; the integument is very flexible and the body
virtually illoricate; sulci obscure and difficult to trace. Corona strongly oblique and

~
decidedly convex without projecting lips. Foot fairly large and robust; tail prominent
and near mid-length. Toes long, slender, wide apart at the base, recurved and slightly
tapering, with conical tips formed by an abrupt dorsal curvature of the dorsal edge; at
this point there is often a septum. Tip usually a short, bristle-like nib forming a con-
tinuation of the ventral edge; in dorsal view, the toes are parallel but diverge suddenly 122
at the septum. Footglands large and pYriform. Mastax large 'and type B: fulcrum
spatulate, manubria strongly crutched. Ganglion very long and saccate; eyespot and
ReO absent.
trtr;(So(/. '19g(]l~ 04fJJ/v:dli /l ttfi9c! I ~124
Total length 200-245 !lID, toes 42-55 !-lm, trophus 40 ~m.
e
06J lf/M"7 cl c:'· 't/i flow. .
Ecology: Acidophil, North America, Europe.. P061V·Z1l iJt~o O(el"4' I
Literature: Wulfert 1950, 1960, Kutikova 1970.
Code: 0102, 0201, 0302, 0407, 0504, 0802, 0902, 1001, 1006, 1101, 1201, 1301,
130~1311, 1402, 1506, 150~ 1808, 1903.
Fig. 121. C. hyalina laterally, dorsally, toes dorsally, trophi.
Fig. 122. C. incila (c,d): different toes; (e,f): trophi; (g): young animal.
Fig. 123. C. innesi.
Cephalodella incila Wulfert, 1937 Fig. 124. C. intuta. (121: Harring & Myers 1924, Wulfert 1960a; 122: Wulfert 1937; 123: Barring &
Fig. 122 Myers 1924; 124: Harring & Myers 1924, Wu1fert 1960a).

C. incila Wulfert 1937, p.616-617, Fig. 39.

Type locality: In ditches, Germany.

 92
Holotype: Not designated
Description: Slender, small animal, not flattened laterally. Lorica soft, sulci widen
tenninally. Neck well visible, foot short, tail usually covers foot. Toes stout, slightly
curved dorsally and, in dorsal view, inward. Corona very convex, with lips. Four sali-
vary glands, gastric glands large, ganglion long, eyespot absent. Trophus type B: ful-
crum spatulate with basal apophysis, rami triangular without alulae, fine teeth (or
striation) terminally, manubria crutched, with double lamellae. There is a U-shaped
pleural apparatus that almost reaches the end of the fulcmm. C. paxi may be an
ecotype. Variable species according to Donner (1964, p. 272). Close to jorjicata,
which has spatulate manubria and no pleural apparatus.
Total length 130-180 ~m, toes 26-40 !lm, trophus 18-~3 [+m.
Ecology: Benthic, in saprobic sediment. ~OO(QJ~I .ttH., OI..~~lUI fI.ai.Jo./i
Literature: Donner 1964, 1970, Kutikova 1970, Koste 1976, 1987 (as limosa)
Code: 0102, 0201, 0302, 0407, 0504, 0601, 0702, 0801, 0901, 1002, 1102, 1201,
1302, 1307, 1402, 1501, 1601, 1607, 1808.
Cephalodella innesi Myers, 1924
Fig. 123
C. innesi Myers, Harring & Myers 1924, p.470, Figs. 26:3-5.
Type locality: Atlantic City, NJ, D.S.A.
Holotype: AMNH 486
Description: Body elongate, slightly compressed laterally 'and strongly gibbous
dorsally. Head small, distinctly deflexed and strongly oblique anteriorly; neck well
marked. Lorica flexible, but the plates are fairly distinct; the lateral clefts are narrow
anteriorly and increase slightly in width towards the posterior end. Foot long, conical
and extremely broad at the base; tail small. Toes relatively short and robust, very
slightly enlarged at the base and taper gradually to slender, acute tips. Corona strongly
oblique and convex without projecting lips. Footglands very large and pyriform, gas-
tric glands small. Ganglion moderately long, eyespot frontal and double, the two
halves fairly wide apart. No RCO. Trophus fairly large and of type B: fulcrum rela-
tively short, without any expansion at the posterior end; rami somewhat asymlnetrical,
the tip wavy, manubria long, crutched. Swims rapidly.
Total length 200-21 0 ~m, toes 40-44 !lm, trophus 40 !lm.
E~ology: AcidophiL.. ('J .., . . ' . _I ')
I'
LIterature: Not seen SInce dIscovery. (J..e({)~0je£i {Jy(h~," $-'hll ue.J f:;,,(. &&l.I:,O~"wL
Code: 0102,0203,0205,0302,0402,0504,0802,0902, 1001, 1006, 1101, 120f: !'J'~~
1302, 1308, 1402, 1501, 1606, 1607, 1803, 1903.
Cephalodella intuta Myers, 1924
Fig. 124
C. intuta Myers, Harring & Myers 1924, p. 500-501, Figs. 35:2-5.
93
Type locality: Loon Lake, WI, U.S.A.
Holotype: AMNH 566 (co-type)
Description: Body moderately elongate and laterally compressed; corona markedly
oblique; lorica rigid, plates distinct; toes long and slender, often curved dorsally, ta-
pering to acute tips with transverse basal septa. Mastax large with salivary glands;
trophus type B: fulcrum expanded distally, manubria crutched and with a1ulae, the
ventral one with a small median tip; rami ends strongly toothed; rami with small alulae
(easily 10sti~~!ea.~h used to prepare trophus). Gastric glands red to red-brown; ducted
RCO present; no eyespot. May be confused with forjicata. but has relatively longer
toes (toe:totallength ratio <4 in intuta, >4 inf01.1icata). According to Harring & Myers
1924,forjicata, tenuior, collactea and intuta show considerable similarity (pigmented
gastric glands, crutched manubria), but are distinguishable by the form of the toes.
Total length 115-225 !lm, toes 30-60 !lm, trophus 30-40 !lm. The North American
specimens were smaller than the European or Australian ones.
Ecology: Cosmopolitan but rare, acidophil.
Literature: Wu1fert 1960, Kutikova 1970, Koste & Shie1 1991.
Code: 0102, 0201, 0302, 0404, 0407, 0504, 0802, 0901, 1003, 1007, 1102, 1201,
1301, 1307, 1402, 1501, 1508, 1606, 1607, 1804,1807, 1903.
C. flA{u.la jw,14Ctlc.Qci~js kDSTC' 124 ~').p, t+f,;,
Cephalodella inquilina Myers, 1924
Fig. 125
C. inquilina Myers, Harring & Myers 1924, p.502, Fig. 36:1.
Type locality: WI, NJ, U.S.A.
Holotype: AMNH 387
Description: Body moderately elongate, nearly parallel-sided and laterally com-
pressed. Head short and broad, somewhat longer dorsally than ventrally; neck is well
marked. Foot large and very broad at the base; tail very small. Toes long, slender,
nearly straight, so.mewhat enlarged at the base, suddenly reduced at the ends to
minute, but very distinct claws. Corona decidedly oblique and strongly convex with-
out projecting lips. Trophus type B: the posterior end of the fulcrum is broadly and
abruptly expanded; manubria crotched. Gastric glands brown; ganglion long and
saccate; eyespot and RCO absent. The species builds a spherical container consisting
of mud, diatom shells, and feces in which it turns restlessly. Very fast swimmer when
free.
if'/;.l'
'~*";zf7~1I11 Total length 250-270 !lm, toes 62-68 !lm.
Ecology: Acidophil, in Sphagnum ponds. North America, Germany.
(,'toc Vf/W8t'1 f.cJ;; Literature: Koste 1976.
Code: 0102, 0201, 0302, 0504, 0902, 1003, 1101, 1201, 1301, 1403, 1505, 1807,
" 1903.
if isae
 94 95
Cephalodella labiosa Wulfert, 1940
Fig. 126

C. lablosa Wulfert 1940, p. 565-566, Fig.5.

Type locality: Birkhorst bog, Germany.

Holotype: Not designated

Description: Body stout, transparent, back straight, head short, wide and dorsally
broad, trapezoidal. Neck and lorica well defined, sulci wide. Corona convex with lips. e
Foot very short, tail absent, toes bulbous at base with caudal antenna, otherwise slim,
slightly curved ventrally. No salivary glands, vitellarium with eight nuclei, double
cervical eyespot. Trophus type A with thin, dorsally bent, short manubria. Related to a
hoodl. b c
Total length 94-140 !-tm, toes 21-35 I-tm, trophus 28 I-tm.
Ecology: Acidophil. 125
Literature: Wulfert 1940, 1960, Kutikova 1970.
Code: 0102,0203,0204,0301,0501,0801,0901, 1001, 1201, 1301, 1303, 1308,
1402,1501,1601,1604,1607,1702,1801,1903.

Cephalodella laisi Hauer, 1935

Fig.127

C. lalsi Hauer 1935, p. 64-65, Fig. 5. 126

Type locality: Schluchsee bog, Germany.
Holotype: Not designated

Description: Described from single specimen. Body stout but extremely compressed
laterally, head large, neck well defined, sulci visible. Corona oblique but not very con-
vex. Foot triangular, short tail, toes with basal bulb, slim, and only the ends curve
ventralIy. Trophus (figure not shown) with a fulcrum teffi1inally bent upwards,
crutched manubria. Double frontal eyespot. Koste (1978) suspects that this species is 128
a juvenile tinea.
Total length 180 !Jffi, thickness 22 !Jm, toes 53 !-tm ,
Ecology: Acidophil.
Literature: Not seen since discovery.
Code: 0102, 0203, 0205,0302,0504,0901,1001, 1007, 1101,1201, 1301, 1303, t'

1402, 1503, 1605, 1903. 129 C'J~' r.2Y·~

.......

Fig. 125. C. inquilina (a): after Harring & Myers; (b): photomicrogram ofholotype; (c): after Koste 1976;
Cephalodella lepida Myers, 1934 (d): dorsally; (e): toe laterally and trophi.
Fig. 128 Fig. 126. C. labiosa.
Fig. 127. C. laisi.
C. lepida Myers 1934, p. 10, Fig. 10. Fig. 128. C. lepida.
Fig. 129. C. licina.
Fig. 130. C. limosa. (l25: Harring & Myers 1924, Koste 1976, photo Nogrady; 126: Wulfert 1940; 127:
Type locality: Mount Desert Island, ME, D.S.A. Hauer 1935; 128: Myers 1934; 129: Wulfert 1961; 130: Koste & Poltz 1984, Wulfert 1937).
 96 97
Holotype: AMNH 537

Description: Body slender, cylindrical and almost parallel-sided. Head somewhat

deflexed and oblique, head long, well set off. Lorica flexible, but plates well defined.
[
Foot and tail small, toes long, slender, blade-shaped and curved dorsally, tapering
gradually. Corona oblique, lips present, but no beak. Trophus (not figured) type A?,
fulcrum stout and reduced in the middle portion, manubria expanded in the middle
and end in slightly swollen decurved tips. Gastric glands small, yellowish-red;
retrocerebral sack present, no eyespot. Resembles hyalina, but the retrocerebral organ
and toes distinguish it. Could be a synonym of oxydactyla which has four salivary
glands and no lips, otherwise identical. 131
Total length 225 ~lm, toes 52!lm.
Ecology: Abundant in decaying Sphagnum. also in New Jersey. 133
Literature: Kutikova 1970.
Code: 0102, 0201, 0301, 0902, 1002, 1003, 1101, 1201, 1301, 1307, 1402, 1501,
1601,1804,1807,1903.

132
-
Cephalodella licina Wulfert, 1961
Fig. 129

C. licina Wulfert 1961, p. 82-83, Fig. 6.

Type locality: Diiben Moor, Germany.

Holotype: Not designated

Description: Described from a single individual. Glass-clear animal with a round 134 135
head and an oval gibbous body hanging far over the foot, so that only the tips of toes
are visible viewed dorsally. The lorica is soft, laterally striped. Corona semicircular,
oblique, with lips. Foot small, toes with bulbous base, curved ventrally with sharp tips.
Salivary glands present, gastric glands oval, in the oesophagus a brownish mass, siIni-
lar to C. tenuior "var." pigmentata. Vitellarium grey, with 12 nuclei. Ganglion small,
single cervical eyespot. Trophus type A, rami with false alulae. Related to C.
ventripes and C. physalis.
Total length 144 /-Lm, toes 30 !lm.
Ecology: At pH 6, in October.
Literature: Kutikova 1970.
Code: 0102, 0202, 0204, 0301, 0404, 0901, 1001, 1006, 1101, 1202, 1303, 1402,
1501, 1601,1606, 1607,1703, 1903. 136

Cephalodella licinia Myers, 1924 Fig. 131. C. licinia.

Fig. 131 Fig. 132. C. lindamaya lateral, ventral, trophi, manubrium.
Fig. 133. C. lipara.
C. licinia Myers, Harring & Myers 1924, p. 503, Fig. 34:4 Fig. 134. C. macrodactyla.
Fig. 135. C. marina.
Fig. 136. C. megalocephala (a): Harring & Myers' animal with trophi; (b): Wulfert's animal with trophi
Type locality: Bog pools, Atlantic City, NJ, D.S.A. and bent toe without septum; (c): "var." rotunda. (131: Harring & Myers 1924; 132: Koste & Shiel 1991;
Holotype: Not designated 133,134, 135: Barring & Myers 1924; 136: Barring & Myers 1924, Wulfert 1961, Donner 1950).
 98
Description: Body elongate, slender and somewhat prismatic. Head a litt~e lon~er
than wide and obliquely truncate anteriorly. Neck not strongly marked; lonca faIrly
rigid, plates well marked. Foot conical and moderately large; tail small.and. rou.nded,
unusually far back on the foot. Toes long and slender, the terminal portIOn IS slIghtly
decurved ventrally and tapers to very slender, bristle-like tips. Corona oblique and
slightly convex without projecting lips. Trophus type A: the fulcrum is slightly ex-
panded at the posterior end, manubria slender, rod-like and not crutched. Gastric
glands small ganglion very long and saccate, eyespot and RCO absent.
Total length 105 ~un, toes 28 r-tm.
Ecology: Acidophilic, rare.
Literature: Not seen since discovery.
Code: 0102, 0201,0301,0901, 1002, 1003, 1101, 1201, 1301, 1306, 1402, 1503,
1506, 1605, 1803, 1903.
Cephalodella limosa Wulfert, 1937
Fig. 130
C. limosa Wulfert 1937, p. 615-616, Fig. 25.
Type locality: Saprobic mud and ditches, Germany.
Holotype: Not designated
Description: Fairly slender, back- and ventral line convex, neck well marked. Lorica
distinct, sulci well visible. Foot and tail small, toes straight, uniformly tapere,d. Corona
slightly oblique, very convex, with lips. Four salivary glands, vitellarium with eight
nuclei. Trophus type B: fulcrum slim, slightly spatulate, no teeth on rami, manubria
with fan, with double lamellae. Ganglion long, eyespots absent. Closely related to
inciIa, with which it was synonymised in Koste (1978), but restored to specific status
(in Koste 1984). In this latter publication (p.1 08) Koste compares this very difficult
group of similar small species occupying the same niche of decaying vegetation and
slime; limosa differs from incila by the absence of pleural rod, rami teeth, and perhaps
by fanned (not crutched) manubria. C. tenuior and rigida (which has rami teeth) also
belong to this group.
Total length 130-160 ~m, toes 32-36 ~m, trophus 19-21 ~m.
Ecology: Saprobic, not rare. Europe.
Literature: Donner 1964, 1970, Kutikova 1970, Koste 1984.
Code: 0102, 0201, 0302, 0406, 0506, 0801, 0901, 1001, 1006, 1101, 1201, 1302,
1306, 1402, 1501, 1601, 1607.
Cephalodella lindamaya Koste & Shiel, 1986
Fig. 132
C. lindamaya Koste & Shiel 1986, p. 95-96, Figs. 3-4.
Type locality: Stock dam, 1 km south of Copping, Tasmania.
Holotype: South Australian Museum V4019. ColI. R. J. Shiel, 1985
99
Description: Body short, stout; head broad, deflexed; lorica flexible, plates indistinct;
toes relatively long, basally thickened; terminal claws curved with acute tips, four dis-
tinct spinules in a row on the inside edge of the claw. Mastax large; trophus type B:
fulcrum long, narrowest in the centre, flaring at distal end; manubria unusual, termi-
nally crooked, leaf-shaped; rami with denticulate inner margin behind tips, uncus with
one tooth and basal lamella; foot glands large, club-shaped; eyespot not recorded.
Total length (contracted) 245 ~lm, ; toes 68 !lm, (spinules 4-6 flm, ); trophus 43 r-tm.
Ecology: Tasmanian endemic. Known only from an acid stock dam at Copping, east-
ern Tasmania. 21.7°C, pH <4.0, 80 ~S cm-I.
Literature: Koste & Shie1 1991.
Code: 0102, 0201,0302,0407, 0510, 0511, 0802,0902, 1001, 1101, 1201, 1303,
1306, 1314, 1403, 1501, 1903.
Cephalodella lipara Myers, 1924
Fig. 133
C. lipara Myers, Harring & Myers 1924, p. 512-513, Fig. 36:5.
Type locality: Ditch with Sphagnum NJ, D.S.A.
Holotype: Not designated
Description: Body extremely short, stout and nearly cylindric. Head enOlnlOUS; its
length is nearly half the length of the body. Neck marked by a slight constriction.
Abdomen very short and abruptly truncate posteriorly; lorica very flexible, dorsal and
ventral plates separated by a very wide cleft. Foot indistinct and the tail very small.
Toes short, stout and blade-shaped; dorsal edge decurved, ventral edge nearly
straight, with a slightly decurved, claw-like tip. Corona slightly oblique without pro-
jecting lips. Trophus huge, type A: fulcrum slightly expanded and decurved at the
posterior end; the manubria are slender and rod-like. Length the mastax is fully half
the length of the body. Ganglion very large and saccate; no eyespot and no RCO. Re-
sembles C. physalis, but has shorter toes, is much stouter, and has no eyespot.
Total length 130-140 ~m, toes 22-24 ~m.
Ecology: Acidophil, D.S.A., New Zealand.
Literature: Russel 1960.
Code: 0103, 0201, 0301, 0503, 0802, 1001, 1101, 1201, 1302, 1402, 1502, 1605,
1803, 1903.
CephaIodella macrodactyla (Stenroos), 1898
Fig. 134
Furcularia macrodactyla Stenroos 1898 p. 134, Fig. 2: 15
C. strepta Myers, Harring & Myers 1924, p. 509-510, Fig. 35:1
Type locality: Finland.
Holotype: Not designated
100
Description: Body elongate, fairly slender, s!ightly gibbous dorsally and faintly
curved. Head relatively long and tapers very shgh.tly from ~he neck towards the c~­
rona; it is protected by a distinct lorica, which falls mto defimte folds when th~ head 1.S
retracted, as in the genus Trichocerca. Neck well marked. Foot large and cOl1lcal; tall
very small reaching near the middle of the foot. Toes very. long, recu:ved, stou~ at ~he
base and taper gradually to acute tips. Corona slightly obhque and ~lthout proJectmg
lips; trophus type A, but not well described; the fulcrum and manubna rod-shaped ~nd
slightly curved, but not crutched. Gastric glands large and usually of a deep p.mk
color. Ganglion fairly large and saccate. Neither eyespot nor. RCO present. SWl~S
slowly. Described from single individuals, but has been found 111 Europe as well as 111
North America. Russel (1960) recordedstrepta from New Zealand. Koste (1978) con-
siders it a dubious species.
Total length 100-152 /-lm, toes 40-56 /-lm. North American specimens Cc. strepta)
are small.
Ecology: Sediment, acid pools. Cosmopolitan.
Literature: Harring & Myers 1924, Eriksen 1969.
Code: 0101, 0201,0301,0402,0509,0901, 1002, 1003,1101,1201, 1307, 1402,
1501,1605,1803,1903.
Cephalodella marina Myers, 1924
Fig. 135
C. marina Myers, Harring & Myers 1924 p. 469-470, Fig. 26:7.
C. enitedia Myers, Harring & Myers 1924 p. 468, Fig. 27:7
r
~ I doati'I,l)-.~;{.
0SJJ,tt ~.~ e
Type locality: Tide-pools, NJ, D.S.A. . fO)~)~/'
Holotype: AMNH 466 (co-type) 1O.l'~.~?'.' d?ev..U!tL\
o{t( tf..1fto..J! v
Description: Body moderately elongate, spindle-shaped, very slight~y compress~d
laterally. Lorica very thin and flexible; foot fairly long, stout and comcal, s~al1 tall.
Toes short, very slender, slightly decurved and taper gradually to very acute tIps. Co-
rona decidedly oblique and somewhat convex without projecting lips. Trophus type
A: fulcrum slightly expanded posteriorly, manubria slender, rodlike and not crotched.
Gastric glands small; eyespot frontal and double, the two spheres rather wide apart.
No RCO; dorsal antenna is unusually far back on the head.
Total length 160-165 /-lm, toes 28-30 /-lm.
Ecology: In tidal pools, Atlantic City, NJ, D.S.A.
Literature: Not seen since discovery.
Code: 0103,0203,0205,0301,0901, 1002, 1003, 1101, 1201, 1301, 1308, 1402,
1501, 1605, 1803, 1906.
Cephalodella megalocephala (Glasscott), 1893
Fig. 136
Furcularia megalocephala Glasscott 1893, p. 56, Fig.4:3
C. megalocephala, Harring & Myers 1924, p. 494-496, Figs. 32:5-7.
101
Type locality: Ireland.
Holotype: Not designated
Description: Body stout, dorsally gibbous; head very large, broad, ciliary field ex-
tremely oblique, ventral; apical field with two large cirri; lorica thin, flexible, plates
indistinct; dorsal median sulcus may have convex rather than concave connecting in-
tegument; body (foot ?) 2-segmented; toes short, decurved, sharply pointed, the toes
are bendable but septa are not always observable. Trophus of peculiar type E: simple
rod-shaped fulcrum; manubria two sigmoid curved slender rods; rami appear semicir-
cular from above; unci with numerous teeth, rake-like. Sometimes triangular, lamel-
lar, thin epi-pharynx distinguishable; no eyespot; RCO transparent.
Total length 195-210 /-lm, toes 34-38 /-lm, trophus 30 /-lID.
Several ecotypes have been described as "forms":
Fonn rotunda Donner (1950) is round in cross-section and is also mentioned by
Wulfert (1937). These are large animals, up to 325 /-lm.
Fonn compressa Donner 1950 is laterally strongly compressed.
Ecology: Cosmopolitan; mud flats, beaches, sand, periphyton of freshwater margins
of running water.
Literature: Wulfert 1937,1961, Donner 1950, Kutikova 1970, Koste & Shiell991.
Code: 0103,0201, 0305,0511,1102, 1104, 1101, 1201,1301, 1305, 1308,1402,
1508, 1606, 1604.
) Cephalodella megalotrocha Wiszniewski, 1934
Fig. 140
r
C. megalotrocha Wiszniewski 1934, p. 352, Figs. 28:13-15
Type locality: Lake Wigry, Poland.
Holotype: Not designated
Description: Body long, cylindrical, head unusually long; foot wide, large, tail small;
toes short, decurved ventrally, tightly adjacent in dorsal view. Lorica faintly indi-
cated, lateral sulcus widens distally. Corona slightly oblique. RCO present, on occa-
sion brown (?). Trophus type A, short manubria. Not seen since description (Kutikova
?), probably identical with C. parasitica.
Total length 150-180 /-lm, toes 18-23 /-lID, trophus 24 ~m.
Ecology: In psammon.
Literature: Kutikova 1970, Koste 1978
Code: NOT CODED, NOT IN KEY.
Cephalodella melia Myers, 1924
Fig. 137
C. melia Myers, Harring & Myers 1924, p. 493-494, Fig. 32:2.
Type locality: NJ, D.S.A.
102 103

Holotype: Not designated

Description: Body elongate, somewhat prismatic and gibbous dorsally. Head rather
small, but relatively long in comparison with its width; neck well marked. Foot long,
conical, tail prominent, but does not cover the foot. Toes fairly long, slender, enlarged
at the base and end in small conical tips, prolonged by a fairly long, bristle-like nib,
continuing the line of the dorsal edge. Corona oblique, strongly convex, without lips.
Trophus type A: fulcrum curves upwards at the posterior end; manubria not crutched.
Symbiotic zoochlorellae are present in abundance in the walls of the stomach. Bladder
very large; ganglion long and saccate; eyespot and ReO absent. Close to C. nelitis, but
v;/ the toes and zoochlorellae are diagnostic.
138 Total length 130-145 ~m, toes 28-32 ~m.
137 140 Ecology: Acidophilic; was fairly common, then disappeared.
139 Literature: Not seen since discovery.
Code: 0102, 0201, 0301, 0503, 0901, 1003, 1101, 1201, 1301, 1303, 1308, 1402,
1502, 1506, 1605, 1803,1901, 1903.

Cephalodella mineri Myers, 1924

Fig. 138

C. mineri Myers, Harring & Myers 1924, p. 471, Fig. 26: 1.

Type locality: Brackish stream, NJ, D.S.A.

Holotype: Not designated

Description: Body rather short and nearly cylindric; head moderately large and
somewhat deflexed; lorica thin and flexible, but plates fairly well defined. Foot rela-
tively short, conical and robust; very small tail. Toes short, strongly decurved, ex-
tremely broad at the base, tapering gradually. The foot glands are large and ovate. Co-
143 rona slightly oblique and convex, no lips. Trophus type B: fulcrum rather stout,
slightly expanded posteriorly; manubria long, very slender and crutched. Gastric
glands small. Ganglion elongate and saccate; eyespot double and frontal, the two pig-
ment spheres fairly wide apart, no RCO.
Total length 125 ~m, toes 27 ~m.
Ecology: Was abundant in a brackish stream among Fontinalis.
Literature: Not seen since discovery.
Code: 0102, 0203, 0205, 0302, 0504, 0901, 1001, 1101, 1201, 1302, 1308, 1402,
1501, 1803, 1906.

Cephalodella minora Wulfert, 1960

Fig. 139
Fig. 137. C. melia. Fig. 138. C. mineri. Fig. 139. C. minora with trophus sideways and distal end dorsally
with footglands and bladder. Note toes. Fig. 140. C. megalotrocha. Fig. 141. C. mira. Fig. 142. C.
misgurnus living animals dorsally, laterally without and with hypodermis stripes, toes dorsally and later-
C. minora Wulfert 1960, p. 271-272, Fig. 7.
ally. Trophus; (a): fulcrum and (b): manubrium laterally; (c): unci in two views; (d): inner rami in two
views. Fig. 143. C. montana. Fig. 144. C. mucosa laterally, photomicrograph of cotype. (137,138: Harring Type locality: Wildenhain bog, Germany.
& Myers 1924; 139: Wulfert 1961; 140: Kutikova 1970; 141: Myers 1934; 142: Wulfert 1937, Donner Holotype: Not designated
1950; 143: Myers 1942; 144: Myers 1934, photo Nogrady).
104
Description: Very slender, cylindrical, tra~spar~nt. He,ad,large, dorsally wide; body
consists of two segments, the first long one IS strIped, SImIlarly, the foot has two s~g­
ments. Toes unique: they consist of a cylindrical part, and an equally long needle-lIke
end. Footglands are very large and encase the lowe~ part of the blad~er. Corona en-
tirely ventral (i.e., extremely oblique). One large sallvary gland, gastnc glands large,
clear, filled with spheres. Ganglion large with a small sack, no eyespot. Trophus ap-
parently type E, but only a lateral view is available; epipharynx present. Moves rap-
idly; related to C. megalocephala with the ventral corona and segmented body, but the
toes and segmented foot are unique in the genus.
Total length 105-120 ~m, toes 15 !lm.
Ecology: Acidophilic.
Literature: Kutikova 1970. _ Total length 165-190 ~m, toes 49-61 !lm, trophus 22 !lm.
Code: 0103, 0201, 0305,0901,1002,1003,1203, 1305, 1401,1501, 1606, 1903.
Cephalodella mira Myers, 1934
Fig. 141
C. mira Myers 1934, p. 6-7, Fig. 6.
Type locality: Mount Desert Island, ME, D.S.A.
Holotype: Not designated Ijel l /{2,({'b) I.'l /JA/.:Y ;. .qat
. \J
Description: Body cylindrical, slim, head small, neck not well defined. Lorica very
thin, plates indistinct, FogtTobus.!?conical, half-covered by tail. Toes. short, b~ade­
shaped, slightly curveQ dorsally.Corona slightly convex. Large palr of sallvary
glands, gastric glands Claimed to be absent in the original description, but Koste
(1970) shows two normal gastric glands. Stomach with zooch10rellae and brown stor-
age granules. Double frontal eyespots with crystals. Trophus unique, type F, consist-
ing of delicate rods and lamellae; probably for pumping only.
Total length 124-200 !lm, toes 22-42 !lm, trophus 30-56 !lm.
BerziQ.s (1976) describes a subspecies, C. m. nidicola, that builds a gelatinous
spherical dwelling of 150-160 !lm in diameter, and never leaves this; hence the animal
could not be studied properly. However, the trophus is unmistakeable. BerziQ.s's
specimens were large, 220-270 !lm long, toes 40-44 !lm, trophus 43-50 !lm. The sub-
species was found in an acidic (pH 6.0, 67 !lS) bog near Aneboda, Sweden.
Ecology: Acidophilic, also in decomposing Sphagnum and Utricularia, not rare. Eu-
rope.
Literature: Koste 1970, BerziQ.s 1976.
Code: 0103, 0203, 0205, 0208,0306,0511,0802,0901,1003,1101,1201,1302,
1307, 1402, 1501, 1607,1901, 1903.
Cephalodella misgurnus Wulfert, 1937
Fig.142
C. misgurnus Wulfert 1937, p. 620-621, Fig. 29.
105
Type locality: Muddy streams, Germany.
Holotype: Not designated
Description: Body elongate, widest in posterior third; head and trunk conspicuously
loricate, three large and two smaller plates (see also pentaplax)~ neck clearly defined;
toes long, cca. one-third total length, slightly wider at base, terminating in acute tips;
mastax small with two small salivary glands; trophus type C, symmetrical: fulcrum
straight, flaring distally; manubria with straight shafts (no lamellae), ending distally in
a distinctive ring; unci robust with quadratic plate on outer half; subcerebral gland
present; two frontal eyespots with crystalline lens in common capsule. Related to C.
pachyodon, but has a more rigid lorica.
Ecology: In mud, even just moist mud, in streams; also in ditches. Europe, Australia.
Literature: Donner 1950, Kutikova 1970, Koste & Shiel 1991.
Code: 0102, 0203,0205,0208,0303,0508,0901, 1003, 1102, 1201, 1301,
1402, 1502.
Cephalodella montana Myers, 1942
Fig. 143
C. montana Myers 1942, p. 276, Fig. 23:3.
Type locality: Mountain Lake, Pocono Plateau, PA, U.S.A.
Holotype: PANS 959
Description: Body elongate, cylindrical, somewhat swollen posteriorly. Head long,
lorica thin, plates ill-defined, sulci parallel-sided. Foot stout, conical, toes straight, ta-
pering gradually. Corona convex, without lips. Trophus type B, manubria strongly
crutched (no figure given). No eyespots. Resembles C. elongata and zeteta.
Total length 135 !lm, toes 20 !lm.
Ecology: Among macrophytes, pH 7.0.
Literature: Not seen since discovery.
Code: 0103, 0201, 0302, 0504, 0901, 1003, 1101, 1201, 1302, 1306, 1402, 1501,
1607.
Cephalodella mucosa Myers, 1934
Fig. 144
C. mucosa Myers 1934, p. 4-5, Big.3.
Type locality: Mount Desert Island, ME, D.S.A.
Holotype: AMNH 831 (cotype)
Description: Body prismatic, elongate, head somewhat deflexed, neck well marked.
Lorica firm, plates well defined, foot and tail short, toes very long, blade-shaped and
lanceolate. Foot glands long. Corona obliquely convex, small lips, ganglion long, cer-
 106 107

vical eyespot. Trophus type A (figure not available), manubria very short, slender and
abruptly decurved.
Total length 125 ~m, toes 30 ~m.
Ecology: Acidophil, was not rare in decaying Sphagnum.
Literature: Not seen since discovery.
Code: 87 0102, 0202, 0204, 0301, 0901, 1003, 1101, 1201, 1301, 1306, 1402, 1502,
1606, 1607, 1903.

~
Cephalodella mucronata Myers, 1924
Fig. 145

~
C. mucronata Myers, Harring & Myers 1924, p. 510-511, Fig. 36:2-4

Type locality: WI, NJ, D.S.A. 146

Holotype: AMNH 252 (cotype)
145
Description: Elongate slender body, rigid lorica extending beyond end of foot; longi-
tudinal sulci deep; foot sheath has triangular ventral point and a stout dorsal spine,
separated by a deep, rounded sinus; toes exceptionally long (about body length) slen-
der recurved and sharp; trophus type B: fulcrum long and straight, manubria slender,
distally crooked, rami denticulate on inner margin; RCO present; no eyespot.
Total length 265-275 ~m, toes 120-140 !-lID, trophus 36 !-lID.
Ecology: Pantropical-subtropical warm stenotherm, but widespread in the temperate
D.S.A. as well. Occurred in TasIDania between 9-29°C, pH 3.1-7.6 at low conductiv-
ity. Algivore and occasional raptor.
Literature: Harring & Myers 1924, Kutikova 1970, Koste 1972, Koste & Shiel1991. c
Code: 0101, 0201, 0302,0407, 0504, 0802, 0902, 1002, 1103, 1201, 1301, 1307,
1403, 1501, 1903.
147 148
Cephalodella mus Wu1fert, 1956
Fig. 146
Fig. 145. C. mucronata.
Fig. 146. C. fiUS.
C. mus Wulfert 1956, p. 465, Fig. 7. Fig. 147. C. nana (a): Barring & Myers' C. nana; (b): Barring & Myers' C. cuneata; (c): Wulfert's C.
nana.
Type locality: Teufelssee near Berlin, Germany. Fig. 148. C. nelitis. (145,148: Barring & Myers 1924; 146: Wulfert 1956; 147: Harring & Myers 1924,
Wulfert 1940). .
Holotype: Not designated

Description: Very small, stout, round, and bulbous body. Head short, dorsally two Code: 0102, 0204, 0301, 0404, 0901, 1008, 1103, 1104, 1202, 1302, 1308, 1402,
small auric1es are visible. Lorica well defined, sulci wide, but it does not cover the 1501, 1601, 1604, 1606,1607,1903.
foot (as in ventripes). Foot very short, tail well separated, covers foot and basis of
toes. Caudal antenna conspicuous. Toes soft, uniformly tapering, ventrally deflexed.
Corona oblique, convex, with lips, cervical eyespot. Trophus type A: manubria S- Cephalodella nana Myers, 1924
shaped with median loop, rami with alu1ae. Fig. 147
Total length 87 !lID, toes 27 !lID.
Ecology: Acidophil, pH 4.5-5.0. C. nana Myers, Harring & Myers 1924, p. 491-492, Fig. 29:1
Literature: Kutikova 1970. C. cuneata Myers, Harring & Myers 1924 p. 505, Fig. 29:2
 108
Type locality: Corduroy Creek, NJ, U.S.A.
Holotype: Not designated
Description: Body short, conical, tapering gradually from corona to base of toes;
head large, ca. half the length of body, and wider than abdomen; lorica moderately
flexible, plates distinct; foot short with conspicuous caudal antenna, covered by tail;
toes long, slender, set wide apart at base with gentle sigmoid curve, tapering to
bristle-like tips; corona oblique with prominent beaklike lips; mastax very large;
trophus type A: fulcrum slightly expanded distally; manubria slender, slightly
clubbed and recurved ends but not crutched; salivary glands small; eyespot cervical,
no retrocerebral organ. Wulfert's (1940) animals were not conical.
The synonymy of nana (Fig. 147a) and cuneata (Fig. 147b) has been established
by Myers himself (see Hauer 1935, p. 66). However, in the original description and
figure of Myers, cuneata has no eyespot (Fig. 147b). Very close to belone and
euknema, but the toes are diagnostic: long and straight in belone, abruptly upturned
tips in euknema, and more or less S-shaped in nana.
Total length 105-160 !-tm, toes 35-52 !-tm, trophus 30-34 !-tm.
Ecology: Acidophil, Europe, North America, Australia.
Literature: Wulfert 1940, Kutikova 1970, Koste & Shiel 1991.
Code: 0102,0204,0301,0501,0802,0901,1004,1102,1201,1301, 1309, 1311,
1402, 1501, 1601, 1606, 1803, 1903.
cephalodell~':;.li:t8 Myers, 1924 .eNd., litt'PfC
(,J /
C. nelitis Myers, Harring & Myers 1924, p. 493, Fig. 32: 1. jYi"l.
SPflJ.IOJed ~ow, ~)c.eJ,r DO/Jed OLI
Type locality: Ma~s Landing, NJ, D.S.A. 1~91.-4:'o~ck..e:1e ~Tt:;e~ei2J~' q{'jI.J 1<l'{1iAv~;"\
Holotype: Not deSIgnated ( l,et411 (A,V:P ;61). 1o~ -s:pecH'S
q Type locality: Bornstedt, Germany.
UlCLM a~~ov'J ..J(Jtl..•lL~L.fu'i." I Holotype: Not designated
......j r\ .
Description: Body elongate, very slender and somewhat prismatic. Head fairly
small, strongly oblique; lorica very thin and flexible, but the plates are fairly well
marked; foot short and conical, but broad at the base; the tail is very small. Toes
short, strongly decurved ventrally and very slender; from the slightly bulbous base
they taper very gradually to acute tips. Corona oblique, strongly convex without lips.
Trophus type A: fulcrum stout, slightly expanded posteriorly; manubria rod-like,
very slender and not crutched. Ganglion elongate and saccate; eyespot and ReO are
absent. Related to C. melia, but the latter has different toes and zoochlorellae.
Totallength 135 /lm, toes 22 /lm.
Ecology: Acidophilic, among submerged Sphagnum.
Literature: Not seen since discovery. . Total length 290-320 !lm, toes 65-70 !-tm, trophus 36-39 /lm.
Code: 0103, 0201, 0301, 0901, 1002, 1003, 1101, 1201, 1301, 1308, 1402, 1501,
1605, 1803, 1903.
Or also C. sp.{v1W'1 v'Iee,!,J) i4;USP 1164
109
Cephalodella obvia Donner, 1950
Fig. 149
C. obvia Donner 1950 p. 319-320, Fig. 17.
Type locality: River Thaya, Bohemia, Czech Republic.
Holotype: Not designated
Description: Robust, transparent, cylindrical body, lateral sulci parallel. Foot short,
tail covers foot, toes uniformly tapered; corona strongly covex. Gastric glands large,
vitellarium with eight nuclei, mastax with two pairs of salivary glands. Trophus
asymmetrical, type C (transition to B ?): the left side is stronger; fulcrum spatulate,
rami with three teeth on the right, none on the left, unci with a wide lamella, manubria
end in an open ring, the upper part of which has a small bump. Behind the rami a
subuncus. Ganglion long, two frontal eyespots. Closely related to C. pachyodon and
misgurnus.
Total length 152-172 !-tm, toes 44-46 !lm, trophus 20 !lIn.
Ecology: On mud and sediment in an inundated lawn and in oxbows of the River
Thaya in summer, numerous.
Literature: Kutikova 1970.
Code: 0102, 0203,0205,0303,0402,0407,0508,0509,0801,0901,1002,
1102, -1201, 1307, 1402, 1501, 1607, 1702, 1802.
J~ou;
Cephalodella oxydactyla Wulfert, 1937
Fig. 150
lUJ, ?~
,:;:e~e..."ce ~ C. oxydactyla Wulfert 1937 p. 621-622, Fig. 30.
Description: Body elongate, prismatic, hyalin; head long, foot wide, conical, cov-
ered by tail. Dorsal sulcus wide, lateral sulci naITOW; corona convex, no lips. Foot
short, conical, covered by tail toes dorsally bent, uniformly tapered. Four salivary
glands, large gastric glands, vitellarium with eight nuclei. Retrocerebra1 organ
present, no eyespots. Trophus type A, rami without teeth, separation oval, manubria
S-shaped, end curved and thickened. Related to C. tenuiseta americana, but shorter
toes and different manubrium. The species was not seen since its discovery; perhaps
identical with lepida, from which it differs only in the absence of a lip; there is no
figure of the trophus for C. lepida.
Ecology: In a pond in September.
Literature: Kutikova 1970.
Code: 0102, 0201, 0301, 0510,0802, 0903, 1003, 1102, 1201, 1301, 1307, 1403,
1501, 1607, 1702, 1804.
 110 111

Cephalodella pachydactyla \\lulfert, 1937

Fig. 151

C. pachydactyla Wulfert 1937, p. 606-607, Fig. 15

C. nodosa Wulfert 1937, p. 608-609, Fig. 18.

Type locality: Oxbow of River Elster, Gennany.

Holotype: Not designated

Description: Body sturdy, slightly gibbous, laterally not compressed~ corona some-
what oblique, convex, with slight lips. Foot short, covered by dOIned tail that has a
small terminal appendage. Toes stout, straight, tips abruptly tapered and curved
dorsally; in C. nodosa, tip of toe is a bristle with a small globular swelling before the
bristle (Fig. 151 b) Several small salivary glands plus one large one on the left side;
gastric glands large, with shiny globules. Ganglion long, narrow, ReO present; dou-
ble frontal eyespots. Trophus type B: fulcrum strongly spatulate, rami without teeth
149 with large sharply pointed alulae, unci with one tooth; manubria with very large
150 crutch carrying a semicircular alula on the upper branch of the crutch (this lamella is
missing in nodosa). One (sometimes two) long pleural rod. Related to C. denfala
which is eyeless and has no pleural rod, and C. penfaplax, which has different toes,
two pleural rods, rami toothed and asymmetrical.
Total length 135-160 !lm, toes 20-44 ~m, trophus 24-29 ~.Lm. Individuals found in
acidic habitats are smaller, in alkaline habitats larger.
Ecology: In periphyton and Sphagnum pools.
Literature: Koch-Althaus 1963, Kutikova 1970.
Code: 0102,0203,0205,0302,0404,0504, .0701,0802,0901, 1001, 1102, 1201,
1301, 1306, 1402, 1502, 1503, 1601, 1607.

Cephalodella pachyodon Wulfert, 1937

Fig. 152

C. pachyodon Wulfert 1937, p. 619-620, Fig. 28,

Type locality: Ditch at Zeitz, Germany.

152
151 B large salivary glands, large gastric glands. Long ganglion but no RCO, only
Fig. 149. C. obvia laterally, trophi, uncus, and manubrium.
Fig. 150. C. oxydactyla laterally, trophi, and manubrium.
Fig. J51. C. pachydactyla (A): laterally, dorsally, head with ganglion and ReO, toes; (B): the synonymous
C. nodosa.
Fig. 152. C. pachyodon dorsally and laterally, trophus; manubrium, uncus, and fulcrum, laterally. (149:
Donner 1950; Wulfert 1937; 150,151,152: Wulfert 1937).
Holotype: Not designated
Description: Stout, prismatic, head short; lorica strong, sulci distinct. Foot dorsally
narrower than body, covered by keeled tail. Toes long, base bulbous, slightly
decurved dorsally; in dorsal view the tips flare. Corona straight, convex, no lips. Two
subcerebral glands; double frontal eyespots. Trophus type C, unusually sturdy: ful-
crum dorsally and laterally spatulate, rami have frontal pegs, unci powerful, thick-
ened, with a blunt tooth; manubria with terminal ring. Resembles C. misgurnus,
which, however, has five sturdy lorica-plates.
Totallegth 220-240 !lm, toes 60 !lm, trophus 28 !lm.
Ecology: In a slowly flowing pasture ditch, not rare.
Literature: Kutikova 1970.
Code: 0102,0203,0205,0303,0409, 0508,0801,0902, 1002, 1102, 1201, 1301, -
1303, 1307, 1402, 1502, 1803.
112 ID

Cephalodella paggia Koste & Robertson, 1983

Fig. 153

C. paggia, Koste & Robertson 1983, p. 231-232, Figs. 2 a-e.

Type locality: Lago Camaleao, (Ilha de Marchantaria, Rio Solim6es, Amazonas,

Brazil).
HoLotype: Inst. Nat. de Pesquisas de Amazonia, Manaus, Brazil.

Description: Described from preserved material. Large animal, gibbous; head large,
corona convex but not oblique. Foot and tail small, toes long, dorsally curved ending e
in claws. Salivary glands not well visible, ganglion long, no eyespots. Trophus type d
C: fulcrum dorsally and laterally spatulate, with basal apophysis; has sickle-shaped
projection with lamellae; inside of rami serrated but no teeth, manubria with short
lamella and terminal ring. At first glance similar to the large form of C. gibba
(Wulfert 1937, length 450 [.tm).
Total length 396 ~m, toes 126 [.tm, trophus 72 [.tm.
Ecology: Not specified. Probably endemic.
c
Literature: Not seen since discovery.
Code: 0102, 0201, 0303, 0508, 0601, 0803, 0903, 1001, 1101, 1201, 1301, 1307, A B
1403,1505, 1607, 1801. 153 154

Cephalodella panarista Myers, 1924

Fig. 154

C. panarista Myers, Harring & Myers 1924, p. 478-479, Figs. 31:4-7.

Type locality: Four Mile Run, Washington, DC, D.S.A.

Holotype: Not designated

Description: Body large, elongate, slender; dorsal margin curves downwards

posteriorly to base of foot; integument very flexible, plates indistinct; toes very long,
stout and recurved, tapering to acute tips; occasionally a dorsal tooth-like spine one-
third of length along toes; footglands extremely long, clubbed; mastax large, trophus
robust, type D: fulcrum long, straight, slightly expanded posteriorly; manubria short, 155
recurved posteriorly but not crooked; with large basal plate; unci have typical single
tooth. No RCO; eyespot frontal with front part of capsule colourless resembling a
lens.
Total length 280-500 ~m, toes 80-124 /lm, trophus 55-72 ~m. The Australian 156 157
specimens of Koste (1981) are enormous (480-500 /lm). Those listed by Koste &
Shiel (1991) are smaller (360- 375 ~m), the same as the North American ones. Fig. 153. C. paggia (a): laterally; (b): trophusventrally; (c): claw; (d): manubrium; (e): rami and fulcrum
Ecology: Rare but widespread, North America, Roumania, Australia. In Australia pH laterally. .
5.4, temp. 310C, 25 /lS.cm-1. Fig. 154. C. panarista (A): Australian specimen of Koste, not fully extended; (B): Amencan and Austral-
Literature: Kutikova 1970, Koste 1981, Koste & Shiel 1991. ian specimen of Harring & Myers and of Koste & Shiel.
Fig. 155. C. papillosa.
Code: 0102, 0202, 0205, 0208; 0304, 0407, 0503, 0802, 0903, 1002, 1101, 1201, Fig. 156. C. parasitica Scale bars 50 and 10 f . L m . . .
1302, 1307,1312, 1403, 1501, 1606, 1803. Fig. 157. C.paxi. (153: Koste & Robertson 1983; 154: Koste 1981, Harnng & Myers 1924; 155: Harrmg
& Myers 1924; 156: Koste & Shiel 1991; 157: Wulfert 1959).
114 115

Cephalodella papillosaMyers, 1924 Cephalodella paxi Wulfert, 1959

Fig. 155 Fig. 157

C. papillosa Myers, Harring & Myers 1924, p. 506-507, Fig. 34:7 C. paxi, Wulfert 1959, p. 61-62, Fig. 12.

Type locality: Oceanville, NJ, U.S.A. Type locality: Wet moss in a spring, Germany.
Holotype: Not designated Holotype: Not designated

Description: Body relatively short, laterally compressed and strongly gibbous
dorsally~ head fairly large and obliquely truncate anteriorly; neck deeply constricted.
Lorica fairly rigid and the plates well marked. Foot moderately short and broad at the
base; the tail is very small; toes very long, slender and slightly decurved; from the
base they taper evenly and gradually to acute tips. The lateral antennae are of unusual
fonn; small tufts of sensory setae are seated on small, slender conical tubules (there is
no category for this unique detail in the code). Corona oblique and strongly convex
without projecting lips. Trophus type A: fulcrum slightly expanded posteriorly;
manubria rodlike and not crutched. Ganglion elongate, saccate; no RCO and eyespot.
Total length 135 !lm, toes 43 !lm.
Ecology: Rare, acidophilic.
Literature: Not seen since discovery.
Code: 0102,0201,0301,0901,1006,1101,1201,1301,1308, 1402, 1501, 1606,
1607,1803,1903.
Cephalodella parasitica (Jennings), 1900
Fig. 156
Pleurotrocha parasitica Jennings 1900, p. 84, Fig. 16: 13.
C. parasitica, Harring & Myers 1924, p. 511-512, Fig. 36:6.
Type locality: Pond near Lake St. Clair, MI, D.S.A.
Holotype: Not designated I/AA~e : k;',u.A9 b
,i;lIe. TO 15Sv]{U (,2,l;t:7c) p. /..,0(;,
Description: Body fusifonn, curved and gibbous dorsally; head unusually long, ta-
pers from neck to corona; integument flexible, no sign of lorica; foot short and coni-
cal; toes slightly decurved to slightly sigmoid, tapering to acute tips; mastax large
with two large salivary glands; trophus type A with sharply pointed unci and
rounded, oval rami which have curved alulae on their outer margins; manubria short
and outward curved. Gastric glands brownish to black; no eyespot.
Total length 110-200 !lm, toes 28-35 !lm, trophus 32 !lm.
Ecology: Europe, N. America, Australia(?) occasionally free-swimming, but more
commonly ectoparasitic on oligochaetes (Chaetogaster, Nais, Stylaria)
Literature: Kutikova 1970, Koste & Shiel 1991.
Code: 0103, 0201, 0301, 0404, 0510, 0802,0901,1002,1006,1101,1201,1301,-
1306,1402,1501,1606,1807,1902.
Description: Very transparent, slim, 10rica pliable but visible, lateral sulcus wide.
Laterally compressed, foot short, almost covered by tail. Corona almost straight, con-
vex, no lips. Toes very long, S-shaped, wide at the apical end, usually spread.
Footglands round, several salivary glands, vitellarium with eight nuclei. No eyespots.
Trophus type B: rami symmetrical, teeth on both sides, manubrium with double la-
mella, crotched; the pleural rod dissolves quickly in bleach. Very restless, rare, pos-
sibly an ecotype of C. incila with longer S-shaped toes but identical trophus.
Total length 180 !lm, toes 56 !lm.
Ecology: In moss moistened by a trickling spring. Gennany, Russia.
Literature: Kutikova 1970.
Code: 0102, 0201,0302,0407,0504,0701,0901, 1002, 1007, 1101, 1201, 1301,
1309,1402,1501,1507,1607,1702,1905.
Cephalodella paxilla Myers, 1924
Fig. 158
C. paxilla Myers, Harring & Myers 1924, p. 468-469, Fig. 26:6.
Type locality: NJ, U.S.A.
Holotype: Not designated
Description: Body elongate, slender, cylindric and very nearly parallel-sided. In-
tegument thin, flexible, plates indistinct, the lateral clefts flaring. Toes short and
nearly straight, ending in acute tips. Corona only slightly oblique, convex and with-
out projecting lips. Trophus type A, delicate; the fulcrum is a long, straight, very
slender rod, not expanded posteriorly, the manubria not crutched. Gastric glands
small, ganglion moderately long and saccate; no RCO, eyespot frontal and double,
the two spheres fairly wide apart.
Total length 210-220 !lm, toes 36-40 !lID.
Ecology: Among t10ating Sphagnum, rare.
Literature: Not seen since discovery.
Code: 0103, 0203,0205,0301,0902,1002,1003,1101,1201,1302, 1306,1402,
1501, 1607, 1803, 1903.
116 117

Description: Stout, round, strong lorica that is divided into five plates by strong
folds: two dorsal, two dorsolateral and one ventral plate (another species with five
plates is misgurnus). Head short, wide, dorsal edge strongly flared. Foot very large,
tail longer than foot with double keel, toes thick, tips separated by septa, straight of
faintly decurved, in dorsal view widely separated. Corona very oblique, moderately
convex, no lips. Ganglion long, double frontal eyespots with lens. Trophus type B,

,l
rami asymmetrical with teeth on both sides and alula on the left side, unci asymmetri-
cally thick and pointed; manubria crotched. Closely related to C. pach.-vdact.vla:
Wulfert suspected that pachydactyla is an ecotype of this species. We consider it to
be quite different and a valid species, based on the lorica, toes, rami and unci.

~w Total length 100-150 !-lm, toes 28-31 !-lm, trophus 25-31 !-lm.
Ecology: In a concentrated saltwater pool devoid of vegetation, pH 9. Also found in

~ ~~ shallow mountain pools, pH 5.3-6.0.

Literature: Godeanu 1964, Kutikova 1970.
Code: 102 0102, 0203, 0205, 0302, 0402, 0405, 0407, 0504, 0801, 0901, 1001,
158 159 160
1103, 1104, 1201, 1302, 1304, 1305, 1306, 140~ 1502, 1508, 1607, 1608, 1906.

Cephalodella pheloma Myers, 1924

Fig. 160

C. pheloma Myers, Harring & Myers 1924, p. 496-497, Fig. 32:3.

Type locality: Atlantic City, NJ, D.S.A.

Holotype: Not designated

Description: Body elongate, slender and nearly cylindric. Head unusually long and
extremely oblique; neck slightly constricted dorsoventrally and excessively com-
pressed transversely, its width being less than half the width of the head. Abdomen
8 fusiform; the' integument is so flexible that it can hardly be called a lorica, and the
plates are very obscure. Foot large and robust; the tail is fairly prominent. Toes short,
stout and decurved, with the ventral edges somewhat undulate; basal portion nearly
161 cylindric, and the posterior tapers gradually to acute tips. Footglands very long and
tubular. Corona excessively oblique and slightly convex without projecting lips.
Fig. 158. C. paxilla. Mastax large, trophus type A: fulcrum broad and very slightly expanded posterorly,
Fig. 159. C. pentaplax. manubria unusually long, nearly straight, rod-like and stout. Ganglion very long and
Fig. 160. C. pheloma. pyriform; eyespot and RCO absent. Related to C. megalocephala, but the footglands
Fig. 161. C. physalis CA): Harring & Myers; (B): Wulfert 1940, laterally, trophi, toes, and eyespot. and the very narrow neck distinguishes it.
(158,160: Harring & Myers 1924; 159: Wulfert 1943; 161: Harring & Myers 1924 and Wulfert 1940).
Total length 200 !-lm, toes 35 !-lm.
Ecology: In an acid pool among Sphagnum and Fontinalis; rare.
Literature: Kutikova 1970.
Cephalodella pentaplax Wulfert, 1943 Code: 0103, 0201, 0301, 0503, 0902, 1002, 1003, 1101, 1201, 1302, 1308, 1402,
Fig. 159 1501,1606, 1803, 1903.

C. pentaplax Wulfert 1943, p. 166-168, Fig. 2.

Type locality: Saltwater pool, Ciechocinek, Belarus (in 1943 Herrmansbad).

Holotype: Not designated
 118 'I 119
Cephalodella phy!wlis Myers, 1924
Fig. 161

C. physalis Myers, Barring & Myers 1924, p. 484-485, Figs. 29:3-5.

Type locality: Acid ponds near Atlantic City, NJ, D.S.A.

Holotype: Not designated

Description: Body very short, stout and gibbous dorsally; head very large and ob-
liquely truncate anteriorly; its length is less than the dorsoventral depth. Lorica quite
firm, lateral cleft wide. Foot very short and stout; tail small and knob-like and near the
posterior end of the foot. The toes are blade-shaped, decurved and have acute tips; the
dorsal edge is evenly curved and the ventral edge has a blunt cusp about one third of
its length from the base. Corona strongly oblique, small beak. The mastax is large and 163
the trophus type A: fulcrum slightly expanded posteriorly, manubria rodlike, not
crutched, strongly curved inside and dorsally; on the European specimens described
162 164
by Wulfert (1940), (but not by Hauer, 1935), the manubria have a large median loop.
Ganglion very large and saccate; the double eyespot is at the posterior end. No RCO.
Resembles mus, which is much smaller.
165
Total length 103-160 ~m, toes 35-40 ""m, trophus 45 ""m.
\I
Ecology: Acidophil, rare; North America, Europe, New Zealand. •

Literature: Barring & Myers 1924, Hauer 1935, Wulfert 1940, Russel 1960, --"-1 .
Kutikova 1970.
Code: 0102,0203,0204,0301,0501, (0507), 0510, 0802, 0901, 1001, 1006, 1007,
1102,12021301,1304,1308,1402,1501,1602,1803,1903.

Cephalodella planera Myers, 1940

Fig. 162

C. planera Myers 1940, p. 4, Fig. 2:4

Type locality: Long Pond, Monroe County, PA, D.S.A. 166 167
Holotype: Not designated
Fig. 162. C. planera.
Description: Body elongate, cylindric, head oblique, no lips. Lorica flexible, plates Fig. 163. C. plicata laterally, and optical cross section.
poorly defined. Foot short, obscurely two-jointed, toes very long, abruptly swollen at Fig. 164. C. poitera.
base. Trophus type B (not figured), no eyespot. Resembles C. tenuiseta and mis- Fig. 165. C. praelonga.
gurnus. Fig. 166. C. psammophila.
Fig. 167. C. pseudeva. (162: Myers 1940; 163: Harring & Myers 1924; 164,165: Myers 1934;
Total length 207 ""m, toes 70 ""m. 166: Koch-Althaus 1962; 167: BerziI{s 1976).
Ecology: Fairly common among Utricularia and Myriophyllum in summer. pH 6.6.
Literature: Not seen since discovery.
Code: 0101,0201,0302,0504,0802,0902,1002,1302,1303, 1306, 1501. Cephalodella plicata Myers, 1924
Fig. 163

C. plicata Myers, Harring & Myers 1924, p. 483, Figs. 28:3-4.

Type locality: WI and NJ, U.S.A.

 120
• Holotype: Not designated
Description: Body rather short, strongly gibbous dorsally. Lorica very firm and the
plates well marked; posteriorly it projects over the foot nearly to the base of the toes.
Dorsal and lateral clefts are abnormally deep and their edges project as very distinct
ridges, as shown in the optical section of the body in Fig. 163; dorsal cl~ft.is straight-
sided and acute-angled, the lateral clefts rounded at the bottom. Sy~blOtIC zoochlo-
rellae are invariably present and arranged in a fairly regular and qUIte constant ~at­
tern. Foot rather small; the minute tail is near the posterior end; toes rather short, faIrly
stout, tapering, and slightly decurved. Corona moderately obli~ue and convex,. no lips.
Trophus type A: fulcrum very slightly expanded at the postenor end; manubna sh~rt,
slender, without terminal crutch. No gastric glands, or RCO, eyespot at the postenor
end of the ganglion. Related to C. gracilis, hoodi, licinia.
Total length 105-110 /lm, toes 24-27 !lm.
Ecology: Acidophilic, not rare. N. America, Europe, New Zealand.
Literature: Russell 1960, Kutikova 1970.
Code: 0102, 0202, 0204, 0301, 0901,1001,1006,1101,1202,1301,1308,1402,
1501, 1605, 1607, 1901, 1903.
Cephalodella poitera Myers, 1934
Fig. 164
C. poitera, Myers 1934, p. 10-11, Fig. 11.
Type locality: Mount Desert Island, ME, D.S.A.
Holotype: Not designated
Description: Body short and stout, head large and oblique. Neck well marked, lorica
firm and well defined. Foot and tail short, toes blade-shaped, ventrally curved with
sharp tips. Corona very convex with beak. Trophus type B (no figure): fulcrum long
and abruptly spatulate, manubria expanded in the middle, and crutched. No eyespots.
Rapid and restless movement. Resembles C. globata, and physalis, but differs by the
absence of eyespot, trophus and the shape of the toe.
Total length 120 !lm, toes 18!lm.
Ecology: In a tide-plain stream, pH 6.0-6.4, occasionally abundant.
Literature: Kutikova 1970. Not seen since discovery ?
Code: 0103, 0201, 0302, 0504, 0901, 1003, 1101, 1201, 1302, 1401, 1501, 1602,
1607.
Cephalodella praelonga Myers, 1934
Fig. 165
C. praelonga, Myers 1934, p. 12-13, Fig. 12.
Type locality: Mount Desert Island, ME, D.S.A.
Holotype: Not designated
]2 J
Description: Body extremely elongate and slender, head relatively short; stiff lorica
which fonns distinct folds when the corona is retracted. Foot very short, almost cov-
ered by tail; toes long, slender, straight, the fine bristle-like tips abruptly curved
ventrally. Corona apparently never extends beyond the opening of the head-sheath,
not even in live animals. Mastax with a pair of salivary glands, trophus type A: ful-
crum straight and slightly expanded terminally, manubria very short, bacillar, slightly
decurved. No eyespot. Close to akrobeles, but has much longer toes.
Total length 157 !lm, toes 34 /lm.
Ecology: In acid pools among submerged Sphagnum.
Literature: Not seen since discovery.
Code: 0102, 0201, 0301, 0501, 0503, 0901, 1002, 1003, 1101, 1201, 1301, 1306,
1402, 1503, 1506, 1608, 1903.
Cephalodella psammophila Koch-Althaus, 1962
Fig. 166
C. psammophila, Koch-Althaus 1962, p. 76, Figs. 8a-f.
Type locality: Stechlinsee, Gennany.
Holotype: Not designated
Description: Body slender, gibbous, laterally compressed. Head well separated,
small lips, lorica overlaps foot, short tail. Foot short with caudal antenna and
long bristle, toes long, slightly S-shaped, tip a thin bristle. Trophus type A, ma-
nubria dorsally curved with median loop, fulcrum with basal apophysis. Rami with
median Hare but no alulae. Cervical eyespot. Close to subsecunda, but no hyaline
spot.
Total length 180-190 !lm, toes 68-72 !lm, trophus 31 /lm.
Ecology: Psammophilic, abundant at type locality, but not reported since.
Literature: Not seen since discovery.
Code: 1090101,0204,0301,0501,0507,0601,0802,0901,1002, 1007, 1301,1309,
1403, 1506, 1907.
Cephalodella pseudeva Berzivs, 1976
Fig. 167
C. pseudeva Berzivs 1976, p. 14, Figs. 34-36
Type locality: Lake Stnlken, Sweden.
Holotype: Not designated
Description: Body fairly slender, somewhat gibbous, flattened dorsoventrall~. Head
well separated, no lips. Foot stout, toes unique: thick apically, unifonnly tapenng, but
in the middle (viewed dorsally) they flare out and then run parallel agai~; thi~ ~rrange­
ment is rigid. No eyespots. Trophus type A (erroneously stated as B m ongmal de-
 122 123

scription): fulcrum thin, spatulate, rami simple, without teeth, manubria thin cres-
cents, slightly swollen distally. It differs from C. eva by the different toes which are, in
C. eva, very soft. It is also reminiscent of C. apocolea, but the vacuoles in the toes are
missing.
Total length 180-185 ~m, toes 38-40 ~m, trophus 26-29 ~tm.
Ecology: Found over several years in Lake Stnlken, a circumneutral softwater lake.
Literature: Not seen since discovery.
Code: 0102, 0201, 0301, 0502, 0510,0801,0901, 1002,1301, 1311, 1402,
1506.

Cephalodella reimanni Donner, 1950

Fig. 168 168
C. reimanni, Donner 1950, p. 309-311, Fig. 5.

Type locality: In a ditch, Czech Republic.

Holotype: Not designated

Description: Stout, gibbous, round, head large and oblique, lorica well visible; lateral
sulci flaring, ventral sulcus missing. Foot small, characteristically bulging ventrally,
almost covered by tail; toes sharp, ventrally decurved. Corona strongly convex, with
lips. Large mastax with salivary glands; trophus type B: fulcrum spatulate, rami asym-
metrical with a few teeth on the righthand side, none on the left; small alulae on both
sides. Manubria characteristically crutched, with double lamellae; one pleural rod.
Ganglion large with two subcerebral glands, double frontal eyespot with lens. Male
known. Related to C. sterea, globata and cyclops: cyclops has a symmetrical trophus
with very long alulae and two pleural rods; globata a U-shaped pleural rod and only
pseudoalulae on the rami. C. sterea is a very variable species, but is usually much
slimmer than the others in the group.
Total length 122-132 ~m, toes 23-31 ~m, trophus 25 ~m.
Ecology: In slighty alkaline water, ?urope;_ n~t _~~r~. ~ .(~. t id, f eco 1,o(S 'l.
Literature: Kutikova 1970. f:<.l10t t d ("'(,l.t"'';;3.;:>(iOl. .
Code: 0102, 0203, 0205, 0302, 0402, 0404, 0407, 0504, 0701, 0801, 0901, 1001,
1008, 1101, 1201, 1302, 1308, 1402, 1501, 1601, 1606, 1607.

Cephalodella rigida Donner, 1950

Fig. 169

C. rigida, Donner 1950, p. 317, Fig. 14.

Type locality: Tiny puddle, Czech Republic.

Holotype: Not designated
Description: Stout, prismatic, strongly compressed laterally. Head wide laterally as
well as dorsally, not oblique, dorsal edge bulging. Lateral sulci flaring, and continuing
into a fold on the head; ventral sulcus absent. Foot covered by tail, toes straight. Co-
~c
Fig. 168. C. reimanni laterally and dorsally, incus, male, manubrium, and fulcrum.
Fig. 169. C. rigida laterally, trophi, manubrium, three different form~ of rami. . .
Fig. 170. C. rostrum dorsally and laterally, trophi; (a): right manubnum; (b): left manubnum; (c): tip of
toe. (168: DOlmer 1950; 169: Donner 1950, 1964; 170: Koste 1978).
 124 125
rona convex with lips; ganglion long, tenninally a small colourless lens without pig-
ment. Trophus modified type B: fulcrum spatulate with apophysis, rami symmetrical
with strong teeth (sometimes reduced), manubria with lamellae, end fan-like; behind
the rami a branched sub-uncus that does not dissolve in bleach. V-shaped pleural rod.
Closely related to (perhaps identical with ?) C. tenuior; differences only in the straight
head, thin toes, lateral compression, ganglion-lens, and minor differences in the
trophus. However, Donner found and retained the species in later publications; so did
Koste (1978). Also related to limosa, which has no rami teeth (see there regarding the
affinities of this group).
Total length 132-175 !lm, toes 33-43 ~m, trophus 22 !lm.
Ecology: Small pools, mud, submerged mosses.
Literature: Donner 1964, 1970, Kutikova 1970, Koste 1984.
Code: 0102, 0201, 0208,0302,0407,0506,0601,0702,0801,0901, 1001, 1007,
1102, 1201, 1301, 1402, 1501, 1601, 1607. 173
171 Vir> _:,
'J

f ('Ho)
Cephalodella rostrum Reed, 1978
Fig. 170

C. rostrum Reed, Koste 1978, p. 345, Figs. 49a: 1a-g

Type locality: Thursley Common, Surrey, England.

Ho1otype: Not designated

Description: Described by Koste (1978) from personal communication of Reed. A

Body slim, slightly gibbous; head short, dorsal edge bulging. Lorica well defined,
sulci wide. Foot and tail short; toes almost straight, ending in dorsally bent claws or
tips. Corona narrow, oblique, convex, with a divergent rostrum originating in the
apical field and not part of the trophus, reminiscent of the rostrum of some
Dicranophorus species. No eyespots. Trophus type A: rami with alulae, manubria
short, asymmetrical, the left one much thicker, ending in an abrupt dorsal kink; the 172
right one is S-shaped in lateral view.
Total length 139 ~m, toes 42 ~m, trophus 23 ~m. Fig. 171. C. rotunda laterally, trophi, manubrium, toes laterally and dorsally.
Ecology: Acidophilic. Fig. 172. C. rotunda bryophila (A): after Wulfert (B): after Donner.
Literature: Not seen since discovery. Fig. 173. C. speciosa. (Fig. 171: Wulfert 1937; 172: Wulfert 1950, Donner 1970; 173: Harring & Myers
1924).
Code: 0102, 0201, 0301, 0404, 0509, 0901, 1002, 1005, 1101, 1201, 1301, 1306,
1402, 1501, 1602, 1903.
short, almost covered by foot, toes straight, with a strong "root". Caudal antenna
present. Corona oblique, convex. No eyespots, one crystal at the end of the ganglion
Cephalodella rotunda Wulfert, 1937 (see C. rigida). No RCO. Mastax with two salivary glands, trophus type B: rami with
Fig. 171 sharp apical teeth and strong curved alulae; manubria cTIltched, with one lamella.
Similar to C. tenuior and bryophila; these are difficult to differentiate, to the extent
C. rotunda Wulfert 1937, p. 613-614, Fig. 23. that Donner (1970) designated bryophila as a subspecies of rotunda, and Koste (1978)
went along with this idea. We retained bryophila as a subspecies, even though Donner
Type locality: Ditch near Merseburg, Gennany. (1978) suggested synonymization.
Holotype: Not designated Total length 124-140 !lm, toes 23-28 !lm, trophus 21-28 ~m.
Ecology: Has been found in both alkaline and acidic waters.
Description: Cylindrical, slim, head large. Lateral sulci flaring, head with folds. Foot Literature: Donner 1964, 1970, 1978, Kutikova 1970.
126
Code: 0103, 0201,0208,0302,0404,0504,0801,0901,1001,1101, 1201, 1301,
1306, 1402, 1501, 1606, 1607, 1803.
Cephalodella rotunda bryophila (Pawlovski), 1938
Fig. 172
Cephalodella bryophila Pawlowski 1938 (not seen)
C. rotunda bryophila Wulfert 1937 (Donner 1970)
Type locality: Tree-moss in Lituania.
Holotype: Not designated
Description: Body stout, widening posteriorly, ventrally somewhat concave, dorsal
line variable. Head stout, well separated, neck distinct, lorica soft, sulci wide. Foot
strongly tapered, no tail; toes uniformly tapered, slightly curved ventrally, rarely S-
shaped. Corona not oblique, convex, no lips. Stomach green, intestine yellow, gan-
glion very long, no eyes. Trophi type B, rami with long pointed alulae, manubria
crutched with basal lamellae. As to synonymy, both Donner (1978) and Koste (1978)
combine C. rotunda and C. bryophila.
Total length 122-128 ~m, toes 23-28 ~m.
Ecology: In moss and Sphagnum.
Literature: Wulfert 1937, 1950; Donner 1970,1978; Kutikova 1970, Koste 1978.
Code: 0103, 0201, 0302, 0404, 0504, 0801, 0901, 1001, 1201, 1306, 1402, 1501,
1607, 1904.
Keyed under both medium- and short-toed groups.
Cephalodella speciosa Myers, 1924
Fig. 173
C. speciosa Myers, Harring & Myers 1924, p. 504, Fig. 34:6.
Type locality: Oceanville, NJ, U.S.A.
Holotype: Not designated
Description: Body moderately elongate, conical; head very large; its dorsoventral
width is greater than the width of the abdomen. Toes very long, slender, tapering, very
slightly decurved ventrally; near the blunt tip is a transverse septum, giving the toes an
appearance of being clawed. Corona oblique, no lips. Ganglion large, saccate; no eye-
spot or ReO. Trophus large, type A: fulcrum crutched, manubria slender and rodlike.
Superficially resembles the nana, be/one group, but has no eyes and the toe shows a
septum.
Total length 145-155 ~m, toes 45-47 !-tm.
Ecology: Acidophil, among Sphagnum and Riccia.
Literature: Not seen since discovery.
Code: 0102, 0201, 0301, 0503, 0902, 1002, 1004, 1101, 1201, 1301, 1402, 1501,
1505, 1508, 1803, 1903.
127
Cephalodella stenroosi Wulfert, 1937
Fig. 174
C. stenroosi, Wulfert 1937,p. 624-625, Fig. 32.
C. bertonicensis Manfredi 1927
C. deformis Donner 1950
Type locality: On anaerobic sediment (sapropel) in alkaline ditches, Germany.
Holotype: Not designated
Description: Stout, powerful, cylindrical/prismatic, the trunk divided into a upper
larger and a lower segment. Lorica rigid, sulci narrow terminally, dorsal sulcus absent
(?). Head short, laterally compressed, foot short, conical, toes stout, thick decurved
dorsally with a dorsal lump or dull tooth in the middle; viewed dorsally, the toes touch
on the inner edge and are wide frontally, tapering to a common sharp tip. Footglands
large with reservoir, gastric glands quite large, ganglion long with subcerebral glands
but no sac; eyespots absent. Mastax round, without salivary glands; trophus type 0:
fulcrum long, spatulate, rami semicircular, opening carrot-shaped; subunci present,
quickly dissolved in bleach. Manubria with inward curved knob and double round la-
mellae, similar toforficula, which it resembles; however the lack of eyes is diagnostic.
Total length 190-240 ~m, toes 55-65 Ilm, trophus 30-34 Ilm.
Ecology: Benthic.
Literature: Wulfert 1937, Donner 1950, Kutikova 1970.
Code: 1180102,0201,0304,0503,0802,0902,1001,1003,1101, 1201, 1302,1307,
1312, 1402, 1501, 1603, 1605, 1607.
A subspecies, C. stenroosi austriaca Donner 1972 (Fig. 174 C) differs from the type
by the absence of a lump or tooth on the toe, which ends in a bristle. The animals are
also smaller: total length 159 !-tm, toes 40 !-tm; (this is C. stenroosi n.var. Donner in
Koste 1978; see also Donner 1978).
Cephalodella sterea sterea (Gosse), 1887
Fig. 175
Furcularia sterea Gosse 1887b, p. 864, Fig. 14:8.
Diaschiza sterea , Dixon-Nuttall & Freeman 1903, p.8, Fig.I:3
Type locality: Watcombe Park, Torquay, England.
Holotype: Not designated
Description: Body fusiform; head large; lorica firm, plates well mar~ed; foot large,
robust; tail extends beyond distal end of foot; toes short, stout slIghtly recurved
posteriorly, may have slightly undulate margins, variable; foot glands !arge and
pyriform; mastax large with four small salivary glands; trophus asymmetrIC type B:
manubria strongly cnltched; rami with variable inner margin denticulation; left ramus
with a small alula; three pleural rods. Retrocerebral sac present; two red frontal eye-
spots in single capsule.
128 129

Total length 140-250 !lm, toes 26-56 !lm, trophus 37-39 !lID.
Ecology: Cosmopolitan, sediment, moss, rivers.
Literature: Harring & Myers 1924, Wulfert 1937, Donner 1964, Kutikova 1970,
Koste & Shiel 1991.
Code: 0102, 0103, 0203, 0205, 0302, 0402,0405,0504,0701,0802,0902, 1001,
1103, 1201, 1302, 1306,1402,1501, 1605.

Variable species, forming a "species group". The following "forms" have been de-
scribed; their assigned subspecific status is uncertain and assumed, as all names point
to phenotypic differences:

Cs. dentata Donner, 1950 (Fig. 175 E). Dorsal plates and tail very long, both rami are
toothed.

Cs. exoculis Berzil}s, 1976, p.14. (Fig. 175 F). Eyes and alula on left ramus absent.
pH 5.7.
174
Cs. minor Donner, 1950, p.311-312. (Fig. 175 C). Small habitus ((105-122 !lm), both
rami carry alulae, the left one is larger.

Cs. mutata Donner, 1950, p.311. (Fig. 175 D). Both rami toothed, lamellae on
manubrium large, covering only half the shaft.

Cs. serrata (Wulfert), 1937, p. 609-610 as C serrata (Fig. 175 G). Tail very long,
unci, manubria and pleural rods asymmetrical, rami with serrated inner edge; two
retrocerebral sacs, brown gastric glands.

Cephalodella subsecunda Myers, 1942

Fig. 176

C subsecunda Myers 1942, p. 276-277, Fig. 23:1.

Type locality: Pocono Lake, PA, D.S.A.

Holotype: Philadelphia Academy of Natural Sciences 665

175 B Related to tantilla, but the toes are different.
Fig. 174. C. stenroosi (A): Wu1fert's original figure, toe after Stenroos; (B): Donner's C. deformis; (C): C.
stenroosi austriaca laterally, toes dorsally and laterally, trophus, tips of rami.
Fig. 175. C. sterea Group A: C. s. sterea (a,b): slim form; (c): stout form; (d): aberrant toes; (e): trophus;
(t): fulcrum and ramus laterally; (g): left ramus; (h,i): manubrium from different sides. Group B: male,
copulation. Group C: C. s. "form" minor laterally, different toe, rami dorsally and laterally, trophus later-
ally. (174: A: Wu1fert 1937, B: Donner 1950, C: Donner 1972, Koste 1978; 175: A,B: Wu1fert 1937, C:
Donner 1950).
Description: Body stout, cylindric/prismatic, head large, neck sharply offset, dorsally
arched; a round hyaline area at the position of the lateral antennae on both sides.
Lorica firm, sulci flaring posteriorly. Foot short, conical, toes long, S-shaped uni-
formly tapering. Corona oblique and very convex, no lips. Ganglion long, one cervical
eyespot. Trophus type A, manubria slender rods curving dorsally in a semi-circle.
Total length 90 !lm, toes 40 !lm.
Ecology: Acidopilic in Sphagnum subsecundum.
Literature: Not seen since discovery.
Code:120 0101, 0202,0204,0301,0501,0503,0901,1001, 1006, 1101, 1201, 1301,
1309, 1402, 1501, 1606, 1607, 1903.
130 131

N.B. The hyaline areas are unique; only Monommala grandis and A1. actices have such spots at the identi-
callocation, but those spots are bright red.

Cephalodella tachyphora Myers, 1934

Fig. 177

C. tachyphora Myers 1934, p. 3-4, Fig. 2.

Type locality: Mount Desert Island, ME, D.S.A.

Holotype: Not designated

o E
Description: Body short and stout; head large, somewhat deflexed, neck constriction
visible; lorica finn. Foot and tail short, toes very long, separated at base, ventrally
bent. Corona convex, very prominent beak. Ganglion long, cervical eyespot.
Trophus type A: fulcrum long, abruptly expanded tem1inally, manubria slender rods
curving upwards in a semicircle; epipharynx slender with sharp tip. Resembles
tantilla, but the beak and ventrally curved toes are different.
Total length 95-130 ~m, toes 40-45 ~m.
Ecology: Predator, acidophilic, abundant among Sphagnum in New Jersey.
Literature: Koch-Althaus '1963, Kutikova 1970.
Code:121 0101,0202,0204,0301,0501,0503,0901,1001,1101, 1201, 1301, 1308,
1402, 1501,1602, 1605, 1903.

F Cephalodella tanti/la Myers, 1924

175 Fig. 178

C. tantilla Myers, Harring & Myers 1924, p. 486-487, Fig. 3Q:2.

)/ C. galbina Myers, Harring & Mye~s 1924, p. 490, Fig. 31: 1. IA.() f j dls.J/v101 .r(?C:Ctr"j
c. sabulosa Myers 1942, p. 276, FIg. 23:2 -~ (0,,('( _ 10 Aj.J JP f-"t.... 'JIe-{I·~.f/
~ [;jAJ}>1 't
Type locality: No single record; WI, NJ, FL, D.S.A.
Holotype: Not designated

Description: Body moderately elongate, laterally compressed and strongly gibbous

dorsally; lorica finn, plates well marked, lateral clefts flaring. Foot short and broad;
the fairly prominent tail is a little beyond mid-length. Toes wide apart at the base,
long, slender and recurved. Corona oblique and strongly convex without lips. T:ophus
type A: fulcrum very stout and slightly expanded at the extreme end, manubna very
slender, rod-shaped and recurved, but not crutched. Ganglion long and saccat~; no
RCO. The eyespot is at the posterior end of the ganglion. ~losely. resembles C. gl~ba,
but the smaller size, cervical eyespot and trophus are dIagnostIc. C. sabulosa IS a
newly designated synonym (perhaps an ecotype).
176 177 178 179 Total length 115-120 ~m, toes 38-40 ~m.
Fig. 175. (cont.) C. sterea Group D: C. s. "form" mutata laterally and dorsally, rami and furcrum dorsally, Ecology: Acidophil, cosmopolitan.
trophus laterally, uncus. Group E: C. s. "form" dentata laterally, rami and furcrum dorsally, manubrium Literature: Kutikova 1970, Koste 1978.
laterally, uncus. Group F: C. s. "form" exoculis trophus. Group G: C. s. "form" serrata.
Fig. 176. C. subsecunda.
Code: 0101, 0202, 0204, 0301, 0501, 0503, 0901, 1003, 1006, 1101, 1201, 1301,
Fig. 177. C. tachyphora. 1307, 1402, 1501, 1606, 1607, 1803, 1903.
Fig. 178. C. tantilla, with optical cross section.
Fig. 179. C. tantilloides. Scale bars 50 and 10 !Lm. (175: D,E: Donner 1950, F: B-erzinGs 1976, G: Wulfert
1937; 176: Myers 1942; 177: Myers 1934; 176,177,178: Harring & Myers 1924, Myers 1942; 179: Hauer
1935).
 132
133
Cephalodella tantilloides Hauer, 1935
Fig. 179

C. tantilloides Hauer 1935, p. 69-70, Fig. 9.

Type locality: Schluchsee bog, Black Forest, Germany.

Holotype: Not designated

Description: Body like C. gibba and tantilla; very gibbous, laterally strongly com-
pressed, sulci well visible, abdomen curves sharply to clearly demarcated foot; dorsal
sulcus relatively deep. Toes very long, dorsally curved to a distinctly segmented tip.
Corona oblique but not convex, prominent lips. Trophus type A: fulcrum dilated
distally into a broad plate; rodlike manubria curve upwards; rami with inner denticles.
Cerebral eye present; retrocerebral sac not recorded.
Total length 160-180 !-lm, toes 45-56 !-lm, toe tips 14 !-lm, trophus 34 !-lm. (N.B. The
total length given as 104 !-lm by Hauer is an obvious error, considering the toe length, 180 181
as pointed out by Wulfert (1950).
Ecology: Acidophilic, cosmopolitan.
Literature: Wulfert 1950, Kutikova 1970, Green 1981, Koste & Shiel 1991.
Code: 0102,0202,0204,0301,0407,0501,0503,0802,0901, 1001, 1006 1101,
1201, 1301, 1305, 1307, 1508, 1601, 1605, 1803, 1903.

Cephalodella tecta Donner, 1950

Fig. 180

C. tecta Donner 1950, p. 318-319, Fig. 16.

Type locality: Sediment of River Thaya, Slovakia. 8

Holotype: Not designated
Description: Body prismatic, slender, laterally compressed; transparent, but lorica
well developed. Foot large, well separated, covered by tail. Toes straight, separated,
tips abruptly tapered (with septum? claw? figures not clear). Corona convex, small
lips. Mastax small, salivary glands large. Double frontal eyespots. Trophus type C:
rami with fine teeth, manubrium with terminal loop that is not entirely closed, also
with wide lamellae. Y-shaped pleural rod that dissolves in bleach. Related to obvia ?
Total length 155-168 !-lm, toes 36 !-lm, trophus 19 !-lm.
Ecology: Not rare, in sediment.
Literature: Kutikova 1970.
Code: 0102, 0203, 0205, 0303, 0407, 0508, 0702, 0801, 0901, 1002, 1102, 1201,
1301, 1305, 1306, 1402,1502, 1505, 1601, 1607.
Cephalodella tempesta Wulfert, 1956
Fig. 181
C. tempesta Wu1fert 1956, p. 467, Fig. 9.
A
182
Fig. 180. C. tecta laterally, dorsally, trophus, uncus.
Fig. 181. C. tempesta.
Fig. 182. C. tenuior Group A: after Donner, laterally, rami and fulcrum with pleural rod. Group B: after
Wulfert. Group C: after Koste & Polz. K: crystal on central ganglion, Mei: reddish- brown pigmentation
on oesophagus and stomach, PI: pleural rod, Kr: crutch on manubrium. Toes ventrally. (180: Donner 1950;
181: Wulfert 1956; 182: A: Donner 1950, B: Wulfert 1956, C: Koste & Poltz 1984).
Type locality: Teufelssee, Friedrichshagen, Germany.
Holotype: Not designated
Description: Transparent, narrow, slender, laterally compressed, body cylindrical;
head large and characteristically tilted at a 45° angle. Sulcus narrow, foot and tail
short, toes straight, uniformly tapering. Corona "nose-like", very convex. Two very
134
small separated frontal eyespots. Mastax with two salivary glands, trophus type B:
fulcrum thin , manubria crutched, two pleural rods. Resembles C. montana which,
however, is eyeless and does not have the tilted head.
Total length 185 ~tm, toes 25-30 ~lm, trophus 25-28 ~lm.
Ecology: Very rare, benthic, solitary.
Literature: Kutikova 1970.
Code: 0103, 0203, 0205,0302,0504,0701,0801,0902,1003,1007,1101,1201,
1302, 1306, 1402, 1501, 1606, 1608.
Cephalodella tenuior (Gosse), 1886
Fig. 182
Diaschiza tenuior, Hudson & Gosse 1886, p. 2:81, Fig. 22:14.
C. tenuior, Harring & Myers 1924, p. 497, Fig. 33:3.
Type locality: Woolston Pond, Dundee, U.K.
Holotype: Not designated
Description: Body elongate, slender and nearly cylindric. Lorica very flexible, plates
indistinct; the lateral clefts are narrow for about two thirds of their length; at this point
the dorsal plates become rounded and the cleft widens rapidly. Foot relatively short
and broadly conical; the tail covers the foot only partly. Toes short, straight or slightly
decurved dorsally, with septum, sometimes with a "vacuole". Corona oblique,
strongly convex, no lips. Trophus type B: fulcrum slightly expanded posteriorly and
the manubria crutched; U-shaped pleural rod. Gastric glands small and occasionally
have a brownish tint (Fig. 182 C). Ganglion very long and pyriform, with one or two
colourless crystals terminally (Fig. 182 C); no eyespot or RCO.
As mentioned under C. limosa, the group containing tenuior, clara, limosa, rigida,
forficata, intuta and incila is difficult to differentiate, even though they all seem to be
good species. Using numerical techniques, intuta proved to be closest to tenuior, but
can be differentiated by ventrally and inward curved toes. Only biochemical and ge-
netic investigations will discriminiate between these species and clarify the problems.
Wulfert (1950) has described a "variety" C. t. pigmentata, differentiated by pres-
ence of red ''jumping, skipping" spheres (hiipfende Kugeln) in the enlarged oesopha-
gus, no lenses in the ganglion, the tips of the toes are more separated and two straight
and one Y-shaped pleural rods are present.
Total length, 115-148 ~m, toes 22-25 ~m, trophus 20-21 ~m, male 80-90 ~m (?).
Ecology: Cosmopolitan but not common; in submerged mosses, psammon.
Literature: Harring & Myers 1924, Wulfert 1956, Donner 1970, Kutikova 1970,
Koste 1984, Koste~&ie1991
Code: 0103,(~{f2 . /~ 0302, 0407, 0504, 0702, 0801,__~01, 1002, 1101,
V'I·'1
7,
1201, 1302, 13U 1 6,1 4021"1501, 1508, 1606, 1607, 1803~
I , eye CjJI.
,
,3fAs\:,r\(jta vl{)G r~lIriJWf1
Cl e I U'3fllll{j'li?~S ~ .,
(jilSt;w1 Q f:..y€ :- -':' IJCCWJI)vt
, JJtJ(/JIJ/ ,S~.
(~.g~ ft. (~1{()
135
Cephalodella fermis Koch-Althaus, 1962
Fig. 183
C. tenuis Koch-Althaus 1962, p. 73-74, Fig. 6.
Type locality: Stechlinsee, Germany.
Holotype: Not designated
Description: Cylindrical, gibbous, dorsal lorica plates extend beyond foot and toes,
so that they are invisible when viewed dorsally. Foot small, hemispherical, toes small,
curved dorsally. Gastric glands connected to ganglion, red cervical eyespot with lens.
Trophus type A, not described but figure supplied. It seems that rami have a single
median tooth and a curved "supraramus" enclosing a circular space. Manubria
dorsally curved, unci simple, fulcrum terminally notched. The habitus of the animal
is, however, most characteristic. Found twice.
Total length 110-130 ~m, toes 10-11 ~m, trophus 27 ~m.
Ecology: Among Chara, June
Literature: Not seen since discovery.
Code: 0103, 0204,0301,0401,0501,0802,0901,1001,1103,1202, 1101, 1401,
1503, 1606, 1808.
C. tenuiseta (Burn), 1890
Fig. 184
Furcularia tenuiseta Bum 1890, p. 34, text figure.
Diaschiza tenuiseta, Dixon-Nuttall & Freeman 1903, p. 138, Fig. 1:2.
C. tenuiseta, Harring & Myers 1924, p. 508, Fig. 35:7
Type locality : England.
Holotype: Not designated
Description: Body elongate, fairly slender and nearly cylindric (Harring & Myers
1924), but slim, prismatic, straight-backed and laterally compressed in the European
ecotype. Head large, but relatively short, obliquely truncate anteriorly. Abdomen unu-
sually long and slightly gibbous posteriorly. Foot short, bluntly conical with a small,
rounded tail. Toes half as long as the body, very slender, slightly recurved dorsally.
Corona slightly oblique without projecting lips. Trophus type D: fulcrum slightly ex-
panded posteriorly, manubria rodlike, not crotched. No RCO and eyespot.
Total length 380-390 ~m, (Harring & Myers 1924), 350-475 ~m, (Wulfert 1937),
205-314 ~m, (Donner 1950), toes 120-125 ~m, (Harring & Myers 1924), 96-110 ~m,
(Wulfert 1937), 59-96 ~m, (Donner 1950), trophus 35-39 ~m.
In a detailed analysis Donner (1950) concluded that there are two ecotypes: those
of Bum (1890) and Dixon-Nuttall (1903) (both published without measurements) on
'. f()
the one hand, and the type of Rousselet, Harring & Myers (1924) (Fig. 184 A),
i waft, Wulfert (1937) (Fig. 184 B), and Donner (1950)on the other hand. Harring & Myers
*&/ . found their specimens in brackish and saltwater ditches around Atlantic City, New
Jersey, D.S.A.; all the other specimens came from freshwater. Donner (1950) de-
 136
137

scribed his animals found in Bohemia as a subspecies, C. t. americana (Fig. 184 C).
This is distinguished by a smaller size but more robust build, shorter foot, much wider
toes; right ramus carries a striped lamella and a tooth, the left one two unequal teeth;
the manubria bend dorsally instead of ventrally. Koste (1978) synonymizes this sub-
species; we think it should be maintained as an ecotype, i.e., a subspecies until further
study. Berzilfs (1976) describes another subspecies, C. t. simplex (Fig. 184 D) from
Lake Lygnen at Aneboda, Sweden. The toes are very thin even at the base (as in C.
gigantea) , and the trophus much more simple; also, the fulcrum is shorter than the
manubria. Measurements agree with those published by Donner (1950).
Ecology: Benthic, rare, Europe, North America, Australia.
Literature: see text, above, Kutikova 1970, Koste & Shiel 1991.
Code: 0102, 0201, 0304, 0407, 0502, 0503, 0802, 0605, 0902, 0903, 1002, 1101,
a I 1201, 1301, 1307, 1301, 1403, 1501, 1605.

Cephalodella theodora Koch-Althaus, 1961

Fig. 185
c
C. theodora Koch-Althaus 1961, p. 219, Figs. la-g.
183 Type locality: Stechlinsee, Germany.
Holotype: Not designated

Description: Short, stout, cylindrical, strongly gibbous; head large. Foot very short,
displaced ventrally, covered by tail. Toes thick, blunt, curved ventrally; tip can have a
septum. Corona slightly oblique, convex, with lips. One cervical eyespot. Trophus
type Band D, or only B: rami symmetrical with eight and ten blunt teeth (or a fan of
teeth under the rami ?); fulcrum clubbed, manubria with median and terminal loop,
bent ventrally; on the Canadian specimens these loops were absent. Related to C.
eatellina, erassipes, xeniea (crutched manubria) and ventripes.
Total length 165-200 ~m, toes 25-33 !-Lm, trophus 27-32 ~m.
Ecology: pH 7.8-9.0, 18-20°C.
Literature: Kutikova 1970, Nogrady 1988.
Code: 0103, 0202, 0204, 0302, 0407, 0502, 0507, 0508, 0801, 0901, 1001, 1005,

~
::\ .,:
.. 1008, 1102, 1202, 1312, 1308, 1310, 1402, 1507, 1508, 1601.
.

Cephalodella tinea Wulfert, 1937

Fig. 186
B
C. tinea Wulfert 1937, p. 622-624, Fig. 31.
184
Fig. 183. C. tenuis two different animals laterally and dorsally, trophi. Type locality: Saprobic drain, Germany.
!
Fig. 84. C. tenuiseta Group A: after Barring & Myers, Group B: after Wu1fert Group C: C. t. Holotype: Not designated
amerIcana. (183: Koch-Althaus 1962; 184: Barring & Myers 1924, Wu1fert 1937, 184: A: Barring &
Myers 1924, B: Wu1fert 1937, C: Donner 1950). Description: Bodly elongate, laterally compressed; abdomen slightly bulbous. Tail
distinct, extending beyond foot; plates and sulci distinct; toes somewhat short, ca.
one-third of total body length, thickened at base, with slight median swellings; mastax
138 139

with two large salivary glands; trophus type D: fulcrum spatulate, proximally broad-
bladed (when seen laterally); manubria with blunt, slightly enlarged tips; paired fron-
tal eyespots in single capsule. Donner (1950) described a "variety", C. t. canspicua.
that differs from the nominate species by thinner and longer toes that are usually
crossed, asymmetrical rami, and branched subunci. These differences may not justify
a new taxon.
Total length 260-280 [.tm, toes 52-70 ~lm, trophus 29-31; male 160 ~lm; parthenoge-
netic egg 60 [.tm.
Ecology: Saprobic species in drains, manure, decomposing vegetation. Cosmopo-
litan?
Literature: Kutikova 1970, Koste 1976 Koste & Shiel 1991.
Code: 0102, 0203,0205,0304,0407,0503,0601,0801,0902, 1003, 1103, 1201,
1301,1303,1307, 1402, 1403, 1501, 1601, 1605, 1607.
1840
Cephalodella tineafarmis Koste & B6ttger, 1992
Fig. 187

C. tineafarmis Koste 1992, p. 286-287, Fig. 6.

Type locality: Lake Laguna de Colta, Ecuador.

Holotype: Not designated

Description: Similar to C. tinea, but longer toes; rami with double tips, without inner
teeth; manubria terminally flared and curved inward.
A Total length 226-240 [.tm,. toes 55-60 ~m, trophus 29-30 ~m.
'''W't" 185 Ecology: Found in a shallow hypereutrophic littoral; 19°C, pH 9.0, 1222 ~S cm-I,
sediment smelled ofB2S.
Literature: Not seen since discovery.
Code: 0102, 0203, 0205, 0304, 0406, 0511, 0601, 0801, 0902, 1009, 1101, 1201,
1301, 1307, 1402, 1501, 1605, 1607.

Cephalodella ventripes (Dixon-Nuttall), 1901

Fig. 188

Diasehiza ventripes Dixon-Nuttall 1901, p. 25, Figs. 2:1-3.

C. ventripes, Barring & Myers 1924, p.484, Fig. 28:5.
"186
Type locality: Knowsley Park, Lancashire, England.
Holotype: Not designated

Description: Body short, stocky, gibbous dorsally, oval in back view; dorsal lorica
may extend beyond distal end of the foot; plates and sulci distinct; dorsal sulcus a dis-
Fig. 184 (cont.). Group D: C. t. simplex laterally, trophus, toes, uncus. . tinct V-shaped groove. Foot ventral, small; toes short and stout decurved ventrally;
Fig. 185. C. theodora Group A: after Nogrady (a): laterally; (b): dorsally; (c): trophus; (d): incus pressed;
(e): trophus laterally; Group B: laterally, trophus laterally, pressed, dorsally, uncus; (a): toe of C.
corona with prominent lips; mastax large, trophus type A: fulcrum spatulate, distal
theodora; (b); toe of C. crassipes. ends of manubria sickle shaped, not crutched, also bent dorsally. Double cervical eye,
Fig. 186. C. tinea. (184: D: Berzil}.s 1976; 185: A: Nogrady 1988, B: Koch-Althaus 1961; 186: Wulfert no ReO. Close to aurieulata, which has a single cerebral eye and foot in body axis
1937).
140 14\

(N.B. cf. with C. v. angustior, below).

Total length 135-140 f.lm, toes 25-28 f.lm, trophus 30-34 f.lm.
Donner (1950) described a Cv. "var." angustior, that he equates with Wuffert's
animal marked with a question mark (Fig. 188 B), and this "variety" corresponds
more to the animal described by Harring & Myers (1924) (Fig. 188 A) than to Dixon-
Nuttall's and Wulfert's (1938a, Fig. 17) figures, referred to as the "true" C. v ventripes
that Wulfert (1937) also observed (Fig. 188 C). c.v. angustior has a foot that is not
ventrally displaced, has a single cerebral eyespot and a median loop on the manubria.
Ecology: Cosmopolitan, common. 13-21°C, pH 4.8-7.1,18-351 f.lS.
Literature: Harring & Myers 1924, Wulfert 1937, Donner 1950, Kutikova 1970, de
Smet 1988, Koste & Shiel 1991.
Code: 0103, 0203,0204,0301,0501,0510,0801,0901, 1001,1006,1103,1202,
1301, 1308, 1402, 1501, 1601, 1803.

CephalodeUa viteUa Wulfert, 1956

Fig. 189
187 189
C. vitella Wulfert 1956, p. 467-468, Fig. 11.

Type locality: Teufelssee, Friedrichshagen, Germany.

Holotype: Not designated

Description: Colourless, stout, cylindrical, gibbous. Head large, oblique. Foot strong,
covered by tail; toes straight. Corona semicircular with lips. Ganglion long; faint but
large cervical eyespot. Trophus type B, like those of C. tenuior: rami with tilted termi-
nal teeth, manubria crotched with a single lamella; U-shaped pleural rod that dissolves
in bleach. Related to limosa and tenuior, except that it has an eyespot. B
Total length 135-140 f.lm, toes 32 f.lm, trophus 21 ~lm.
Ecology: Acidophilic (pH 4.5-5.0).
Literature: Kutikova 1970.
Code: 0102,0202,0204, 0302, 0407, 0504,0702, 0801, 0901, 1001, 1005, 1102,
1201, 1301, 1306, 1402, 1501, 1601, 1607, 1903.

Cephalodella vittata Kutikova, 1985

Fig. 190

C. vittata Kutikova 1985, p. 55-56, Fig. 1.

190
Type locality: Lake Baikal.
Holotype: Not designated 188
Description: (Prepared from figure, not from text). Robust, cylindrical animal with Fig. 187. C. tincaformis.
Fig. 188. C. ventripes A: Harring & Myers, Group B: Donner's c.v. angustior, corresponding to Wulfert's
straight ventral lorica, slightly gibbous. Lorica distinct, lateral sulcus narrow. Head "Dixon-Nuttall & Freeman ?" form. Group C: Wu1fert's "true" Dixon-Nuttall fonn.
large, not oblique, strongly convex, no lips. Foot small, tail wide but does not cover Fig. 189. C. vitella.
foot; toes uniformly tapered, decurved ventrally. Gastric glands very large, large Fig. 190. C. vittata. (I87: Koste & Bottger 1992; 188 A: Barring & Myers 1924, B,C; Wu1fert 1937; 189:
Wu1fert 1956; 190: Kutikova 1985).
142 143

round cervical eye. Trophus simple, type A; fulcrum spatulate, unci with large alulae
(?), manubria almost straight, rod-like. Resembles C. megalotrocha, and dorystoma.
Totallenth 205-225 !lm, toes 30 /lm, trophus 35 ~tm.
Ecology: Littoral of Lake Baikal, 6°C.
Literature: Not seen since discovery.
Code: 0103, 0202, 0204, 0301, 0503, 0802, 0902, 1001, 1003, 1101, 1201, 1301,
(f)
~"
1308,1402,1501,1607.

Cephalodella wrighti Wulfert, 1960

Fig. 191
192

C. wrighti Wulfert 1960b, p. 318-319, Fig. 40. 191

Type locality: Krebsscheren Pond, Diiben moor, Germany.
Holotype: Not designated

Description: Hyaline, straight cylindrical body, head slightly bent, edge puffed.
Lorica distinct. Foot, short, conical, tail does not cover foot; toes short, straight. Co-
rona very convex, no lips but epipharynx visible. No eyespot. Trophus type B: ful-
crum very long, spatulate, manubria crutched, Y-shaped pleural rod. Related to C.
tenuior.
Total length 125-130 !lm, toes 18 !lm.
Ecology: pH 6-7, not rare at type locality.
Literature: Kutikova 1970.
Code: 0103, 0201, 0302, 0504, 0702, 0901, 1002, 1003, 1101, 1201, 1301, 1306, 193
1401,1501,1606, 1608.
194 195
Cephalodella xenica Myers, 1924
Fig. 192

C. xenica Myers, Harring & Myers 1924, p. 492-493, Fig. 29:6.

Type locality: Huron River, Ann Arbor, MI, D.S.A.

Holotype: Not designated

Description: Body moderately elongate, arched dorsally, its greatest depth near mid-
length. Foot large and robust; the prominent tail does not cover the foot; toes short,
blade-shaped and very broad; very slender, conical tips, slightly blunted at the ex-
treme ends (resembling those of C. crassipes and C. theodora). Corona oblique and 197 199
strongly convex without projecting lips. Trophus type B: fulcrum very slightly ex- 196 198
panded at the posterior end; manubria long with very pronounced tenninal crutch.
Gastric glands small and oval; ganglion elongate and pyriform; no RCO. Eyespot at Fig. 191. C. wrighti. Fig. 192. C. xenica. Fig. 193. C. zeteta. Fig. 194. C. eunoma. Fig. 195. C. conjuncta.
the posterior end of the ganglion. Fig. 196. C. derbyi. Fig. 197. C. retusa. Fig. 198. C. unguitata. Fig. 199. C. vacuna. (191: Wulfert 1960b;
192: Harring & Myers 1924, Hauer 1959; 193: Wulfert 1961; 194,195: Myers 1940; 196: Wulfert 1950,
Total length 128 !lm, toes 22 !lm.
1961a; 197, 199: Harring & Myers 1924; 198: Hauer 1935).
There is some confusion in the literature: Koste (1978) synonylnizes this species
with C. crassipes, but provides a xenica figure; he shows C. donneri Hauer (1959)
 144 145
(that is a synonym of crassipes), which has crutched manubria and is eyeless (like Cephalodella derbyi (Dixon-Nuttall), 1903
eupoda). We consider xenica a valid species. . c
Fig. 196
Ecology: Free-living (not parasitic as crassipes), not rare. -!((lU<'\Se.e '~9.1:1fl
Literature: Hauer 1959. Diaschiza derbyi Dixon-Nuttall and Freeman 1903, p. 131, Fig.4: 13.
Code: 0103,0202, 0203,0302,0404, 0504, 0901, 1001, 1005, 1302, 1402, 1502,
1507, 1803. Type locality: Unknown.
Holotype: Not designated
Cephalodella zeteta Wulfert, 1961 Description: Body strongly gibbous, laterally compressed; head large, neck very dis-
Fig. 193 tinct, deeply cleaved. Lorica distinct, sulcus flaring distally. Foot short, half-covered
by short tail. Toes thin, soft, staight or slightly curved dorsally, spread. Vitellarium
C. zeteta Wu1fert 1961, p. 83~84, Fig. 8.
with six nuclei; stomach green or gold; intestine well separated. Corona somewhat
oblique, not very concave, prominent lips. Ganglion small; cervical eyespot. Mastax
Type locality: Diiben moor, Germany.
large, salivary glands present. Trophus type A: fulcrum wide, strong, manubria
Holotype: Not designated
semicircu1arly bent in second half (no drawing of trophus in literature). Acidophil,
slow. Wulfert (1940, p. 564), discussing C. euderbyi, designates C. derhyi a very vari-
Description: Body short, stout, bulbous, very wide dorsally. Foot and tail short, toes
able and even doubtful species, quoting Myers' doubts as well. Needs more study.
fairly long, decurved ventrally or straight. Lorica soft, lateral sulci wide. Corona
Total length 110-165 flm, toes 34-42 flm, trophus 25-35 /lm.
slightly oblique and not very convex, with lips. Ganglion very long, with one eyespot
Ecology: Bog-ponds and Sphagnum.
on the dorsal side of the ganglion, and sometimes one additional eyespot on the ven-
tral side, unique. Trophus type A, manubria very short, inward curved. Related to C. Literature: Wulfert 1940, 1950, 1960a; Kutikova 1970.
labiosa and tachyphora, also similar to ventripes, derbyi. Code: 0102,0204,0301,0510,0802,0901,1001,1101,1201, 1301,1307,1402,
Total length 129-140 flm, width 70 !-tm, toes 38-40 /lm, trophus 25 /lID. 1501,1508,1601,1605,1701,1903.
Ecology: pH 6.0.
Literature: Kutikova 1970.
Code: 0102, 0202, 0203, 0204, 0301, 0503, 0801, 0901, 1001, 1008, 1101, 1201, Cephalodella eunoma Myers, 1940
1301, 1308, 1402, 1501,1601, 1605, 1903. Fig. 194

C. eunoma Myers 1940, p. 3, Fig. 1: 1

SPECIES INQUIRENDAE
Type location: Pocono Lake, PA, U.S.A.
Cephalodella conjuncta Myers, 1940 Holotype: Philadelphia Acad. Nat. Sci. No. 130 sf. curved wVl1~af- rl/(JJ[J f:X)
Fig. 195 /

C. conjuncta Myers 1940, p. 3-4, Fig. 2:3
Type locality: Pocono and Naomi lakes, PA, U.S.A.
Holotype: Not designated
Description: Body cylindric, robust, head large, corona oblique, no lips. No lorica
visible. Gastric glands small, eyespot absent. Mastax modified virgate (type not men-
tioned), manubria with well developed basal plate (not figured). Resembles C.
forficula, but differs by its larger size, absence of eyespot, prominent tail, stouter toes
without spines on their dorsal margin. Conspicuous by its milky appearance in the liv-
ing state. Only a habitus figure supplied, description too short.
Total length 243 !-tm, toes 47 flm..
Ecology: Among submerged plants, fairly common in summer at type location.
Literature: Not seen since discovery.
NOT IN KEY
Description: Body elongate, fusiform, head small, corona oblique, convex, no lips.
Abdomen gradually diminishing to short foot; toes short, straight, somewhat undulate
with acute recurving tips. Lateral plates indefinite. (Trophus neither descibed nor fig-
ured). Resembles C. dorystoma, but differs by the absence of a refringent vacuole
simulating an eyespot, different toes and recurved manubria.
Total length 195 flm, toes 30 flm.
Ecology: Rare, among Nitella in summer.
Literature: Not seen since discovery.
NOT IN KEY
Cephalodella retusa Myers, 1924
Fig. 197
C. retusa Myers, Ran-ing & Myers 1924, p. 498, Fig. 33 :6.
 146 147

Type locaIit)': Absecon, NJ, U.S.A. Description and figure insufficient for identification. Considered nomen nudum.
Holotype: Not designated

Description: Body rather short, fusiform, fairly stout. Head large and obliquely trun- Genus Paracephalodella Berzins 1976
cate anteriorly. Foot moderately large and robust; the tail is very small. Toes short and
stout with an acute tip. Corona strongly oblique and convex without projecting lips. Monotype: Paracephalodella latffztlcrum.
Trophus type B: the fulcrum is expanded posteriorly and the manubria crutched (no Type locality: Lake Frejen, Aneboda, Sweden.
figure). The ganglion is very large and saccate; eyespot and RCO absent. Related to C.
forficata, but much smaller; toes are different. The description and figure do not seem Description: Distinguished from Cephalodella by two movable palps on the trochal
to make the species identifiable. discs. The palps are not ciliated. The foot and toes are also some\vhat different from
Total length 100 !-tm, toes 24 ~tm. those seen in Cephalodella.
Ecology: Acidophilic, rare.
Literature: Not seen since discovery.
Code: 0102, 0201, 0302, 0504, 0901, 1001, 1101, 1201, 1302, 1402, 1501, 1606,
1607, 1803, 1903. Paracephalodella latifulcrum BerziQ.s 1976
Fig. 200
NOT IN KEY
P. latffulcrum BerziQ.s 1976, p. 8-9, Figs. 5-9

Type locality: Lake Frejen, Aneboda, Sweden.

Cephalodella unguitata Hauer, 1935 Holotype: Not designated
Fig. 198
Description: Body dorsoventrally flattened, foot very wide, toes long, straight. Eye-
C. unguitata Hauer 1935, p. 70-71, Fig. 10. spot on distal end of ganglion, two palps on trochus. Inner organs correspond to a
Cephalodella type, zoochlorellae present. The trophus is type A, uncus simple "ir-
Description: Insufficiently described, no mastax figure, drawing from partially con- regular" (?) with a long pointed subuncus. Rami curved with wide alulae, thin lamel-
tracted animal. Considered nomen nudum. lae between the rami. Fulcrum extremely wide, ends in a "pediform" half-crutch. No
epipharynx.
Total length 120-135 !-tm, toes 34-43 !-tm, mastax 25-31 !-tm, fulcrum length 14-17
Cephalodella vacuna Myers, 1924 !-tm, width 5-7 !-tm, palpus 7-8 !-tm.
Fig. 199 BerziQ.s assumed that the animals have previously been mistaken for C. tachyphora
if the palps and trophi were overlooked. Koste (personal communication) is doubtful
C. vacuna Myers, Harring & Myers 1924, p. 503-504, Fig.35:6 about the validity of the genus.
Ecology: Acidophilic, found regularly in several lakes and bogs in the Aneboda area
during 1950-1968, either in bottom sediment, among Sphagnum, or Utricularia. pH:
Type location: Near Atlantic City, NJ, D.S.A. 4.7-6.0, conductivity 31-48 !-tS, Pt: 115-125 mgL- 1, KMn0 4 : 102-108 mg L- 1, total Fe:
Holotype: Not designated
1
0.1-0.4 mg L- 1, alk. 0.1 mg L- .
Literature: Not seen outside type area.
j' 'IJ...
/u;Ltt:Wl SpecIes:
'c' /,/1/
• 50 V~L{~t;aett5IS S~
Description: Body elongate, fairly slender, nearly parallel sided and somewhat pris- Code: 13,16,25,31,34,42,45,52,61,64,71,81, 91. Cl~~lla,v{) (11IJ (I]l.li dl11i1
matic. Body obliquely truncate posteriorly; foot large and conical, projecting slightly
('''' ,/ N · )....::,00...
C. 0fUI,10{Orl~ f

beyond the small, rounded tail. Toes very long, faintly recurved and slender, tapering (];rod;;.iJ
gradually to acute tips. Corona convex and strongly oblique without projecting lips. Genus Pleurotrochopsis Berzil1s, 1973
Trophus type B: the fulcrum is slightly expanded posteriorly and the manubria
strongly crutched (no figure supplied). Ganglion large and elongate saccate. Neither Pleurotrochopsis Berzil}.s 1973 p. 449.
eyespot nor RCO are present.
Total length 220 ~tm, toes 62 !-tm. Type: Pleurotrochopsis anebodica Berzil}.s 1973.
Ecology: Acidophil, rare. Type locality: Aneboda, Sweden.
Literature: Ban-ing & Myers 1924.
 148 149

Description: Body stout, gibbous, ventral side flat; foot with t\vo joints, toes short,
blunt. Integument and foot carries thorns or hooks and rows of thorns of variable con-
figuration, some (if present) are assumed to serve as an anchoring device. Cerebral
eyespot, trophus virgate. Single species with three ecotypes, some poorly described.

Pleurotrochopsis anebodica BerziQ-s 1973

Fig. 201

Pleurotrochopsis anebodica Berzil}.s 1973 p. 449-451, Fig.l.

Pleurotrocha multispinosa Fadeew 1925 p. 72, Figs. 1: 1-2

Type locality: Aneboda, Sweden.

Holotype: Not designated

Description: Body stout, gibbous, ventral side flat, but body outline varies in the liv-
ing animal. Dorsal side of head with variable appendages. Stomach and intestine well
separated; stomach with a series of parallel lines in P. anebodica anebodica, but
200 smooth in the Russian ecotype P. anebodica multispinosa, and with pronounced
round cellular structure in the Canadian (unnamed) ecotype. Spination also different
in the three ecotypes. Eyespot on ventral or distal end of ganglion, with diffuse red
pigment granules in the Canadian ecotype. Trophus also variable as far as the unsatis-
factory drawings of Fadeew and Berzil}.s allow a comparison with Nogrady's (1989)
figure. Rami triangular or square with one or two terminal teeth, Unci with two strong
teeth, manubria short, with one, two or no basal plates or one median alula. Fulcrum
short, large paddle in lateral view.

Comparison of variable features of subspecies:

multispinosa Canadian
Large pointed Large pointed
lamella lamella and
small horn

Smooth Smooth

dorsally, 5 distal
horizontal 4x3 ventral
rows; no lateral
Fig. 200. Paracephalodella latifulcrum dorsally, laterally, trophi, head. Scale bar 50 /lm. no ventral;
F.ig. 201. Pleurotrochopsis anebodica Group A: laterally, foot dorsally, trophus, fulcrum laterally, adhe- 3 rows la-
SIOn organ; Group B: laterally, head of another animal, foot dorsally, trophi dorsally and laterally. (201: terally?
BerziJ;ls 1967; 201: A: Berzil}.s 1973, B: Nogrady 1989).

Lateral hooked present on double on absent

lamella both sides both sides?
 150 151

Total length 180-220 ~tm, toes 13-15 ~un, trophus 32 ~tm.

Ecology: In acidic humic ponds (Sweden): pH 6.7, 18°C, 64 ~S cm-I, Fe 0.2 mg L-I,
°2
8.1 mg L-I, KMn0 4 77 mg L-I. In Canada pH 8.2, 17°C, 640 ~S cm-I; no data from
Kharkov, Russia.
Literature: Fadeyew 1925, Nogrady 1989.

[.
~~[
Genus Taphrocampa Gosse~ 1851

Taphrocampa Gosse, 1851, p. 199

Type: Taphrocampa annu.losa Gosse

Type locality: Leamington, England.

Description: Velmifoffi1, stretchable, dorsally plicated (accordion-like) body, ob- 202 203
lique corona with auric1es. Cerebral eyespot. Mastax asymmetric, virgate, unci with
2-3 teeth, rami with large alulae.

KEY TO TAPHROCAMPA

1. Right manubrium only 2/3 of left one; body slim clavigera

Manubria of almost equal length; body stout 2.
2(1). Toes long, curved; rami with inner teeth selenura
Toes short, conical; ramus with one blunt tooth annulosa
I
Taphrocampa annulosa Gosse, 1851

~
Fig. 202

T. annulosa Gosse, 1851, p. 199. 204

205
Type locality: Leamington, England.
Holotype: Not designated
Fig. 202. Taphrocampa annulosa laterally, dorsally, trophus, unci.
Fig. 203. Taphrocampa selenura dorsally, laterally, trophus, unci.
Description: Variable body shape; sluggish, capable of swimming as well as inching/ Fig. 204. Taphrocampa clavigera.
creeping, but auric1es seldom extended. Tail, foot, and toes short. Trophus asymmetri- Fig. 205. Eothinia carogaensis laterally, trophus ventrally, tip of ramus, foot laterally. (202,203,204:
cal, left ramus with indistinct single inner tooth, unci with two teeth each; manubria Harring & Myers 1924; 205: Myers 1937).
almost equal length.
Total length 130-230 /lm, toes 11-15 /lm, trophus 25-30 f-tm. Type locality: England.
Ecology: Cosmopolitan, in detritus. Holotype: Not designated
Literature: Harring & Myers 1924, Wulfert 1956, Koste & Shiel 1991.
Description: Body cylindrical, tapering to medium long foot that is covered by broad
tail; tail separated from abdomen by groove. Toes long, slender, semicircular when
Taphrocampa selenura Gosse 1887 viewed dorsally. Trophus asymmetrical, but manubria almost equal in length; right
Fig. 203 ramus with eight teeth, left ramus with a blunt tooth, a lamella, and then two blunt
teeth. Left uncus with two teeth, right uncus with three. Integument usually sticky and
T. selenura Gosse, 1887a p. 1, Fig. 1: 1 covered by detritus.
152 153

Total length 220-290 ~lm, toes 25-33 !Im, trophus 36 !Im . ent otherwise. Koste 1978 and Koste & Shiel 1991 describe E. elongata only. Eight
Ecology: Cosmopolitan but not abundant in eutrophic detritus; limnosaprobic. species are known; the American ones are acidophils, some have not been seen since
Literature: Harring & Myers 1924, Koste & Shiel 1991. discovery. Six species.

Taphrocampa clavigera Stokes, 1896 KEY TO EOTHINIA

Fig. 204
1. Corona extends down the ventral side 2.
T. clavigera Stokes, 1896, p. 18, Fig 7:2. Corona apical only 3.
2. Rami asymmetrical, row of rami teeth longer on the left side .
Type locality: Trenton, NJ, U.S.A. ................................................................................................. carogaensls . *
Holotype: Not designated Rami with uniform row of teeth on both sides triphaea
3. Rami asymmetrical: 4+2 and 5+1+2 teeth argus *
Description: Body elongate, fusifonn and slender; plication less exuberant than 011 Rami symmetrical 4.
congenerics and disappears on fully extended swimming animal. No true foot or tail, 4. Ramus with tenninal tooth and rows of 5-6 teeth eukolpa *
toes small; between the toes a minute papilla. Corona extends all the way down the Ramus without tenninal tooth 5.
ventral surface of the head, prominent rostrum curves over corona. Right salivary 5. Rami with 4-5 short and 3 very long teeth poitera * ,<

gland very large, left one rudimentary; eyespot a mass of pigment granules. Mastax Rami with a row of fine teeth elongata
strongly asymmetric: alula of left ramus almost as long as the ventral part and fonns Rami without teeth or with 3 teeth; fulcrum possibly with dorsal lamella ...
an acute angle with the fulcrum. Right alula reinforced by a rib. Right manubrium .................................................................................................... lamellata(?)
only two-thirds of the left one. Unci with two teeth.
Total length 120-200 Ilm, toes 10 !-tm, trophus 26-38 [lm.
Ecology: Rare, periphytic, also in Sphagnum. North America, Europe. Eothinia carogaensisMyers, 1937
Literature: Harring & Myers 1924, Koste 1978. Fig. 205

E. carogaensis Myers, 1937, p. 9-10, Figs. 3,6,9,10.

Species inquirendae: T. lemurensis BerziI].s 1982
T. levinseni Bergendal 1892 Type locality: Lake Caroga, Adirondack Mountains, NY, U.S.A.
insufficiently described. Holotype: AMNH 878 (lost ?)

Genus Eothinia Harring & Myers, 1922
Eothinia Harring & Myers, 1922, p. 555 and 646.
Type: Eosphora elongata Ehrenberg 1832 = Eothinia elongata.
Type locality: Berlin.
Description: Body long, slender, the usual notommatid shape, head and neck well
separated by folds; trunk transparent, shows longitudinal striation. Foot two-seg-
mented, two toes. Corona oblique, may have a ventral chin; marginal cilia sparse, ex-
cept on auricles. Oesophagus long, stomach and intestine separated. Retrocerebral
organ distinct, both sac and glands present; cerebral eye and two widely spaced fron-
tal eyes. Single salivary gland. Trophus virgate, triangular rami with a row of fine
denticles on inner margin, unci with one to four teeth. Manubria rod-shaped with
clubbed end, fulcrum long, tenninally widened; pleural rods present. The genus was
created to accommodate Eosphora elongata, with trophus different from other
Eosphora species; it shows close similarity to Sphyrias but is morphologically differ-
Description: Body cylindric, parallel sided, neck-fold distinct; foot short, character-
istically wrinkled, toes short with narrow base, flaring out. Corona apically convex,
continuing down the ventral side. RCO present, one cervical and two frontal eyespots.
Trophus virgate/forcipate combination. Fulcrum long, formed of two plates joined
dorsally, thus V-shaped in cross-section. Rami triangular, distal third abruptly curved
dorsally with broad decurved alulae. On the apical inner margin fan-shaped combs of
fine teeth, larger on the left side. Unci with single tooth, manubria curved rods,
clubbed at base. Epipharynx just two thin triangular plates, righthand salivary gland
larger than left. Trophus similar to E. elongata, but the wrinkled foot and strongly
ventral corona distinguishes it.
Total length 260 !-tm, toes 25 !-tm.
Ecology: Rare, among Sphagnum, Nitella, and Utricularia, pH 6.4-7.0.
Literature: Not seen since discovery.
154 155

Description: Body fusifonn, transparent, flexible, transverse folds deep~ head,

dorsally viewed, is trapezoidal. Foot has two joints, toes with bulbous base, very long,
incurved and decurved ventrally. Corona extends down the ventral side. Trophus
virgate, fulcrum extremely broad at base, tapering abruptly to a slender incurved sec-
tion. Rami broadly triangular, with a unifonn series of short sharp teeth along the
whole inner margin. Unci have a subsquare basal plate \vith a single clubbed principal
tooth which carries two slender auxiliary teeth at its ventral margin. A second tooth
crosses the basal plate diagonally to its dorsal edge. Basal plate of manubrium large,
subsquare, handle stout and decurved. Two slender pleural rods.
E. lasiobiotica differs only in the absence of rami teeth and the shape of the basal
foot joint; Berzil1s' figures are not in agreement with his description and are not clear.
Total length 175-250 !lm, toes 25-35 ~lm, 28 !lm long, 35 !lm wide.
Ecology: Rare, among Sphagnum in shallow ponds. ,.,,,,:,,,,.....
208 Literature: Berzil1s 1949b.

206 Eothinia argus Harring & Myers 1924

Fig. 207

E. argus Harring & Myers 1924, p. 530-531, Figs. 41 :6-11.

Type locality: Lenape Lake, NJ, U.S.A.

Holotype: Not designated

207
Fig. 206. Eothinia triphaea laterally, dorsally, trophi, unci.
Fig. 207. E. argus dorsally, laterally, trophi ventraIly, laterally, oblique frontal view, unci.
Fig. 208. E. euko/pa dorsally, trophus, manubrium, uncus. (206,207: Harring & Myers 1924; 208: Myers
1940).
Description: Body cylindrical, slightly broadened and rounded distally~ tail small,
subsquare, trilobate. Foot has two short joints of equal length, toes short, conical,
slightly decurved. Ganglion large, saccate, RCO long; one cervical and two frontal
eyespots. Trophus virgate, rami slightly asymmetric, triangular with large alulae; they
form a heart-shaped opening above the fulcrum. Right ramus carries four small teeth
above this opening, followed by two larger apical teeth and two small dorsal teeth. On
the left ramus the sequence is 5+1+2. Rami carry two unique blade-like lamellae
dorsally. Unci irregularly shaped with large clubbed tooth and two small accessory
teeth attached to ventral side of the clubbed tip of the main tooth. Adjoining the tooth
on the dorsal side is a narrow denticulate portion. Manubria straight with two basal
lamellae, inner lamella carries a terminal expansion. Fulcrum long and broad, taper-
ing to an incurved terminal tip. Two undulate pleural rods with spooned anterior end.
Total length 250-300 !lm, toes 18-22 !lm, trophus 38!lm.
Ecology: Softwater species, rare.
Literature: Not seen since discovery.

Eothinia triphaea Harring & Myers, 1924 Eothinia eukolpa Myers 1940
Fig. 206 Fig. 208
E. triphaea Harring & Myers, 1924, p. 528-530, Figs. 41:1-5 E. euklopa (eukolpa injigures) Myers 1940 p. 5, Figs. 1:2; 2:5,10,12.
E. lasiobiotica Berzil1s, 1949b, p. 318, Figs. 18-21.
E. eukolpa Koste 1978, p. 314.
Type locality: Vilas and Oneida Counties, WI, U.S.A. Type locality: Round Pond, Carbon County, PA, D.S.A.
Holotype: Not designated
 156 157

Holotype: Not designated

Description: Body stout, hyaline, head short, neck well marked. Foot single-jointed
(shown double-jointed in figure), base of toes swollen with short papillose tips.
Subcerebral glands huge, with clustered black bacteroids along margins. Trophus
virgate, rami triangular, symmetric, with a denticulate comb of five or six slender
teeth; tips of rami expanded then suddenly diminishing into a blunt terminal tooth.
Unci with a single clubbed tooth on a semicircular lamellar plate.
l~\ Total length 200-210 !-tm, toes 12 !-tm, trophus 38 J.tm.
Ecology: Fairly common among algae and Utricularia, pH around 6.6.
o Literature: Not seen since discovery.

c Eothinia poitera Myers, 1933

Fig. 209

E. poitera Myers, 1933b, p. 10-11, Fig. 6.

Type locality: Barcelona River, Mount Desert Island, ME, D.S.A.

Holotype: Not designated

Description: Body short, stout, hyaline and t1exible; head also stout with well defined
209 neck-fold. Foot short, single-jointed, toes narrow and sigmoidal in side view, incurved
dorsally. .c_QIQna~~tend_s_JaLdQWJ1JJ;:H~ ..Y~ntmL§.i.g~. Trophus virgate; fulcrum broad
tapering to a rod. Rami triangular and symmetrical with pointy alulae; on the inner
margin 4-5 short teeth are followed by three very long acute teeth. Unci with single
tooth attached to web-like plate. Manubria stout, apical portion with three fenestrae,
clubbed terminally. Two slender pleural rods.
Total length 190-220 !-tm, toes 13-18 ~m.
Ecology: Among decaying Utricularia, rare.
Literature: Not seen since discovery.
'~
Eothinia elongata (Ehrenberg) 1932
Fig. 210

Eosphora elongata Ehrenberg 1832 p. 140.

Eothinia elongata, Harring & Myers 1922, p. 555, 646-648, Figs. 61:1-5.
E. elongata macra Berzit;ls, 1949b, p. 317-318, Figs. 16, 17.
E. lamellata Berzit;ls 1949b, p. 317, Figs. 12-15.

Type locality: River Saale, Germany.

A B
210
Fig. 209. Eothinia poitera laterally, trophus laterally, oblique dorsally, toes. Fig. 210. E. elongata Group
A: dorsally, trophi dorsally, laterally and oblique ventral; Group B: Wulfert's figure laterally, ganglion
wIth subcerebral glands, pleural rods, mastax with single salivary gland; C: trophus of E.lamellata, note
fulcrum lamella; Group D: Donner's figure of E.lamellata, incus dorsally and laterally, manubrium, and
crotched pleural rod. (209: Myers 1933; 210: A: Harring & Myers 1922, B: Wulfert 1960b, C: Berzins
1949b, D: Donner 1978). '
Holotype: Not designated
Description: Cylindrical, slightly gibbous body, transverse folds indistinct, longitudi-
nal folds marked, lobed tail. Extremely transparent, sluggish. Corona frontal only,
vitellarium a band, footglands the length of the foot, right salivary gland huge.
Trophus with triangular rami, inner edges with numerous denticles, symmetrical.
 158
Unci with one tooth, manubria straight, clubbed on both ends, fulcrum fused from two
plates, distal end fringed. Predator. Koste (1978) questions the relationship of this
species with the others of the genus. Male known.
Total length 350-500 ~lm, toes 32-42 ~lm, trophus 56-65 ~lm, male 150-215 /lm.
£.e. macra BerziQs, 1949 is distinguished by its larger size (460 [.lm), shorter foot,
and differences in the trophus. On Potamogeton natans in Lake Fiolen, Sweden; we
consider it a synonym.
£. lamellata (Fig. 210 C) is synonymised, as differences from nominate species are
insignificant or unproven, and descriptions confusing. Rami symmetrical, usually
three teeth, large alulae, covered by small oral plates. Small sharp epipharynx present,
Donner (1978) (Fig. 210 D) shows two crutched pleural rods but not the lamella on the
fulcrum (there is no crutch in E. elongata, see Fig. 210 B). Unci with an appendix on
the underside. The fulcrum and/or the short manubria differentiate elongata and
lamellata on the basis of the manubria being longer or shorter than the fulcrum. How-
ever, the manubria of both species are shorter than the fulcrum.
Total length 340 /lm, toes 26-33 /lm, trophus 53-60 [.lm.
Ecology (E. lamellata): Acidophilic, but also found at pH 7.9-8.0 in an oxbow of the
Danube near Vienna, Austria.
Literature: see text, Donner 1978.
Ecology: Carnivore, particularly belloids. Rare but cosmopolitan.
Literature: BerziQs 1949b, Wulfert 1960b, Koste & Shiel 1991.
Genus Eosphora Ehrenberg, 1930
Eosphora Ehrenberg, 1930, p. 47.
Type: Eosphora najas Ehrenberg, 1830 pp. 47,88, Fig. 7:3.
Type locality: Berlin, Gennany.
Description: Body stout, saccate, pseudo-segmented with round or lobed tail. Foot
with or without segments, two toes with long footglands. Corona apical, circumapical
band with dorsal gap, lateral ciliary bundles and a sparsely ciliated buccal field.
Apical sensory antennae common, cerebral eye at distal end of ganglion (except in E.
anthadis). Retrocerebral organ and subcerebral glands present. Mastax three-lobed,
unci usually with one tooth; rami triangular, strongly fenestrated, with varying
number of teeth, often with basal apophysis. Fulcrum wide, plank-shaped, epipharynx
paired. Six species.
KEY TO EOSPHORA
1. Eyespot absent anthadis
Eyespot(s) present 2.
2(1). Footgland without mucus reservoir. ehrenbergi
Footgland with mucus reservoir 3.
3(2). Salivary gland rudimentary, animal very transparent. therina *
Salivary gland large 4.
159
4(3). Salivary gland asymmetrical; papilla at base of toe najas
Salivary gland symmetrical ~. 5.
5( 4). Fulcrum in lateral view broad thoa
Fulcrum slim, narrow thoides
N.B. Myers (see ':Vulfert 1940, p. 585) warned against relying on salivary glands; they are used here be-
cause they may stIll be a better taxonomic criterion than foot segmentation or trophus length. Obviouslv.
further work on the genus is overdue. ~ -
Eosphora anthadis Harring & Myers, 1922
Fig. 211
E. anthadis Harring & Myers 1922, p. 641-642, Figs. 58:9-13.
Type locality: Not specified, "widely distributed".
Holotype: Not designated
Description: Body robust, head wide, trapezoidal, integument transparent and soft.
Foot not (or rarely) segmented, toes short, wide, dorsal outline of the pair is hemi-
spherical, with small claws. Salivary gland absent, gastric glands very large, ovoid;
footglands elongated, consisting of the gland proper and a terminal mucus reservoir
(see also E. thoa). Ganglion small, RCO present, no eyespot. Trophus modified
virgate, delicate. Rami triangular, bent at right angle. At this bend between the two
openings four or five small teeth on each ramus (visible in apical view only; Kutikova
(1970) and Koste & Shiel (1991) do not show that view). In the Borneo (Koste
1988b)(Fig. 211 B) and Brazilian specimens (Koste et al. 1984) the rami have a
denticulate membrane instead of teeth, and are not strongly bent. Manubria slightly
bent, unci small plates with one long tooth, fulcrum consists of two plates fused
dorsally in a V-shape, distal end flaring and scalloped.
Total length 276-410 /lm, toes 16-28 /lm, trophus 33-35 /lm.
Ecology: Described from acidic water, but has also been found in strongly alkaline
waters (e.g., Neusiedlersee, Austria), and actually seems to prefer those (Harring &
Myers 1924, p. 525). Such major ecological discrepancies make the homogeneity of
the taxon suspect. Cosmopolitan but rare, sluggish.
Literature: Russel 1960, Donner 1975, Koste et al. 1984, Koste 1988b, Koste &
Shiel 1991.
Eosphora ehrenbergi (Ehrenberg), 1832
Fig. 212
Notommata najas Ehrenberg 1832, p. 132.
E. ehrenbergi, Weber & Montet 1918, p. 123.
Type locality: Berlin, Germany.
Holotype : Not designated
Description: Body robust, brownish in living specimens, body folds indistinct, abdo-
 160 161

men terminally rounded with small tail. Foot double-jointed, toes short, conical. Large
salivary glands are asymmetrical (right one larger), footglands long without mucus
reservoir. RCO and cerebral eyespot present. Trophus virgate, rami triangular, bent,
strongly fenestrated with double tenninal teeth; fulcrum very broad with diagonal ter-
minal edge; unci with single tooth. Manubrium almost straight with triangular anterior
plate directed dorsally. Epipharynx consists oftwo tripods. Male known (Wesenberg-

P
~
20 m .' J~ ,~d
~
Lund 1923, as Notommata najas.)
Total length 350-450 !lm, toes 24-30 !lID, trophus 65 !lm, male 212 !lm.
Ecology: Cosmopolitan, not frequent but occurs in large numbers when present.

m
Predator.
/.:.• LJ, z,· .'. Literature: Barring & Myers 1922, Koste & Shiel 1991.

.,,~

Eosphora najas Ehrenberg, 1830

8 Fig. 213
211 E. najas Ehrenberg 1830, pp. 47, 84, Fig. 7:3

Type locality: Berlin, Gennany.

Holotype: Not designated

Description: Body robust ovoid, sometimes orange in living animals; head and neck
folds distinct, collar-like; integument firm, tail small with three lobes. Foot indis-
tinctly three-segmented, toes straight, slender, with a sensory (?) papilla at their base
and mucus reservoir. Salivary glands large, asymmetrical; one cerebral eyespot at
anterior end of ganglion (according to Barring & Myers 1922 on posterior end), two,
sometimes four frontal eyespots, the lateral ones on a papilla. Rami powerful, triangu-
lar, with two stronglyrecurved apophyses; number of teeth seems to be variable: ac-
cording to Barring & Myers (1922) there is one tooth on each side; Wulfert (l960b)
and Koste (1978) describe 3-4 large and several small teeth; Koste & Shiel (1991)
state the presence of one tooth on the left side, two on the right (potentially not E.
najas (R. Shiel personal communication). Unci with one clubbed tooth and two small
teeth, with small attached dorsal plate. Manubria short, stout, fulcru)Tl.y'~)jY broad... ,vir
Epipharyngeal rods right-angled, two small serrated oral plates. ~~/1'~fi::er1 t~~e
Total length 260-610 !lm, toes 36-48 !lID, trophus 80 ~m, male up to 300 flID, rest- (-<00" .>'
ing egg 130x170 flm. '
212 Ecology: Cosmopolitan, predator living on small rotifers, often limnosaprob. pB 6-8,
13-22°C, 60-400 flS cm-I.
Literature: Harring & Myers 1922, Pourriot 1960, Wulfert 1960b, Koste & Shiel
213 1991.

Fig. 211. Eosphora anthadis Group A: North American specimens; Group B: Indonesian specimen' ~~te
lamella on ramus. . ' Eosphora therina Barring & Myers 1922
Fig. 212. E. ehrenbergi. Fig. 214
Fig. 213. E. najas. Scale ~ars 50 and 10 f-lm. (211 A: Harring & Myers 1922, B: Koste 1988a' 212: Harrin
& Myers 1922,213: Hamng & Myers 1922). ' g
E. therina Barring & Myer~ 1922, p. 639-641, Figs. 59:1-5.

Type locality: Small puddle near Potomac River, Washington, DC, U.S.A.
162 163

Holotyp,e: Not designated

Description: Body broad, very robust, extremely transparent; head not separated but
a fold between neck and trunk is present. Foot three-jointed, tapering, toes small with
a minute claw. Gastric glands large, salivary glands rudimentary, footglands with
mucus reservoir. Cervical eyespot large. Trophi very similar to those of E. ehren-
bergi: they are virgate, rami triangular, almost symmetrical, bent, strongly fenestrated
with double terminal teeth; fulcrum very broad with diagonal terminal edge; unci with
a single tooth on a basal plate. At the dorsal side of this plate a triangular striated ex-
pansion (auxiliary tooth ?). Manubrium almost straight with triangular anterior plate
directed dorsally. Epipharynx consists of two tripods. The small tail, tapering foot,
transparency and extreme swiftness distinguishes this species from E. ehrenbergi.
Total length 450-475 ~m, toes 25-28 ~m, trophus 72 ~m.
Ecology: Found in a tiny muddy puddle covered with Euglena, in enormous numbers.
Literature: Not seen since discovery.

Eosphora thoa Harring & Myers 1924

Fig. 215

E. thoa Harring & Myers 1924, p. 523-524, Figs. 39:1-5.

Type locality: Cemetery Pond, near Eagle River, Vilas County, WI, U.S.A.
Holotype: Not designated

Description: Resembles E. anthadis in appearance: body broad, robust, head and

neck fused but separated from trunk by a fold, body striated. Foot conical, wrinkled,
two toes heart-shaped ending in fine points. Footglands with mucus reservoir. Large
cerebral eyespot. Rami strongly curved, with two very large apophyses proj ecting
ventrally. Near the middle of rami a blunt tooth on either side, posteriorly from those
about 20 small conical teeth. Unci with single large tooth, fulcrum very short and
broad. Epipharyngeal plates very large, covering rami (in Fig. 215d they are shown
turned up; their normal position is seen in Fig. 215c). The presence of an eyespot, the
epipharyngeal plates, rapid swimming, and avoidance of alkaline water distinguishes
this species from E. anthadis.
Total length 300-500 ~m, toes 20-35 ~m, trophus 50 ~m.
Ecology: Soft water, rare, pH 6.5-6.7, 16-24°C, 260-310 ~S cm-I.
Literature: Koste & Shiel 1991.

215 216
Eosphora thoides Wulfert 1935
Fig. 216 Fig. 214. Eosphora therina.
Fig. 215. E. thoa. . 4
E. thoides Wulfert 1935, p. 600, Figs. 15 a-d. Fig. 216. E. thoides Scale bars 50 and 10 !J.rn. (214: Harring & Myers 1922; 215: Hamng & Myers 192 ;
216: Wulfert 1935).
Type locality: River Saale, Germany.
Holotype: Not designated
164
Description: Habitus resembles E. thoa; foot wide, tapered, 3-4 segmented, toes tri-
angular in side view, foot-glands asymmetric, mucus reservoirs large. Trophus more
delicate than those of E. thoa: fulcrum long, narrow, distally widened; rami triangular,
curved, apparently without teeth. Uncus with one tooth, base plate fenestrated;
manubria somewhat asymmetric, wing-shaped single alula at base. Epipharynx bifur-
cate.
Total length 460-510 flm, toes 26-35 !-tm, trophus 37 !-Lm .
Ecology: Benthic, rare, Europe, Australia, New Zealand.
Literature: Koste & Shiel 1991.
165
The study of living animals and of the trophi is mandatory, as it is almost impossi-
ble to obtain preserved specimens with extended auricles, even using narcosis. About
half the species described are known only from North American acid waters and there
are several Australian endemic taxa. About 52 species are described in the literature.
NOTOMMATA DESCRIPTORS
01 TOTAL LENGTH including toes

01 > 500!-Lm
Genus Notommata Ehrenberg, 1930 02 300-500 flm
03 < 300!-Lm

ROGER POURRIOT 02 DIFFERENTIATED BLADDER

Laboratoire de Geologie App1iquee, B 123, Universite Paris 6, F 75232 Paris Cedex
05, France. 01 present
02 absent
Notommata Ehrenberg 1830, p. 46.
03 RETROCEREBRALORGAN
Type: Vorticella aurita Muller 1786 = Notommata aurita (Miiller)
Type locality: Copenhagen. 01 overlaps mastax
02 does not overlap mastax
Description: De Beauchamp (1907, 1965) considered the Notommata genus to be 03 subcerebral glands absent
relatively primitive. The species vary in size from 130 flm to 1 mm, their bodies being 04 subcerebral glands overlap sac
more or less elongated with an abdominal bulge narrowing into a well differenciated
retractile head and foot. While body size is variable and subject to fluctuation depend- 04 TOES
ing on the ecotype or geography, the genus can still be considered a crude taxono-
mical unit. Transverse and longitudinal folds on the trunk are variable. The two- 01 >40!-Lm
jointed foot ends in two more or less tapering toes. The foot is often covered by a 02 20-40 !-Lm
cuticular expansion, incorrectly called a tail and frequently bears a sensory papilla, or 03 <20 !-Lm
a short spur between the toes. A retrocerebral organ, a virgate mastax (functioning
like a pump) and a reduced, strongly oblique or ventral corona often bearing retractile 05 "TAIL"
auric1es is characteristic. The dista1 post-oral end of the corona is also known as a
"chin". The eye, when present, is cervical but often concealed by the opaque granules 01 long, conical
of the retrocerebral sac. 02 spur-like
The trophi are more or less asymmetrical, the left side - viewed dorsally - being 03 broad, usually covers first foot joint
more robust. Characteristic preuncial teeth (ventrally to the first larger tooth), and a 04 shorter than first foot joint
cuticular epipharynx are present. The absence of a well-differentiated ventral bladder 05 indented
in some of the species is sufficiently remarkable to be retained as a taxonomic crite- 06 trilobate
rion (Pourriot, 1989). 07 reduced or absent
The Notommata are benthic; while they are able to swim, none of them is truly
planktonic. Their mastax, adapted for suction, enables them to ingest food of various 06 TROPHI
types and sizes, though this does not exclude a certain specialization in their diet (see
Pourriot, 1965, 1977). A highly developed tooth on each uncus is used for piercing of 01 asymmetric
algal cell walls; many species are raptorial. Males and resting eggs are unknown in 02 symmetric
some species. The biology of reproduction has been studied, particularly in N. copeus, 03 specialized or modified, small or slender
by Pourriot and Clement (Pourriot 1963, Clement 1977, Pourriot & Clement 1981). 04 preuncial teeth present
 166 167
07 DISTAL FOOT JOINT 13,21,31,43, 57?, 61, 72, 81,94? mera *
13,21,31,43,57,61'1,64,71,81,93 voigti
01 with short spur, papilla or tuft of cilia 13,21,31,43,5~ 61,72,83,93 cyrtopus paracyrtopus
02 none of the above 13,21,31,43,57,63,73,83,94 sylphoides
03 foot absent 13,21,32,41,54,62,64,72,81,93 doneta
13,21,33,42,54,62,72,83,94 tithasa 'I< moved to Pleurata
08 EYESPOT 13,21,33,43,52,61,64,71,82,93 tripus
13,21,33,43,54,61,72,81,93 apochaeta'l<
01 present 13,21,33,43,54,62,72,82,92 stitista '"
02 reduced to pigment granules 13,21,33,43,57,61,71,82,94 silpha
03 absent 13,21,33,43,57,61,72,827,93 venusta *
13,21,34,43,54,62,72,81,94 fasciola *
09 AURICLES 13,227,33,43,57,62,72,82,92 rugosa '"
13,22,31,43,54,61,72,81,930r94 omentata
01 long, T-shaped 13,22,32,43,54,61,647,71,82,93 groenlandica
02 medium length 13,22,32,43,54,62,72,81,93 placida '"
03 short 13,22,32,43,56,63,72,82,94 contorta
04 rudimentary or absent 13,22,33,43,53,62,72,83,94 tyleri '" moved to Pleurata
13,22,33,43,54,63,72,81,94 trypeta '" moved to Pleurata
13,22,33,43,57,62,64,72,83,94 aethis '"
NUMERICAL KEY TO NOTOMMATA
* species marked with an asterisk have not been seen since description.
11, 22, 31, 41, 52, 61, 71, 81, 91 copeus
11, 22, 31, 41, 53, 61, 71, 81, 91 cerberus N.B. Species descriptions are arranged alphabetically.
11,22,31,41,53,61,71,81,92 p. pachyura
11,22,31,42,53,61,71,81,92 collaris
11,22,31,42,56,61,71,81,92 falcinella DICHOTOMOUS KEY TO NOTOMMATA
11,22,31,42,56,61,72,81,93 endoxa
11,22,31,42,56,62,72,81,92 pseudocerberus -I. RCO well developed 2.
12,21,31,41,53,61,717,81,92 allantois Subcerebral glands absent 33.
12,21,31,41,54,62,64,72,81,93 diasema 2(1). Differentiated bladder 3.

!~~~d:;~gS~::: : : : : : : : : : : : : : : : : : : : : : : : : : : ::::::::::::::::::::::::::::::IL;~;s
12,21,31,41,54,63,72,81,93 avena *
12,21,31,42,53,62,72,81,92 longina 3(2).
12,21,31,42,54,61,64,727,81,94 peridia
12,21,31,4~55,61,71,81,92 cerberus 4(3). Bladder double a 5
12,21,31,43,53,61,64,72,81,93 codonella Bladder single ;
12,21,31,43,54,61,64,72,81,93 angusta * 5(4). Trophus slender, aberrant avena
12,21,31,43,54,61,72,81,92 glyphura Normal virgate trophus :d?
12,21,31,43,54,61,72,81,92 haueri 6(5). Lateral antennae tubular with long setae pen la
12,21,33,42,54,61,72,81,937 prodota * .
Lateral antennae 0 th erwIlse . ...... 7.
~~~usta*
12,21,33,43,56,61,64,72,81,93 lenis 7(6). RCO hangs on long tubu ar stem .
12,22,31,43,54,61,72,81,93 saccigera .
RC O otherwlse .. 8.
donella
12,22,32,43,57,61,72,81,93 cherada * 8(7). Ta~1 broad ::::::::::::::::::::::::::.~~ 9.
13,217,34,42,57,63,64,73,81,94 bennetchi * Tall rounded 10
13,21,31,42, 55, 61,64,72,81,0~ 82,93 c. cyrtopus 9(8). Median lobe of tail with notch......... '.
13,21,31,43,54,61,64,72,81,93 aurita .
WIthout notch .. . I hauerl
hura
13,21,31,43,54,61,72,81,93 thopica * 10(9). Long subcerebral glands gyp 11
13,21,31,43,54,61,72,81,93 pygmaea * Short subcerebral glands .
 168
169
11 (l 0). Tail simple cerberus Body <250 !-lm 35.
Tail double longina 34(33). Double dorsal antenna; toes <12 !Lm leni.";'
12(3). Eyes present 13. Single d~rsal antenn~: to.es about 35 ~m prodota*
Eyes absent '" 21. 35(33). Foot rudImentary, tall WIth rounded projection tripus
13(12). Toes curved 14. Foot normal, no spur 36.
Toes straight 15. 36(35). Oesophagus expanded distally aetlzis.
14( 13). Toes <30 !-lm c. cyrtopus Oesophagus normal 37.
Toes >40 !-lm doneta
~~~~~: ~:~:::::::::::::::::::::::::::::::::::::::::::::::::::::::::::::::::::::::::::::::::::~:::::.~~~~~
37(36).
15(13). One pair of subcerebral glands : 16.
Two pairs of subcerebral glands diasema 38(37). Trophi symmetric 39.
16(15). Glands with dark globular bacteroids thopica* Trophi asymmetric 40.
Glands without such aggregation 17. 39(38). Rami with long alulae rugosa.
17( 16). Slender symmetric trophi fas ciola * Trophi slender, delicate stitista *
Robust asymmetric trophi 18. 40(38). Manubria with long spur venusta *
18(17) Fulcrum very short bennetchi* Manubria without spur apochaeta *
Fulcrum long 19.
19(18). Toes < 12 !-lm, body <220 !-lm 20. * Species marked with an asterisk have not been seen since description.
Toes> 15 !-lm, body >250 !Lm aurita
20(19). Body <100 !-lm pygmaea* N.B. Species descriptions that follow are arranged alphabetically.
Body ± 165 !-lm mera *
Body >200 ~Lm voigti
21(12). Free-swimming, small trophus silphoides Notommata aethis Myers, 1933
Ectoparasite on Asellus cyrtopus paracyrtopus Fig. 217
22(2). Body >400 !-lm 23.
Body <350 ~m 28. N. aethis Myers 1933a, p. 4-6, Fig. 2.
23(22). Tail long, conical, with articulated tip copeus
Small recurved spur at base of toes falcinella Type locality: Witch Hole, Mount Desert Island, ME, U.S.A.
Neither tail nor spur present 24. Holotype: AMNH 690 (paratype)
24(23). Trophi robust, asymmetric 25.
Trophi slender, symmetric pseudocerberus Description: Body slender and cylindrical without skin fold at the neck, corona very
25(24). Tail large 26. long; foot short and tubular with two short straight toes. Trophus slightly asymmetric:
Tail small endoxa right uncus has a large ventral tooth preceded by three pre-uncial teeth and followed
26(25). Toes <32 !-lm '" collaris by an accessory tooth; left uncus has a large ventral tooth followed by two accessory
Toes >35 !-lm 27. teeth. The structure of the gut is also peculiar: the oesophagus is long and posteriorly
27(26). Without lateral alae pachyura pachyura expanded in a proventriculus. The large and fusiform gastric glands are attached to the
Two lateral and one dorsal ala pachyura triangularis mastax anteriorly and to the stomach posteriorly. The stomach and intestine are sepa-
Two lateral alae ending in distal tooth pachyura spinata * rated by a deep constriction. An expansion of the cloaca functions as a bladder. The
28(22). Unci with large ventral tooth, fulcrum short saccigera retrocerebral sac is long and clear. There is no eye. Related to the contorta-silpha-
l)nci with normal ventral tooth, fulcrum long 29. tithasa group which has no auricles, only tufts of locomotor cilia.
29(28). Suocerebral glands much longer than sac omentata Total length 237-256 ~m, toes 12-16 ~m, trophus 25 ~m.
Subcerebral glands < than sac 30. Ecology: Acidophilic, rare; also found in New Jersey.
30(29). Toes <12 !-lm 31. Literature: Not seen since discovery.
Toes>12 ~m cherada* Code: 13,22,33,43,57,62,64,7~ 83,94.
31(30). Body <150 !-lm placida*
Body> 150 !-lm 32.
32(31) Body and trophi slender contorta
Body and trophi more robust. groenlandica
33(1). Body >300 !-lm 34.
 170
171

short and pseudosegmented, entirely covered by a characteristic large V-shaped and

hooded tail. Toes long, conical. The two lateral antennae are short tubules with a tuft
of setae. Mastax very robust, similar to that of N pachyura with a very strong hooked
ventral tooth on the unci. Two subunci have been observed by Wulfert in N. allantois.
also described in N. pachyura. The double bladder is a specific character. This species
resembles N. pachyura but the double bladder is diagnostic.
Total length 350-550 ~m, toes 35-54 ~m, trophus 75-100 !-Lm .
Ecology: Very common in acid waters and Sphagnum pools, feeding on Desmids.
First found in North-America by Stokes (1896, reported as Copeus quinquelobatus)
and more recently by Shoemaker et al. (1987). Koste & Jose de Paggi (1982) recorded
it from the Neotropis but, again, confusion with N. pachyura cannot be excluded.
Literature: see text and Wulfert (1940), Koste (1969), Pourriot 1989.
217 Code: 12,21,31,41,53,61, 7l?, 81, 92.

Notommata angusta Harring & Myers, 1922

~
Fig. 219

N angusta Harring & Myers 1922, p. 580-582, Figs. 52:9-12.

Type locality: Brick Haven, VA, U.S.A.

~"
", ",.
j
Holotype: AMNH 632

219
Fig. 217. Notommata aethis laterally, toes, trophus dorsally, unci.
Fig. 218. N. allantois (a,b) dorsal and lateral view; (c): foot; (d): trophus dorsally; (e): fulcrum laterally;
(:f): manubria; (g): unci; (h): subunci; (i): right ramus; 0): pleural rods.
Fig. 219. N. angusta.
(217: Harring & Myers 1933a; 218: Wulfert 1935; 219: Harring & Myers 1922).
Description: Body elongate and slender, head bluntly pointed anteriorly, small
auricles. Toes short. The shape of the long retrocerebral sac is characteristic: it is more
than one-third of body length and consists of an ampulle-shaped sack truncated at the
distal end, hanging on a long cylindrical section which then narrows abruptly to a
slender tube. Trophy slightly asymmetric, rami with large alulae. The unci have two
teeth, the ventral tooth larger than the second tooth.
Total length 300-325 ~m, toes 15 !lm, trophus 36 ~m.
Ecology: Found in a polluted creek.
Literature: Not seen since discovery.
Code: 12,21,31,43,54,61,64,72,81,93.

Notommata apochaeta Myers, 1933

Fig. 220
Notommatq allantois Wulfert, 1935
Fig. 218 N apochaeta Myers 1933a p. 8-9, Fig. 4.
N allantois Wulfert 1935 p. 591-593, Figs. 8:a-h
Type locality: Mount Desert Island, ME, D.S.A.
Copeus quinquelobatus Stokes 1896; N. pachyura Donner 1954; N pachyura Wulfert
Holotype: Not designated
1956 (?).
Description: Body rather stout, cylindric; toes small and conical. Trophus a~ymmet­
Type locality: Peat-bog at Luckenwalde, Gennany. ric: unci have a short ventral tooth followed by three accessory teeth. A rudImentary
Holotype: Not designated tooth is attached to the ventral side of the larger tooth of the right uncus. ReO round
and clear, no subcerebral glands.
Description: Large broad and stout body, trunk ovoid, auricles with a wide base; foot Total length 200-212 !lID, toes 10-12 !lID, trophus 34 !lm.
 172 173

Ecology: In Sphagnum; also found in New Jersey and Wisconsin.

Literature: Not seen since discovery.
Code: 13,21,33,43,54,61,72,81,93.

Notommata aurita (O.F.Miiller), 1786

Fig. 221

Vorticella aurita O.F. MUller 1786, p. 288, Figs. 41: 1-3

N. aurita Ehrenberg 1830 p. 46 (also in Ehrenberg, 1838).
N. epaxia Barring & Myers 1924, p. 443-444. Figs. 21:1-5.

Type locality: Copenhagen.

Holotype: Not designated

Description: Of moderate size, has no typical morphological characteristics: body

somewhat stout, tail present and rounded, a two-jointed foot with two straight and
short toes (16-20 f.Lm). According to Wulfert (1960b p. 325) a papilla between the toes
220 may have been overlooked. Chin pronounced. Trophus somewhat asymmetric: unci
221 have only a single well-developed tooth and 1-2 rudimentary teeth, triangular oral
plate and pleural rods present. RCO large and opaque, subcerebral glands reach down
to the sac. Resembles N glyphura, but the smaller rounded tail and trophi are differ-
ent. Male described by Wesenberg-Lund (1'923).
Total length 250-350 f.Lm, toes 16-20 ~m, trophus 34-36 f.Lm, male 180 f.Lm.
N. epaxia Barring & Myers 1922 is distinguished only by a more slender and slightly
smaller body. Not seen since discovery, considered a synonym.
Ecology: Cosmopolitan, eurytopic, predator.
Literature: Harring & Myers 1922, Wesenberg-Lund 1923, Koste & Shiel 1991.
Code: 13,21,31,43,54,61,64,72,81,93.

Notommata avena Myers, 1933

Fig. 222

N. avena Myers 1933a, p. 6-8, Fig. 3.

~
Type locality: Mount Desert Island, ME, D.S.A.
Holotype: Not designated

222 Description: Body cylindrical, slender, transparent; auric1es small, foot has two
joints, toes long, acute. Stomach and intestine not separated, bladder present. Retro-
cerebral sac and glands are equally long, contain bacteroids. Trophus aberrant,
strongly resembling Monommata:short straight board-like fulcrum, rami lyrate and
223 curved, manubria curved rods without lamellae, unci with one clubbed tooth; there are
Fig. 220. Notommata apochaeta. two supporting rods passing under the manubria.
Fig. 221. N. aurita. Total length 340-365 f.Lm, toes 40-45 ~m.
Fig. 222. N. avena.
Fig. 223. N. bennetchi (a,b): laterally and dorsally; (c): trophi ventrally; (d): unci; (e): incus laterally; (t):
Ecology: Acidophil, in Sphagnum, rare.
manubrium. (220,222: Myers 1933a; 221: Harring & Myers 1922; 223: Myers 1942). Literature: Not seen since discovery.
 174 175

Code: 12,21,31,41,54,63,72,81,93.

Notommata bennetchi Myers, 1942

Fig. 223

N. bennetchi Myers 1942 p. 277-278, Figs. 24:4-9.

Type locality: Poponoming Lake, Poconos, PA, U.S.A.

Holotype: Not designated

Description: Body short and stout with an indistinct foot; toes stout and slightly
decurved. Very small auricles. Mastax highly specialized: the pumping action is lost.
A
Fulcrum short, right ramus has a larger alula than the left. Unci with a stout ventral
tooth (with a pre-uncial tooth) followed by a second smaller tooth and two rudimen-
tary teeth. ReO well developed; the retrocerebral glands go beyond the long and duct-
c
less sac. Habitus resembles N. tripus without the appendage between the toes.
Total length 165-212 ~m, toes 24 ~m, trophus 34 ~m.
Ecology: Psammobiontic species, found in two lakes in the Poconos.
Literature: Not seen since discovery.
Code: 13,21?, 34, 42, 57, 63, 64, 73, 81, 94.
y
Notommata cerberus(Gosse, 1886)
Fig. 224-

Copeus cerberus Gosse 1886, p. 34, Figs. 41: 1-3.

N. cerberus De Beauchamp 1908, p. 401, Figs. 1-3.

Type locality: Sandhurst Wood, Berkshire, England.

Holotype: Not designated

~
Description: Fairly slender body of variable length; tail has a rounded median lobe . "~
with a distinct median notch (seen also in N.falcinella, N. glyphura, N. cyrtopus), and ~" .
"
J - •• j
two rudimentary lateral lobes. The foot has two joints, the posterior one carrying a --"
very small tuft of setae. Toes short, straight, abruptly tapering. Trophus somewhat
asymmetric, robust; fulcrum is of moderate size with aY-shaped lamella at the base;
h
unci with three to five teeth. Although variable in size, the granulated retrocerebral
sac reaches nearly to the end of the mastax. Eyespot large, lenticular; four footglands.
Total length 300-600 ~m, toes 15-35 ~m, trophus 46-60 ~m.
Ecology: Cosmopolitan, acidophil, omnivore.
Literature: Harring and Myers 1922, Wulfert 1940, Koste & Shiel 1991.
Code: 12,21,31,42,55,61,71,81,92.
B
Myers (1937) described a closely related species, N. parvida, distinguished by the
shape of toes and two "bosses" (protuberances, bumps) on the dorsal median line of
224
the abdomen. This structure cannot be seen on Myers' figure. The trophus is ~lso Fig. 224. Notommata cerberus Group A: (a): dorsally; (b): foot laterally; (c): foot dorsally;. (d,e,£): tr~phi.
slightly different from N. cerberus. We do not believe specific status is warranted and Group B: (d) male dorsally; (e): rami, fulcrum; (f,g): unci; (h): rami; (i): fulcrum laterally; (j): manubnum.
(224 A: de Beauchamp 1903, Harring & Myers 1922; B: Wulfert 1940).
 176 177

treat this taxon as N. cerberus "fonn" parvida (Myers). Not seen since discovery.

Notommata cherada Myers, 1938

Fig. 225

N. cherada Myers 1938 p. 6-7, Figs. 3, 5, 8,17,18,21

Type locality: Bargaintown, NJ, U.S.A.

Holotype: AMNH 952

Description: Body cylindrical and vennifonn; foot stout with two small toes.
Trophus asymmetric: rami are triangular without teeth, fenestration (basal apophysis 225
?) asymmetric. Unci symmetric with four slender teeth followed by two rudimentary
teeth. RCO rounded and clear, subcerebral glands equal to one half the length of the
sac. The males appeared to be common.
Total length 275-335 !!m, toes 12-15 !!m, trophus 40 !!m.
Ecology: Periphyton, acidophilic, made sudden appearance after many years of sam-
pling at type locality.
Literature: Not seen since discovery.
Code: 12,22,32,43,57,61,72,81,93.

Notommata codonella Harring and Myers, 1924

Fig. 226

N. codonella Harring & Myers 1924, p. 444-446, Figs. 21 :6-10.

Type locality: Town Line Lake, WI, U.S.A.

Holotype: AMNH 556

Description: Body elongate and slender; short two-jointed foot and two straight and
short toes. Auricles medium, chin present. Tail broad, covering a large part of the

~
foot; has a large median lobe and two minute lateral lobes. RCO elongate and conical,
granulated on the posterior margin. Subcerebral glands fusifonn, moderately long, ~
heavily granulated posterior to the eyespot. Trophus relatively small: unci have four '.I' . ,
~ ~~
f' " .. .. .
j':'.
, '
to five teeth; the first ventral tooth is clubbed. The male and spiny resting eggs were
observed in culture by Pourriot (1966). The male is morphologically similar to the
female with a broad characteristic tail. T~is specie~ is ea~i1'y recognized"by the r~tro- I J

ffl Notom at~c~ra:

cerebral organ and the broad truncated taIl. SE)/! 1}RS. et
HOpt.vd k reJh~ ~ 1'-1, h>(,(1a.,gttil
Total length 300-350 !!m, toes 18-21 !!m, trophus 40 !!m, resting egg 120x90 !!m,lx/Aeb"r
male 210 !!m. (J..oo1, ~J,)
Ecology: Acidophilic to circumneutra1, also in Sphagnum. Omnivore, often occurs
together with N cerberus. Fig. 225
Literature: BerziQ.s (1954/55), Pourriot (1966).
B Fig. 226. Group A: N. codonella; Group B: N. c.
• galena.
Code: 12,21,31,43,53,61,64,72,81,93. Fig. 227. N. collaris. (225: Myers 1938; 226: (A):
226 Harring & Myers 1924, (B): Harring & Myers 1922;
227: Harring & Myers 1922, Pourriot 1967).
 178 179

N. codonella galena (Fig. 226 B.)

Harring and Myers (1922) described the closely related N. galena, that differs from N.
codonella by its slightly smaller size (250-300 ~m), a narrower tail and a more
rounded retrocerebral sac. The trophi are nearly identical, including the presence of
the two pre-uncial teeth, absent in N. cerberus. Curiously, this "widely and abun-
dantly distributed" form closely related to N. codonella and N. cerberus has not been
found since its first description. We consider it a subspecific taxon only.

Notommata collaris Ehrenberg, 1,~g6/ I(~ j 1

Fig. 227 .'

Notommata collaris Hudson and Gosse 1886 pl.17, Fig. 6; ? Copeus collaris Voigt c
1912. 228
Type locality: Berlin.
Holotype: Not designated

Description: This large species is recognizable by a broad tail covering the first joint
of the foot. Auricles moderately long and stout with a continuous ciliation. Abdomen
bulges to twice the anterior width. Foot with two joints, terminal joint bears a small
papilla with a tuft of setae. Toes short, straight and conical. Dorsal and lateral anten-
nae small, setigerous. Mastax large and robust, the unci have three teeth; ventral tooth
very large, others reduced. Right ramus with denticulate broad tooth, manubria with
long lamella on dorsal margin, fulcrum very wide, curved. Pleural rods present. RCO
long, clear and vacuolate, subcerebral glands small with sparse granulation. The male
and resting egghave been described by Pourriot (1967) As in N copeus, the external
sheath of resting eggs is thick and dark but not spiny. 5[/1 fJA' 01
~Dec,;vvtlM v~, itA t-o,JavIIA.)
Total length 500-750 ~m, toes 25-32 ~m, trophus 1OO~m; male 240 ~m, resting egg '-< /Aht,f'
170x215 + 15M-m. (0Ji]L, ftl()
Ecology: This species is cosmopolitan but not common, collected in acid waters and
feeding on desmids (Closterium) and filamentous algae.
B
Fig. 228. Notommata contorta.
Literature: Harring and Myers 1922. Fig. 230. N. cyrtopus Group A: Barring & Myers 1922; Group B: N telmata; Group C: (a,b): dorsally
Code: 11,22,31,42,53,61,71,81,92. and laterally; (c): foot; (d): mastax with salivary gland; (e,t): trophi; (g): epipharynx. (228,230A,230B:
Barring & Myers 1922, Donner 1949; 230C: Wulfert 1956).

Notommata contorta (Stokes), 1897

Fig. 228 rudimentary and rarely extended, no chin. Dorsal antenna single (Harring and Myers,
1922) or paired (Donner, 1949). Mastax highly specialized, limited to suction: ful-
Diglena contorta Stokes 1897 p. 630, Fig. 14:5 crum and manubrium long and slender with terminal curve, rami strongly curved and
N. contorta Harring 1913. have prominent alulae. Unci consist of two straight, very slender divergent rods. RCO
consists of a small rounded black sac and two large subcerebra1 glands longer
Type locality: Near Trenton, NJ, U.S.A. (Harring & Myers, 1922) or shorter (Donner, 1949) than the sac. No eyespot accord-
Holotype: Not designated ing to Harring & Myers 1922 and Koch-Althaus (1963); a yellow eyespot according
to Donner (1949)
Description: Slender vermifonn elongate body, abdomen tapers rapidly to the base of Totallenght 206-300 ~m, toes 7-10 M-m, trophus 17-20 ~m.
a minute three-lobed tail. Small foot with single joint or two indistinct joints and car- Ecology: Cosmopolitan, acid waters. In Australia 20-22°C, pH 6.4-6.6, 114-274 ~S
ries two small and acute toes (7-10 ~m). Ciliated corona is reduced and the auricles cm-I.
 181

Literature: Harring & Myers 1922, Donner 1949 1972, Koch-Althaus 1963, Koste
1969, Koste & Shie1 1991.
d Code: 13,22,32,43,56,63,72,82,94.

Notommata copeus Ehrenberg, 1834

Fig. 229

Notommata centrura Ehr.1834, Notommata copeus De Beauchamp 1908.

Type locality: Berlin.

Holotype: Not designated

Description: The largest species of the genus: its size reaches about 1 mm. Long,
conical tail with a blunt, articulated, pointed tip; long, tubular, ciliated, T-shaped
auricles; thick mucilaginous sheath around the body. Foot has two joints; the tenninal
joint bears a small, knob-like projection with a tuft of minute setae. Toes straight and
moderately long. Dorsal antenna long and sharply constricted at its base. The two lat-
eral antennae are short with long setae, their unusual length due to the thickness of the
mucilaginous sheath. Trophus strongly asymmetric, more robust on the left side than
on the right. Fulcrum long and stout, slightly curved; rami irregularly triangular with
basal projection; unci with a very large ventral tooth followed by three much smaller
teeth; manubria long and broad. Retrocerebral sac extremely long, clear and vacuo-
late; subcerebral glands less than half the length of the sac displaying black granu-
lations. The dwarf and rudimentary male has been described and figured by Wesen-
berg-Lund (1923) and Donner (1954) who has also described the resting egg.
Total length 500-1100 ~lm, toes 40-65 ~m, trophus 80-100 !lm; male 280-350 ~m,
toes 36 ~m; resting egg 180x230 ~m.
Ecology: A cosmopolitan species common in acid water and feeding on Zygnemata-
les (Spirogyra, Mougeotia).
Literature: Harring and Myers 1922, Donner 1954, Wulfert 1956, Koste 1969, 1981.
Code: 11,22,31,41,51,61,71,81,91.

Notommata cyrtopus Gosse, 1886

Fig. 230

N. cyrtopus Hudson & Gosse 1886, p. 2:22, Fig. 17:7.

N. telmata Harring & Myers 1922, p.484-486, Figs. 49:1-4
• N. carpatica Rodewa1d 1935, p. 209. Fig. 10.
J Type locality: Widcombe Pond, Bath, England.
Holotype: Not designated

Description: Small, has a variable stout body without a tail (Harring & Myers, 1922)
229 or with a small tail with a median notch confused with the posterior end of the trunk
(according to Wulfert (1956) and Pourriot, unpublished). Foot with two (Harring &
Fig. 229. Notommata copeus (h): male ventrally; (i) head with ReO. Myers, 1922) or three (Wulfert, 1956) joints, carries two slender and outcurved toes
(229: (a,b,e,f): Harring & Myers 1922, (c,d,j): Wulfert 1956, (g): Koste 1969, (h): Donner 1954, (i) de
Beauchamp 1908.
 182 183

(20-30 ~m). Trophus slightly asymmetric: rami approximately triangular with a inner
ventral projection and well-developed sharp alulae; unci have a single larger ventral
tooth. RCO large, pyriform and opaque. The subcerebral glands reach the sac.
Total length 175-250 ~m, toes 22-28 ~m, trophus 30-35 ~m,male 139 J-lm.
Ecology: Cosmopolitan, common.
Literature: Harring & Myers 1922, Donner 1954, Wulfert 1956, Koste 1976.
Code: 13,21,31,42,55,61,64,72, 810r 82,93.

N. cyrtopus "form" telmata (Harring & Myers) 1922 can be distinguished by the pecu-
liar ventrally curved toes, the more slender body and some details in the retrocerebral
organ.
Total length 250 ~m, toes 20-26 ~m, trophus 42 J-lm.

Notommata cyrtopus paracyrtopus (de Beauchamp), 1932

231 233 Fig. 231

N. paracyrtopus de Beauchamp 1932b, p. 394-397, Fig. 2.

Type locality: Creeks near Strasbourg, France.

Holotype: Not designated

Description: Very similar to N. cyrtopus including the trophi, but has a longer foot,
and no eyespot. It is, however, an ectoparasite onAsellus aquaticus andA. meridianus.
Male known.
Total length 275 ~m, male 180 ~m.
Ecology: Ectoparasitic.
Literature: Not seen since discovery.
Code: 13,21,31,43,57,61,72,83,93.

A related free-living species, N. gisleni BerziJ}s 1949 (Fig.261) has been described
from semicontracted specimens (length 135 ~m, toes 15 ~m). It has two ciliated pits
on the head; the description is incomplete; see under Species inquirendae on page 210

Notommata diasema Myers, 1936

Fig. 232

232 GSya:;J N diasema Myers 1936, p.11-12, Figs. 5,9,16.

Type locality: Lenape Lake, NJ, D.S.A.

234 @5c§;J Holotype: AMNH 777

Description: Body stout, cylindrical, transparent, variable. Tail small, one-lobed,

Fig. 231. Notommata cyrtopus paracyrtopus.
Fig. 232. N. diasema. foot two-jointed, long, toes long with slender tip. RCO long, clear, two pairs of
Fig. 233. N. doneta. subcerebral glands, unique in this genus (Myers 1938). Trophus symmetrical, unci
Fig. 234. N. endoxa. (231: de Beauchamp 1932b; 232: Myers 1936; 233: Harring & Myers 1924; 234: with 5-6 preuncial teeth on first (ventral) tooth, which is shorter than the second one.
Myers 1933a). Related to N. doneta.
184
Total length 312 r-tm, toes 42 ~tm, trophus 40 r-tm.
Ecology: Acidophil, psammophilic.
Literature: Myers 1938.
Code: 12,21,31,41,54,62,64,72,81,93.
Notommata doneta Barring & Myers, 1924
Fig. 233
N. doneta Barring & Myers 1924, p. 448-450, Figs. 22: 1-4.
Type locality: Starvation Lake, Vilas County, WI, U.S.A.
Holotype: Not designated
Description: Elongate body, prominent tail. Two-jointed foot, two characteristic long
and curved toes. Trophus nearly symmetric: long fulcrum, unci have a single well-
developed ventral tooth followed by a second rudimentary tooth crossing the basal
plate diagonally. RCO large, rounded and clear, subcerebral glands large and reach
the anterior part of the sac. The species is distinguished from N. cyrtopus and N.
cyrtopus telmata by the unusually long toes. The smaller (175 and 112-145 r-tm long)
specimens observed by Rodewald (1935) and Pawlowski (1938), respectively, have
short toes (16-17 and 20 ~m). It seems doubtful whether they can be assigned to N.
doneta and agree better with N. cyrtopus.
Total length 250-300 ~m, toes 45-50 r-tm, trophus 40 ~m.
Ecology: Cosmopolitan but rare species.
Literature: Rodewald 1935, Pawlowski 1938, Koste & Shiel 1991.
Code: 13,21,32,41,54,62,64,72,81,93.
Notommata endoxa Myers, 1933
Fig. 234
N. endoxa Myers 1933a, p. 2-3, Fig. 1.
Type locality: Mount Desert Island, ME, U.S.A.
Holotype: Not designated
Description: Large, slender, tapering body, foot short, two-jointed, tail one small
rounded lobe, toes small ending in papillose tips. Corona ventral, small auricles,
retrocerebral sack long, opaque, large cerebral eyespot. Bladder absent, is only an
expansion of the cloaca. Trophus robust, asymmetric, fulcrum long, Y-shaped in
cross-section; rami strongly curved longitudinally, left alula stronger than right; left
manubrium also stronger and longer. Two supporting rods in the wall of the mastax.
Trophus structure and retrocerebral organ are similar to those of N. pachyura, but N.
endoxa differs by the shape of the body, smaller toes, smaller tail and auricles, and the
retrocerebral sac is never clear.
Total length 530-650 ~m, toes 30-40 r-tm, trophus 60 ~m.
Ecology: Widely distributed in the U.S.A., probably acidophilic. Also found in Ma-
185
laysia but no ecological data were mentioned.
Literature: Fernando & Zankai 1981.
Code: 11,22,31,42,56,61,72,81,93.
Notommata falcinella Barring & Myers, 1922
Fig. 235
N.falcinella Barring & Myers 1922 p. 570-571, Figs. 45:8-12
Type locality: Furnace Creek near Baltimore, MD. U.S.A.
Holotype: AMNB 384
Description: Closely related to N. endoxa but easily differentiated by the small
recurved spur at the base of the toes, and the three-lobed tail (a large median lobe and
two small lateral lobes). At the base of the indentation separating the lobes there is a
minute additional lobe. The tail is less broad than in N. collaris and N. pachyura.
Wulfert (1940), notes the existence of the mastax rods and also describes dark and
rough oral and pleural plates above the mastax in European specimens that seem to
have a role in grinding food. BerziI).s (1949b) has described N. falcinella europea
from Sweden, a subspecies (?) of similar size with slightly larger trophi (85 Ilm) and
without mastax-rods.
Total length 500-550 Ilm, toes 30-33 r-tm, trophus 75 Ilm.
Ecology: Europe, North America, rare, collected from acid waters among submerged
Sphagnum. §rOJ;;( I Kosr€: 11.000 (~5.)
Literature: Wulfert 1940, BerziJ}.s 1949b, Kutikova 1962.
Code: 11,22,31,42,56,61,71,81,92.
Notommata fasciola Myers, 1933
Fig. 236
N.fasciola Myers 1933a, p. 11-12, Fig. 6.
Type locality: Mount Desert Island, ME, U.S.A.
Holotype: AMNB 479
Description: Body slender and cylindrical; distal joint of the foot long and wrinkled;
compressed dorsally, stout and lanceolate laterally. The trophus resembles that of N.
pseudocerberus; the fulcrum is long and slender without posterior enlargment; rami
triangular in ventral view. The right uncus has three teeth; the left uncus has four
teeth. RCO small, rounded and ductless, the subcerebral glands are very long.
Total length 163-2175 r-tm, toes 13-16 Ilm. .b~a!~f (vlnjr~,:Lo.fJO, INtJ.)
Ecology: Acidophilic, in Sphagnum.
Literature: Not seen since discovery.
Code: 13,21,34,43,54,62,72,81,94.
 ::;::...::; I.. ), .'~I.,i'l

186 187
,I 1/

(Se
d er ~
A
236
c

~
9 B ;
-..,.
A.
..:'
~~\:.
. ..
..:,,:' . 2
Fig. 36, NotommatafascLO
I hura Group A' 23 7
. la . (f): resting egg,'( g )'male;Gro
.
up B: (b): male. ( 236', Myers
1933a' 237 A:

g~.
~ ,
Fig. 237. N. Wulfert 1935» . (g)

~j~
Donner 1954, . B' Wulfert, (I): fulcrum,

<-:- 235. '. Notommata

Fig. fa lcinella
(h,i): epipharynx,. U) pleural
Group rods.
A'. (235 A:
. Hamng
Hamng &.Myers;
& Myers 1922," B'. Wulfert 1940).
Group
manubna,

235
 188 189

Notommata gZvphura Wulfert, 1935

Fig. 237

~
N. gZ'vphura Wulfert 1935, p. 590-591, Fig. 7.

Type locality: Common in the Merseburg area, Gennany.

Holotype: Not designated

Description: Moderate size, straight and short toes and a characteristically notched d
tail seen only in N. cerberus. It is easily distinguished from that species by the ReO
and the mastax. Although variable in size, the opaque retrocerebral sac is filled with
dark bacteroids and often extends beyond the end of the mastax; the subcerebral
glands are well developed and are attached to the sac. The asymmetric trophus is ro-
bust; rami with teeth, the left alula is very large and dagger shaped. The unci bear a
moderately strong ventral tooth that can have three tiny side teeth (Wulfert 1943) fol-
9
lowed by three reduced teeth. The parthenogenetic egg has a thin shell carrying nu-
merous short spines (Pourriot 1958, Koste, 1976). The male was described by Donner
(1954) and Koste (1976).
Total length 320-450 /lm, toes 20-24 /lm, trophus 60-70 Ilm, parthenogenetic egg
145xl15 lLm, male 280 Ilm.
Ecology: Common and eurytopic in Europe, reported from West-Africa (De Ridder,
1983). This predator feeds mainly on small benthic bdelloids, or sucks out eggs or
carrion.
Literature: See text.
Code: 12,21,31,43,54,61,72,81,92.

Notommata groenlandica Bergendal, 1892

Fig. 238

N groenlandica Bergendal 1892 p. 56, Table 2, Fig.3 :21

Type locality: J acobshaven, Greenland.

Holotype: Not designated

Description: Pseudosegmented slender body, small auricles, bulging corona. Foot

240
has one joint or two undistinct joints and bears a papilla at the distal end, between the
two short toes. Trophus asymmetric: left uncus has four teeth, the right uncus, three 239
teeth. The manubria bear a curious wing-shaped lamella strenghtened on its sides. No
bladder; RCO opaque, the two clear subcerebral glands reach the median part of the
sac.
Fig. 238. Notommata groenlandica (e): unci; (t): manubrium; (g) photo of swimming female.
Total length 180-240 Ilm, toes 9-11 /lm, trophus 30 /lill.
Fig. 239. N. haueri.
Ecology: Acidophilic, Europe. Fig. 240. N. lenis. (238: Wulfert 1940, photo Koste; 239: Wulfert 1939; 240: Harring & Myers 1922).
Literature: Wulfert 1940, Koste 1969, Francez 1981.
Code: 13,22, 32,43,54,61, 647, 71, 82, 93.
 190 191

Notommata haueri Wulfert, 1939

Fig. 239

N. haueri Wulfert 1939, p. 70., Figs. 3:a-e.

Type locality: River Spree, Germany.

Holotype: Not designated

Description: Body elongate., tail of variable length, rounded; two-jointed foot with
straight and short toes; large auric1es. Mastax robust (70 !lm); rami strongly asymmet-
ric with a larger left alula; unci have six teeth. Related to N. cerberus (see Wulfert
1940) but the rounded tail without median notch is diagnostic. See also N. glyphura
(Fig. 237) and N. galena (Fig. 226 B).
Total length 300-500 !lm, toes 16-22 !lm, trophus 70 !lm.
Ecology: In periphyton, alkaline pH.
Literature: see text and Donner (1972).
Code: 12,21,31,43,54,61,72,81,92.

~~
Notommata lenis Harring and Myers, 1922
Fig. 240

N. lenis Harring & Myers 1922 p. 586-587, Figs. 50:9-13.

Type locality: "Common in weedy ponds". 241

Holotype: Not designated

c~
Description: Body slender and elongate, abdomen tapering posteriorly, ending in a
three-lobed tail. Foot has two joints, the posterior divided in two parts ("fingers")
bearing the short toes. The most specific character is the double dorsal antenna. The
two tufts of setae are about 10 ~m apart and united by a slender muscle. Trophus
somewhat asymmetric, rami roughly semicircular without alulae. Unci have five to
six teeth, with two or three teeth larger than the others. ReO consist of a very large,
clear and nearly circular sac without subcerebral glands.
The morphology of the male is similar to the female, also with a double dorsal an-
tenna. Resting eggs also observed in culture, having a spiny shell (Pourriot, 1966).
Ecology: No data. While designated common by Harring & Myers, it seems to be
rare, known only from the USA and France.
Literature: See text.
Code: 12,21,33,43,56,61,64,72,81,93.

= ()( fO{/I~/~a. PO(/((IO/, /1Q') .~ ICt)/ ~fJ.J:

Notommata longzna (Wulfert), 1961 nom. novo
Fig. 241

N. cerberus var. longinus Wulfert 1961 bp. 89-90, Fig. 22. 242
Fig. 241. Notommata longina.
Type locality: Diiben Moor, Germany. Fig. 242. N. mera (c): ReO; (f): mastax. (241: Wulfert 1961; 242: Donner 1940).
 193
192

:" ,~

m·.
t

243
Fig. 243. Notommata omentata (e): unci, (t) mastax with salivary glands. (243: Wulfert 1939a).

Holotype: Not designated

Description: Body shape variable but usually long and narrow. Neck distinct,
auricles short and round. Foot two-jointed, tails characteristic: it consists of a wide
anterior part folded in like a hat, and a distal lobe bent over half of the second foot
joint. Toes conical. RCO long and narrow, covering mastax. Trophus symmetrical,
inner edge of rami denticulate, unci with seven and eight teeth respectively, the mid-
dle one being the longest. Manubria lamellate. Specific status has been recommended
already by Wulfert (1961) and by Koste & Shiel (1991). 244
Total length 350-430 !lm, toes 18-24 !lm, trophus 42 !lm.
Ecology: Acidophilic, found in Germany and Tasmania, rare. Fig. 244. Notommata pachyura (a-e): N.p. pachyura; (f,g): N.p. triangulata; (h,i): N. p. spinata.(244: (a-g)
Literature: see text. Harring & Myers 1922, (h,i) Koste 1991).
Code: 12,21, 31,42,53,62, 72, 81, 92.
small conical tail small. RCO well developed, eye present. Trophus with delicate
rami ~arrying iong diagonal alulae, manubria bent; trophus figures and description in-
Notommata mera Donner, 1970 complete. Could, perhaps, be transfered to Pleurata n.gen. after further study.
Fig. 242
Total length 165 !lm, toes 8 !lm, trophus 26 !lm.
N. mera Donner 1970, p. 231-232, Fig. 17.
Ecology: In mosses, debris.
Literature: Not seen since discovery.
Code: 13,21,31,43, 57?, 61, 72,81, 94?
Type locality: River Salzach, Austria.
Holotype: Not designated

Description: Body club-shaped, widest at head; striated. Foot small, single joint, toes
194 195

Notommata omentata Wulfert, 1939

Fig. 243

N. omentata Wulfert 1939a, p. 612, Figs. 13 a-d.

Type locality: Ditch near Merseburg, Germany.

Holotype: Not designated

Description: Body cylindrical with pseudosegmentation, head broad, corona bulging

ventrally; no auricles. A two-jointed foot bearing two conical, straight and short toes.
Trophus slightly asymmetric: rami without alulae, the unci have 2/3 and 3/4 teeth, re-
spectively. No bladder, replaced by a cylindrical organ. ReO small and black, the
subcerebral glands are very unusual, much longer than the sac, a specific character.
Ganglion contains two crystals; large red eyespot.
Total length 200-230 [lm, toes 10-14 [lm, trophus 35-48 [lm.
Ecology: Alkaline and acidic waters, Europe.
Literature: Wulfert 1960b, Koch-Althaus 1963, Francez 1981.
Code: 13,22,31,43,54,61,72,81,93 or 94.

Notommata pachyura pachyura (Gosse), 1886

Fig. 244
Copeus pachyurus Gosse, Hudson and Gosse 1886 p.31
Notommatapachyura Harring and Myers 1922 p. 565-567, Figs. 42:1,2 & 43:1-5.
nec N. quinquelobatus Stokes 1896.
Type locality: Sandhurst, England.
Holotype: Not designated
Description: Body pear-shaped, lobed tail covers first foot joint. Small papilla be-
tween toes. Auricles large, post-oral chin prominent. Retrocerebral sac extremely
long, subcerebral glands small. Trophus asymmetric, fulcrum long, posterior cross-
section Y-shaped. Right ramus with broad tooth, unci with 1+4 and 1+3 teeth respec-
tively. Two sigmoid pleural rods and an oral plate present. Closely related to N.
collaris, distinguished by the much larger tail, the longer toes, the less robust trophus
and some details in the mastax (number of unci teeth). There is confusion regarding
the identity of this species. For example, in his "N. pachyura ", Wulfert (1956) ob-
served a bladder, normally absent in the animal described by Harring and Myers. A
bladder is also visible on the figure of Donner (1954) where the size (350-560 [lm),
the external morphology and the mastax are those of N. allantois. On the basis of the
presence/absence of a well-differenciated bladder, an important taxonomic character
often neglected, N pachyura can be confused with N. collaris but not with N.
allantois, a more common and quite different species (see below). Because of this
confusions and because the drawings are often reproduced from Harring and Myers,
the presence of the true N. pachyura in the fauna of Roumania (Rudescu, 1960), of
Czechoslovakia (Bartos 1959) and URSS (Kutikova, 1970) must be reviewed. There
245
Fig. 245. Notommata pachyura spinata semi-contracted, dorsally. (245: photo Koste)
is also some doubt about the specific identity of the male described by Dixon-Nuttall
(1894) and by Wesenberg-Lund (1923).
Total length 300-800 [lm, toes 35-75 [lm, trophus 75-102 [l~; m~le 300 [lm. I:I0w-
ever, these measurements are subject to the same caveat as the IdentIty of some lItera-
ture data.
Ecology: Cosmopolitan, acidophil, eats desmids, occasional predator.
Literature: see text.
Code: 11,22,31,41,53,61,71,81,92.
Notommata pachyura "form" triangulata (Kirkman), 1906
Fig. 244 f, g
Copeus triangulatus Kirkman 1906, p. 264, Figs. 12:1,2
N. brachiata Daday 1910, p. 67, Fig. 2.
Description: "Winged" forms which develop trian?u1ar humps, t:w 0 late~al and one
dorsal appear to be ecotypes of N. pachyura. Descnbed from tropIcal habItats: South
Afric~ by Kirkman (1906), from East-Africa by Daday (1910) and by ~oste (l981~ as
N. pachyura form triangularis (Australia), as well as from North Amenca by Harrmg
& Myers 1922, these large forms (600 to 810 [lm) appear to lack a bladder. A com-
plete series of intermediates between the normal humpless fo~and the humped
forms, which appear in warm waters, has been observe~ by Harnng (1913). Koste
1981 suggests that a predator substance may play a role m the development of these
 196
A tough mucilaginous sheath. No auricles, only small ciliated appendages; square head
B Notommata placida Harring & Myers, 1922
246ffl 247
Fig. 246. Notommata peridia Group A: without sheath; Group B: with mucilaginous sheath.
Fig. 247. N. placida. (246 A, 247: Harring & Myers 1922, B: Nogrady 1980).
a:ppe~dages, .sim.ilar to those on Brachionus or Asplanchna. Koste (personal commu-
mcatlOn) mamtams that his Australian forms are not identical with the Kirkman taxon'
Koste & Shiel (1991) suggest specific status under the name N spinata (non Copeu;
197
triangulatus Kirkman). Until further studies we assign subspecific status to Noto-
mmata pachyura spinata Koste & Shiel 1991, (Fig. 244 h) based on trophi, bladder,
and distal tooth on the hump.
Total length 600-720 ~m, toes 58-60 ~m, trophus 140 ~m (Australian specimens).
Literature: See text.
Notommata peridia Harring & Myers, 1922
Fig. 246
N. peridia Harring & Myers 1922, p. 576-578, Figs. 48:1-5.
Type locality: Oceanville, NJ and Wheeler Lake, Vilas County, WI, D.S.A.
Holotype: AMNH 388
Description: Body fusiform or cylindrical, elongate and slender, surrounded by a
with chin. Lateral antennae long, tubular, with a broad base; they end in long setae
emerging from the sheath (see N. copeus). Tail forms a blunt rounded projection and
carries two little knobs on the dorsal side. Foot with two joints, toes tapering and
recurved. Absence of auricles and feeble development of the corona make the animal
unable to swim. Gastric glands mentioned by Harring and Myers were not observed
by Nogrady (1980). RCO very large, disc-shaped, subcerebral glands short, reaching
the level of the eyespot. Trophus somewhat asymmetric. The unci have 4-5 teeth; one
(left uncus) or two (right uncus) ventral teeth are much larger; a rudimentary pre-
uncial tooth is present.
Total length 300-520 ~m, toes 25-32 ~m, trophus 65-70 ~m, lateral antennae 35-37
~m.
Ecology: A rare species known only from North and South America, collected among
the vegetation of acid swamps and lakes.
Literature: Nogrady 1980, Reid & Turner 1988.
Code: 12,21,31,42,54,61,64, 72?, 81,94.
Fig. 247
N. placida Harring & Myers 1922, p. 487-489, Figs. 42:3-6
Type locality: Keni1worth, DC., U.S.A.
Holotype: Not designated
Description: Body of this small and rare species is fusiform and slender. Foot has a
single stout joint and bears two short conical toes. Trophus appears symmetric: ful-
crum relatively short, unci have five (right) or four (left) teeth. The bladder appears to
be fonned by an expansion of the cloaca. RCO well developed.
Total length 130 ~m, toes 9 ~ln, trophus 30 ~m.
Ecology: Rare in weedy ponds.
198 199

e
)(

248~~
e

249
1?
Fig. 249. Notommata pseudocerberus. (249: de Beauchamp 1907, Harring & Myers 1922).

Literature: Not seen since discovery.

Code: 13,22,32,43,54,62,72,81,93.
Fig. 248. Notommata pro-
dota (d): unci; (e): apical Notommata prodota Myers, 1933
teeth of rami. Fig. 248
Fig. 250. N pygmea.
(248: Myers 1933a; 250: N. prodota Myers 1933a, p. 10-11, Fig. 5.
Harring & Myers 1922).

250
200 201

Type locality: Round Pond, Mount Desert Island, ME, U.S.A.

Holotype: Not designated

Description: Elongate, slender body, nearly cylindrical with a small round tail and
two characteristic tapering toes. Mastax appears slightly asymmetric. Rami with un-
equal rows of teeth as well as 3-4 rounded protuberances at the tip; right uncus has
two large subequal teeth and the left uncus only one large tooth. RCO small and clear,
no subcerebral glands. Bladder normal.
Total length 355 /-lm, toes 35 ~m.
Ecology: Rare, acidophilic.
Literature: Not seen since discovery.
Code: 12,21,33,42,54,61, 72,81, 93? 252
jqoz
Notommata pseudocerberus de Beauchamp, 1907
Fig. 249
@3~
6
N. cerberus de Beauchamp 1907 p. 90)': Figs. 1-3.
nec N. cerberus Gosse 1886. b
N. pseudocerberus de Beauchamp 1908, p. 400.

Type locality: Several ponds at Chaville near Paris, France.

Holotype: Not designated

Description: Body slender, fusiform, longitudinally striated; long two-jointed foot

with two straight toes; a rounded tail with a median lobe and two small lateral lobes.
Corona with rostrum and chin, moderately long auricles. I-r0phus very slender and
symmetric; Fulcrum long and thin, rami lamellar(withou.9teeth, unci with a simple
slender tooth. Salivary glands are vestigial. Retrocerebral sac granulated, subcerebral
glands well developed, as long as the sac.
Total length 400-680 ~m, toes 30-35 ~m, trophus 45-56 ~m.
Ecology: Cosmopolitan but rare, preys on sessile ciliates.
Literature: Harring & Myers 1922, Donner 1954, Wulfert 1956, Koste 1969, 1976,
Koste & Shiel 1991. 7* set-or'! row ~ C-O L'l "66..fkete
Code: 11, 22, 31, 42, 56, 62, 72, 81, 92. P(J~J I -"J fl .:Jl.I..4.hU i

Notommata pygmaea Harring & Myers, 1922

253 9
Fig. 250
Fig. 251. Notommata rugosa.
Fig. 252. N. saccigera.
N. pygmaea Harring & Myers 1922, p. 593-594, Figs. 56:5-8 Fig. 253. N. silpha (e): incus laterally; (t): incus dorsally. (251: Myers 1933a; 252,253: Harring & Myers
1922).
Type locality: Lenape Lake, NJ, D.S.A.
Holotype: AMNH 500
Total length 100 ~m, toes 8 ~m, trophus 22 /-lm.
Description: Body of this very small species is stout, spindle-shaped. Foot has two Ecology: Rare, acidophilic?
joints and bears two short toes abruptly reduced to slender tips. The asymmetric Literature: Not seen since discovery.
trophus has triangular rami with alulae and a slender fulcrum and manubria; unci have Code: 13,21,31,43,54,61, 72, 81,93.
a single tooth. RCO well developed.
 202 203

Notommata rogosa Myers, 1933

Fig. 251

N. rogosa Myers 1933a, p. 13-14, Fig. 7.

Type locality: Jordan Mountain Pool, Mount Desert Island, ME, U.S.A.
Holotype: Not designated

Description: Body cylindrical, slender, tapering and the shape changing constantly;
integument ringed. Foot and toes short. Mastax mainly prehensile and has large sali-
vary glands. Rami appear elongate with large symmetric alulae; each uncus has only
one slender tooth. The straight manubria have circular basal plates. No mention of a
bladder in the description and the figure is not clear about it. RCO rounded and clear,
a very small eyespot. The species has affinities to Pleurotrocha (one uncus tooth, al-
most straight incus, salivary glands).
Total length 180 !lm, toes 15 !-lm.
Ecology: Acidophilic, rare.
Literature: Not seen since discovery.
Code: 13, 22?, 33,43, 5~62,72, 82,92. 254 255
Notommata saccigera Ehrenberg, 1832
Fig. 252

N. saccigera Ehrenberg 1832, p. 133, 1838 p. 434, Fig. 50:8.

Type locality: Berlin, Germany.

Holotype: Not designated

Description: Body short and stout, falling off abruptly to the very small tail. Foot
short with two minute toes. Integument is brown, leathery, opaque. Mastax robust
with asymmetric trophus, which has completely lost the pumping action. Fulcrum a
short and broad plate; ventral tooth of the left uncus is very large, followed by three
rudimentary teeth; the right uncus has a large ventral tooth followed by two other teeth
nearly as large and a third small tooth. RCO is large, granulated at the margin and has
a long, slender, and tubular duct. The subcerebral glands are short.
Total length 325-350 !-lm, toes 12-14 !-lm, trophus 75 !-lm.
Ecology: This cosmopolitan species is common in marshes on Utricularia and in sub-
merged or very wet Sphagnum.
Literature: Hudson and Gosse (1886, p.24, Fig. 17; according to Harring & Myers
this is a different taxon), Harring & Myers 1922, Kutikova 1970.
Code: 12,22,31,43,54,61,72,81,93.
256
Fig. 254. Notommata silphoides (b): corona and mastax ventrally; (d) trophus apically; (e) trophus later-
ally.
Notommata silpha (Gosse), 1887 Fig. 255. N. stitista. .
Fig. 253 Fig. 256. N. thopica. (254: de Beauchamp 1923; 255: Myers 1937; Barnng & Myers 1924).

Diglena silpha Gosse 1887b, p. 2, Fig. 1:2.

N.jorcipata Gosse, Hudson and Gosse 18862:23, Fig. 18:1.
nec N. jorcipata Ehr. 1838
204 205

Notommata silpha Harring 1913 p. 79.

Type locality: Ireland.

Holotype: Not designated

Description: Slender, elongate body, posteriorly reduced and wrinkled; short foot
appears to have only a single joint. Toes short and slightly curved; no auricles. Mastax
modified virgate type, without pumping function; the robust and strongly asymmetric
trophus has become adapted to prehension. Fulcrum short, rami asymmetric, right one
with hooked alula. Each uncus has three functionnal teeth. Manubria broadly triangu-
lar with a hood-like process anteriorly. A clear vacuolate sac is only present with a
large ganglion.
Total length 160-200 /lm, toes 8-10 Ilm, trophus 24 /lm.
Ecology: In acid and dystrophic waters, rare.
Literature: Harring & Myers 1922.
Code: 13,21,33,43,57,61,71,82,94.

Notommata silphoides De Beauchamp, 1923

Fig. 254

N. silphoides de Beauchamp 1923, p. 221-223 Fig. p. 223.

Type locality: Canal de Bourgogne, Dijon, France.

Holotype: Not designated

Description: Body elongate, without foot but with two pincer-shaped small toes.
Trophus very small: uncus and ramus are joined in an unique plate; fulcnlm and the
manubria are three slender curved rods. RCO well developed but there is no eye. Fig. 257. Notommata tripus (g): feeding. (257: Harring & Myers 1922, Wu1fert 1939).
Total length 300 /lm, toes 17 /lm, ttophus 50 Ilm.
Ecology: No data.
Literature: Not seen since discovery. Total length 168/lm, toes 13.
Code: 13, 21, 31,43, 57, 63, 73, 83, 94. Ecology: Acidophilic, fairly common at type locality.
Literature: Not seen since discovery.
Code: 13,21,33,43,54,62,72,82,92.
Notommata stitista Myers, 1937
Fig. 255
Notommata thopica Harring & Myers, 1924
N. stitista Myers 1937, p. 7-8, Figs. 2, 5, 8. Fig. 256

Type locality: Caroga Lake, Adirondack Mountains, NY, D.S.A.
Holotype: AMNH 873
Description: Body short, gibbous, and stout, with small toes. Trophus slender, sym-
metric: each uncus has three teeth; manubria with inner truncate lamella. RCO small
and pyrifonn, eyespot present. Stomach invariably filled with Chlorella. This species
is similar to N fasciola but distinguished by the stouter body and the absence of
subcerebral glands.
N. thopica Harring & Myers 1924, p. 446-448, Figs. 22:5-9.
Type locality: Philadelphia, PA, D.S.A.
Holotype: Not designated
Description: Body fusifonn, flexible with longitudinal striae; foot with two short
joints, short tail. ReO very long, subcerebral glands contain very characteristic dark
globular aggregation of bacteroids. Mastax slightly asymmetric, rami with blunt teeth
206 207

and a sharp alula on the right. Unci asymetric with one principal tooth and a series of
serration-like auxiliary teeth (3 and 5). Manubria almost straight with large basal
plates. Closely related to N. aurita, (epaxia), codonella.
Total length 250-300 !lm, toes 15-18 !lm, trophus 36 p.m.
Ecology: Unknown, common near Atlantic City.
Literature: Not seen since discovery.
Code: 13,21,31,43,54,61,72,81,93.

Notommata tripus Ehrenberg, 1838

Fig. 257

N. tripus Ehrenberg 1838 p. 434, Fig. 1:4.

N. pi/arius Gosse, Hudson & Gosse 1886 p. 2:23, Fig. 17:5.

Type locality: Berlin.

Holotype: Not designated

Description: Body short and broad, head and neck a single segment, rudimentary
foot. The rounded tail has a very large spur dorsally, with a knob-like base; this is a
specific character. Toes robust and conical. Trophus asymmetric: left alula broad and
strongly curved; fulcrum long and slender. Ventral tooth of each uncus larger than the
following teeth (one or two); left uncus has a pre-uncial tooth. RCO short and opaque,
ganglion large with the eyespot at his posterior end. No subcerebral glands.
Total length 150-200 !lm, toes 16-20 !lm, trophus 26-30 !lm, caudal process 16-22
!-tm .
Ecology: Cosmopolitan, common.
Literature: Harring & Myers 1922, Wulfert 1939, Donner 1954, Koste & Shiel1991.
Code: 13,21,33,43,52,61,64,71,82,93.

Notommata venusta Harring & Myers, 1922

Fig. 258

N. venusta Harring & Myers 1922, p. 591-592, Figs. 50:1-4.

Type locality: Bargaintown, NI, U.S.A.

Holotype: AMNH 483 (syntype)

Description: Body moderately stout, tail small and indistinct. Toes short, slender,
straight and tapering. Trophus slender and strongly asymmetric (see rami); fulcrum
without any posterior enlargement. Unci have a prominent ventral tooth followed by a
second smaller tooth; manubria slender with a characteristic very long spur curved
inwards from the basal plate and attached to therami. There is a second rod attached to
the manubria and rami. RCO small and pyriform, no subcerebral glands; eyespot
Fig. 258. Notommata venusta. small.
Fig. 259. N. voigti Group A: Donner; Group B: Wulfert (258: Harring & Myers 1922, photo Nogrady;
259 A: Donner 1949, B: Wulfert 1960b). .
208 209

a b

260 261
Fig. 261. Notommata gisIeni. (260: Hudson & Gosse 1886, Weber 1918?; 261: B-erzinGs 1949).
Fig. 260. Notommata brachyota.

Literature: Wulfert 1960b, 1961; Koch-Althaus 1963.

Total leng!h 20~-~25 !lm, toes 15 ""m, trophus 24 !lm. Code: 13,21,31,43,57, 61?, 64, 71, 81, 93.
Ecology: ACIdophIlIc? Found at type locality only.
Literature: Not seen since discovery.
Code: 13,21,33,43,57,61,72, 82?, 93. SPECIES INQUIRENDAE

Notommata brachyota Ehrenberg, 1832

Notommata voigti Donner, 1949
Fig. 259
N. voigti Donner 1949, p. 8-10, Fig. 4.
Type locality: Admont, Austria.
Holotype: Not designated
D~scriptio~: .Body saccate, flares posteriorly in swimming position; no tail (?). Foot
w~th three Jomts, ~oes s~all. with papi~la in between. Bladder on left side. Trophus
slIghtly asymmetrIc: ramI WIth long thm alulae, teeth on both sides unci with three
teeth. Related to N aurita which, however, has a median bladder a~d no papilla be-
tween ~he toes (N:B. Wulfert ~1960b) suspects that this papilla has been overlooked in
N aurzta). TrophI show no dIfference, but the males are different.
Totallen¥th 200-210 !lm, toes 10-12 !lm, trophus 30-35 !lm.
Ecology: CIrcumneutral pH occurrence, not rare.
Fig. 260
Description: This small species (total length 170-200 !lm) has a morphology similar
to that of N. groenlandica Bergenda1 and N. weberi Voigt 1957 with a rudimentary
foot carrying two short toes (10 !lm ?) but without tail. Koste (1978) synonymized this
species with N weberi but provided only a tiny figure, no description. The mastax is
insufficiently illustrated in all sources. A bladder can be seen on the figure of
Ehrenberg(?), but is not mentioned in the text; Voigt (1957) specifies a true bladder
("Protonephridialb1ase"). The existence of cerebral glands appears probable (the so-
called "calcareous masses") but that of the sac is doubtful. Most of the literature
descriptions and figures reproduce those of Weber (1898), but all of them are insuffi-
cient, including those of Rudescu (1960). According to Hudson and Gosse (1886),
this species sucks out Nitella.
NOT IN KEY
210 211

Notommata gisIeni Berzins, 1949 03 TOES

Fig. 261
01 base bulbous
N. gisIelli Berzil1s 1949b p. 314-315, Figs. 1-5. 02 base not bulbous
03 claws present
Type locality: Lakes Fiolen and Straken, Aneboda, Sweden.
Holotype: Not designated 04 STOMACH

Description: Small, stout, very transparent body, head well separated, foot wide, 01 walls scalloped
conical, toes medium length. Corona ventral with funnel-shaped pit. Stomach divided 02 walls smooth
into two parts, footglands very large. Eyespot absent. Trophus: fulcrum dorsally bent,
rami asymmetric, left alula longer than right. Left ramus carries a knob on outer edge. 05 SALIVARY GLANDS
Manubria curved, asymmetric wide basal plates, distal ends curved. At the proximal
end two pleural rods. Unci unequal, powerful teeth (number not stated). Trophi re- 01 symmetrical
semble those of N. paracyrtopus. Considerable discrepancies between description and 02 asymmetrical
figures; appears to be a semi-contracted preservation artefact. 03 on one side only
Total length 134-138 ~m, toes 14-15 ~m, trophi 24-25 ~m. 04 rudimentary
Ecology: No data.
Literature: Not seen since description. 06 FULCRUM 07 ROSTRUM

NOT IN KEY 01 distal end normally widened 01 present

02 distal end conspicuously widened 02 absent
03 distal end a knob
Genus Resticula Harring & Myers, 1924

Resticula Harring & Myers 1924, p. 518.

NUMERICAL KEY TO REST/CULA
Type: Furcularia melandocus Gosse 1887 = Resticula melandocus (Gosse).
11,22,31,42,53,61,71 melandocus
Type locality: Woolston, England.
11,22,32,41,53,61,71 lestes
12,21,32,42,54,62,72 gelida
Description: Vermiform or spindle-shaped body, short toes with bulbous mucus res-
12,22,31,33,42,52,61,71 nyssa
ervoir. Corona frontal, with ciliary wreath and two small auricle-like tufts of
12,22,31,33,42,52,61,71 vermiculus
locomotory cilia. Salivary glands present, RCO a small ductless sac; eyespots a loose
12,22,31,42,51,63,71 plicata
aggregate of pigment granules in vacuoles of the sac. Mastax virgate, fulcrum long
12,22,32,42,51,61,72 anceps
and slender; rami triangular with a sharp bend at right angle in the middle, unci with a
single tooth, epipharynx absent or rudimentary. Seven species.
DICHOTOMOUS KEY TO REST/CULA
REST/CULA DESCRIPTORS Eye absent or hidden 2.
1.
Eye present, sometimes in a dispersed form 3.
01 EYES Stomach wall scalloped, base of toe normal lestes
2(1).
Stomach wall smooth, base of toe bulbous melandocus
01 absent Rostrum absent or rudimentary 4.
3(1).
02 present Rostrum present 5.
4(3). Fulcrum conspicuously spatulate, foot >'i4 of total length, rami ..
02 FOOT symmetric gelida
Fulcrum normally spatulate, foot short, rami asymmetric anceps
01 long, > 1/4 total length Salivary gland asymmetric 6.
5(3)
02 short
 212 213

Description: Body fusifonn, longitudinally striated, with rostrum; foot conical,

unsegmented, toes relatively long, slim, blunt. Eyespot absent. Salivary gland small,
on right side only. Stomach wall with lobes and terminal caeca, RCO and duct dark
grey. Trophus delicate, rami with long, thin vertical alulae; on the right ramus three
thin sharp teeth, small salivary gland on right side of mastax. Contracts readily on
slightest provocation.
Total length 320 ~m, toes 25 ~m, trophus 50 ~m.
Ecology: Predator; in bottom sediment.
Literature: Kutikova 1970.
Code: 11,22,32,41,53,61,71.

262 Resticula melandocus (Gosse), 1887

Fig. 263

Furcularia melandocus Gosse 1887a, p. 2, Fig. 1:4.

Eosphora melandocus, Harring & Myers 1922, p. 644-646, Figs. 59:6-10.
263 R.melandocus, Harring & Myers 1924, p. 518.

Type locality: Woolston, England.

Holotype: Not designated

264
Fig. 262. Resticula lestes dorsally, mastax with salivary gland (sp).
Fig. 263. R. rnelandocus (b,c): rami laterally and ventrally; (d): rami from different sides; (e): uncus.
Fig. 264. R. gelida females dorsally and laterally, males dorsally and laterally, toes of male, trophus, ful-
crum. (262,264: Wulfert 1935; 263: Wulfert 1939a; Wulfel1 1935).
Salivary gland symmetric; toe comma-shaped
6(5} Toes slim, with claw; rami with outward curving alulae
Toes conical, inner basal part with cone; rami with large
downward curved alulae
Resticula lestes Wulfert, 1935
Fig. 262
R. lestes Wulfert 1935 p. 596-597, Fig. 12.
Type locality: Ditch near Merseburg, Germany.
Holotype: Not designated
Description: Body variable, fusiform with longitudinal striae; foot 3-4 segmented,
toes bulbous at base, curved. Eyespot (?) just a dark granular mass (or hidden behind
this mass). Single salivary gland on right side ofmastax. Trophus complex: rami trian-
gular with downward pointed alulae, dorsally bent with two large teeth on each side.
Unci with single strong tooth, two preuncial teeth on right uncus, one on left.
Manubria sturdy with scalloped base-plates, fulcrum thin, spatulate. Pleural rods em-
bedded in mastax behind upper end of manubria.
Total length 210-320 ~m, toes 25-37 ~m, trophus 34-42~m.
Ecology: Cosmopolitan but not common; carnivore. Possibly in Australia at pH 3.4-
5.4,22-31 QC, 29-59 ~tS cm-I. However, it has been found at alkaline pH elsewhere.
Literature: Donner 1954, Koste & Shie11991. According to Koste 1978, the species
figured by Harring and Myers 1922 is not R. melandocus.
Code: 11,22,31,42,53,61,71.
plicata
vermiculus Resticula gelida (Harring & Myers), 1922
. Fig. 264
nyssa
Eosphora gelida Harring & Myers 1922, p. 642-644, Figs. 60: 1-6.
R. gelida, Harring & Myers 1924, p. 519.
Type locality: Washington, DC, D.S.A.
Holotype: Not designated
Description: Body shape variable from fusiform to elongate-saccate, even wide; ros-
trum absent (although Donner 1970 shows a small wartlike appendage on the corona);
 214 215

small tail; foot long, well separated from trunk, without segmentation, but faint fold-
ing; toes short with blunt points. Integument sometimes with longitudinal folds or
rows of fine dots. RCO just a small sac, no defined eyespot,just a cluster of red gran-
ules at the end of ganglion. Salivary gland rudimentary (present on occasion? see
Donner 1970). Mastax modified virgate: rami triangular with fenestration, bent at
right angle, striated on inner edge; one terminal tooth on both sides. Unci with single
tooth and three preuncial teeth on left uncus, two on right; a supporting structure con-
nects it to the manubria. Manubria slender with a ventral and triangular dorsal lobe;
fulcrum long, terminally strongly spatulate. Male known (Koste 1976).
Total length 312-600 !lm, toes 17-30 !lm, trophus 45-75 !-lm.
Ecology: Cosmopolitan but rare, raptorial. Originally found in cold water (hence the
name), but also occurred at 17°C.
Literature: Wulfert 1935, Donner 1970, Koste 1976, Koste & Shiel 1991.
Code: 12, 21, 32, 42, 54, 62, 72.

265 Resticula anceps Harring & Myers, 1924

Fig. 265

R. anceps Harring & Myers 1924, p. 519-520, Figs. 38:5-8.

Type locality: Helen Creek, Mamie Lake, Vilas County, WI, U.S.A.
Holotype: AMNH 561

Description: Body spindle-shaped, slender, integument soft, transparent, with ros-

trum. Transverse folds distinct. Foot with two joints, toes short, conical. Salivary
glands large, cerebral eyespot large, with lens. Rami triangular with slightly asymmet-
ric alulae and a complex fenestration. Basal apophysis prominent and broad. The
curved dorsal portion of the rami is striate and obtusely dentate. Unci with one main
tooth and 5-6 rudimentary teeth on the tip of the left uncus, none at the right.
Manubrium gently curved, basal plate somewhat pentagonal; slender curved rods in
A the mastax wall parallel to rami, one on each side. Close to R. nyssa, but differs in the
shape of corona, eyespot and simpler toes.
~.
--,/i Total length 250-300 !-Lm, toes 12-15 !lID, trophus 45 !lm.

B
\'(, 268
Ecology: Rare, in a boggy area in Wisconsin and a river in Poland.
Literature: Pawlowski 1958.
Code: 12,22,32,42,51,61,72.

266 Resticula p!icata Wulfert, 1935

Fig. 266
Fig. 265. Resticula anceps.
Fig. 266. R. plicata Group A: dorsally, laterally, trophus; (a): trophus apically; (b): manubrium; (c): incus
R. plicata Wulfert 1935, p.598-600, Fig. 14.
laterally; (d) head contracted; (e): toes. Group B: head and trophus.
Fig. 267. R. vermiculus dorsally, laterally, head with mastax and salivary glands. Type locality: Ponds and ditches near Halle, Gennany.
Fig. 268. R. nyssa. (265: Harring & Myers 1924; 266 A: Wulfert 1935, B: Koste 1987; 267: Wulfert 1935; Holotype: Not designated
268: Koch-Althaus 1962, Harring & Myers 1924).
Description: Fusifonn body with conspicuous longitudinal folds, which may disap-
216 217

pear in well-fed animals; foot three-segmented or also unsegmented, toes bulbous at Pleural rods attached to rami, ends fanned, fibrillate.
base with acute points ventrally curved, i.e., comma-shaped in lateral view. Corona Total length 300-550 ~m, toes 14-20 ~m, trophus 50-60, male 160 ~m.
ventral, large wide rostrum. Light red cerebral eyespot. Rami curved, blunt alulae, Ecology: Acidophil, cosmopolitan but rare.
two teeth. Unci with two equal teeth, between unci and manubria an intramalleus. Literature: Wulfert 1940, Koste 1976, Koste & Shie11991.
Manubria sturdy, basal plate with two cells; fulcrum with distal knob. The species Code: 12,22,31,33,42,52,61,71.
may exist in several morphs (those described by Wulfert 1935, Wulfert 1961, and
Koste & PoItz 1987).
Total length 163-400 ~m, toes 9-17 ~lm, trophus 12-22 ~tm. Genus Pleurotrocha Ehrenberg, 1830
Ecology: Moves very slowly, predator, necrophage. October, November.
Literature: Wulfert 1961, Koste & Poltz 1987. Pleurotrocha Ehrenberg 1830, p .46.
Code: 12,22,31,42,51,63,71.
Type: Pleurotrocha petromyzon Ehrenberg, 1830
Type locality: Berlin, Germany.
Resticula vermiculus Wulfert, 1935
Fig. 267 Description: Body stout, head and neck well separated. Foot long, cylindrical, with or
without segmentation or pleats, two toes (P. robusta with one toe). Corona slightly
R. vermiculus Wulfert 1935, p. 597-598, Fig. 13. oblique with a circum
apical band and lateral locomotory tufts of cilia. Mastax virgate, trophus simple: long
Type locality: Pasture ditch, Gennany. fulcrum, triangular trophus without teeth (exc. atlantica), unci with 1-3 teeth, manu-
Holotype: Not designated bria with small basal plates. RCO often absent, single eyespot, footglands usually
large with a mucus-reservoir. Twelve species in the literature. As listed below, we
Description: Body vermiform with longitudinal folds as well as cross segmentation: moved several species to Pleurata n.gen.
narrow, collar-like neck, long first trunk segment, three slightly plump posterior seg-
ments. Rostrum present. Foot short, toes slim, short, with claws. Eyespot large, dark
red; two small "lenses" with red rim on ganglion. Mastax white, trophus insufficiently PLEUROTROCHA DESCRIPTORS
described: slightly asymmetrical, rami with long outward curving alulae; salivary
gland asymmetrical. 01 RETROCEREBRALORGAN 04 MANUBRIA
Total length 400-600 ~m, toes 15-20 ~m, trophus 50-66 ~m.
Ecology: Benthic sediment, pH 7.5, Gennany, Roumania. 01 extremely long 01 with irregular enlargement and
Literature: Kutikova 1970. 02 nonnal lamella
Code: 12,22,31,33,42,52,61,71. 03 absent 02 with knob on ventral basal plate
03 ending in a "spoon"
04 undulate without basal plates
Resticula nyssa Harring & Myers, 1924 02 CORONA 05 sigmoidal with small basal plates
Fig. 268 06 asymmetric
01 with large sensory papilla 07 apically crutched
R. nyssa Harring & Myers 1924, p. 521-522, Figs. 38:1-4 02 without papilla 08 smooth

Type locality: Bargaintown, NJ, U.S.A.

Holotype: AMNH 505 03 RAMI 05 UNCI

Description: Body vermiform, striated and pseudosegmented; distal half of trunk 01 with teeth 01 no tooth
swollen in side view; corona ventral with small rostrum; small broad tail. Foot 2-3 02 without teeth 02 one tooth
segmented, toes bulbous with claw. No true eyespot, just a mass of red piglnent gran- 03 two teeth
ules without a capsule. Salivary gland asymmetric. Rami triangular with ± asymmetri- 04 five teeth
cal curved alulae, 3-4 apical teeth; basal apophysis prominent. Unci with two teeth, on
the left uncus five auxiliary teeth. Manubria long, slender, fulcrum long, spatulate.
 218
219

NUMERICAL KEY TO PLEUROTROCHA

11, 22, 32, 48,53 channa

12, 22, 31, 46, 51 atlantica
12,22,32,41,51 chalicodis t
12,22,32,42,53 petromyzon
12,22,32,43,52 vernalis t
12,22,32,44,51 thura t
12,21,32,47,54 hyalina
12,22,32,48,52 altila*
13,22,32,41,52 robusta

269
DICHOTOMOUS KEY TO PLEUROTROCHA

1. Rami with numerous teeth atlantica

Rami without teeth 2.
2(1). Retrocerebral organ extremely long channa
Retrocerebral organ n()nnal length 3.
Retrocerebral,Q:rg?:l1 ?:Q~eIit; one toe robusta
3(2). Manubria with lamella and knob in upper half petromyzon' {l..<
Manubria apically crotched hyalina (
Manubria normal, smooth altila *

t N.B. P. chalicodis, vernalis and thura have been moved to Pleurata n.gen. based on their trophi (see
page 223).
:.' . ::f
\·>"!?f
,
\,:
271 Pleurotrocha atlantica Myers, 1936
270 Fig. 269

P. atlantica Myers 1936 p. 428-429, Figs. 53: 1-4

Fig. 269. Pleurotrocha atlantica dorsally, laterally, photo of syntype, trophus dorsally and laterally Type locality: Brackish ditches, Mullica River, NJ, D.S.A.
uncus. ' Holotype: AMNH 828; 844 (syntype)
Fig. 270. P. channa.
Fig. 271. P. petromyzon dorsally and laterally, trophi according to Harring & Myers and Koste. Scale bars Description: Sturdy transparent animal, large square tail, foot fluted, two short coni-
i~;~)~ 10 ~m. (269: Wulfert 1942, photo Nogrady; 270: Myers 1933b;'271: Harring & Myers 1924, Koste cal toes. Ganglion pink, large red cervical eyespot. Gastric glands simple, round in
Myers' description. According to Wulfert (1942) they are uniquely complex with
pockets and terminal appendices that point caudally and are attached to the body wall.
Trophus asymmetrical, rami with large alulae and numerous large teeth; large curved
apical branch carries teeth on the right side, none on the left. Unci with two and three
220 221

teeth, respectively~ fulcrum long, laterally a wide plank. Manubria also asymmetric,
basal plates relatively large.
Total length 290-300 !lID, toes 18-20!lID, trophus 35 /lm.
Ecology: Marine, apparently widespread.
Literature: Wulfert 1942, Kutikova 1970.
Code: 12, 22, 31, 46, 51.

Pleurotrocha channa Myers, 1933

Fig. 270

P. channa Myers 1933b, p. 5-6, Fig..3.

Type locality: Mount Desert Island, ME, U.S.A.

Holotype: Not designated

Description: Body elongate, slender, tapering uniformly to the toes; flexible and
changing shape. Foot long, two-jointed, toes teardrop-shaped with blunt tips. Corona
almost frontal. Retrocerebral sac extremely long, extending for nearly one-half length 272
of the trunk; ganglion large, saccate, small eyespot. Rami triangular, without teeth,
large pointed alulae. Unci with two clubbed teeth, manubria with very small basal
plates.
273
Total length 152-155 /lm, toes 12-15 !lm.
Ecology: Acidophilic, in Sphagnum.
Literature: Berzilfs 1954/55, Kutikova 1970.
Code: 11, 22, 32, 48, 53.

Pleurotrocha petromyzon Ehrenberg, 1830

Fig. 271

P. petromyzon Ehrenberg 1830, p. 46.

Type locality: Berlin.

Holotype: Not designated
274
Description: Body robust, trunk wider than head, gibbous. Foot two-segmented, toes
conical, footglands as long as foot, with reservoir. No RCO, small cervical eyespot.
Trophus simple, rami triangular with large rounded alulae; fulcrum long, slim, termi- Fig. 272. Pleurotrocha hyalina.
nally curved; manubria sigmoidal with a characteristic ventral basal plate carrying a Fig. 273. P. altila.
knob. Unci triangular with one slim ventral and another rudimentary tooth. Pleural Fig. 274. P. robusta. (272: Wulfert 1939; 273: Myers 1940; 274: Harring & Myers 1~24).
rods present.
Total length 220-480 !lm, toes 20-26 !lm, trophus 30-37 /lm; distinct populations of
dwarf and giant forms exist, probably depending on food availability.
Ecology: Necrophage, also eats sessile ciliates. Population explosions in the fall after
cladoceran populations collapse. Observed in Volvox, Ophridium colonies, snail and
trichopteran egg-masses. Cosmopolitan, common, limnosaprobic, occurs in running,
acid and brackish water as well.
 222 223

Literature: Harring & Myers 1924, Koste 1976, Koste & Shiel 1991. Type locality: Unknown, Ireland.
Code: 12, 22, 32, 42, 53. Holotype: Not designated

Description: Body robust, ovoid, dorsally arched; continuous longitudinal creases

Pleurotrocha hyalina Wulfert, 1939 arching around posterior abdomen. Head well separated, broad; foot long, tapering,
Fig. 272 faintly divided into three segments; single toe tapering abruptly, but two pedal glands
as evidence of toe fusion. Dorsal antenna on knob. Trophus virgate: fulcrum long,
P. h}'alina Wulfert 1939a p. 582, Fig. 10. curved ventrally, carrying a tiny terminal thorn. Rami triangular without teeth,
pointed alulae; according to Hauer 1975 they fold like a book and separate only on
Type locality: In a ditch near Merseburg, Germany.
iooh bending; unci with one clubbed tooth; manubria long, basal plate narrow; epipharynx
(l.-, /j'·X
Holotype: Not designated two slender curved rods. No RCO, single eyespot with lens.
Total length 180-220 ~m, foot 40-42 ~lm, toes 16-18 ~m.
Description: Head and trunk cylindrical, occasionally longitudinally striated; foot Ecology: Rare to frequent in acid waters (Ireland, North America, Gennany), but also
smooth with protruding bulges, toes long, conical. Corona frontal, bulging, with large from alkaline Neusiedlersee, Austria.
sensory papilla. RCO with globules and faintly visible duct, eyespot on ventral side of Literature: Harring & Myers 1924; Wulfert 1960a, 1960b; Donner 1975.
ganglion. Trophus slender, rami with large pointed alulae, unci with five teeth each;
k.
manubria straight, crutched apically. Mif/rocodides robustus has been synonymized by Harring & Myers (1924). They
Total length 95-110 ~m, toes 10-14 ~m, trophus 12-14 ~m. stated that only the fused toe - not a generic character - is common with M. chlaena.
Ecology: On alkaline clay sediment, as well as in acidic limnosaprobic fishponds; Furthennore, the trophus is not malleate, it resembles that of P. channa; the RCO is
several locations in Germany. absent. See Wulfert 1960 p. 286 and 1961 p. 89., Donner 1975. Koste 1978 retains
Literature: Berzil}.s 1954/55, Kutikova 1970. both species, just compares P.r. with M.r.
Code: 12, 21, 32, 47, 54.

SPECIES INQUIRENDAE
Pleurotrocha altila Myers, 1940
Fig. 273 Pleurotrocha aurea (Zawadowsky) 1916; Kutikova 1970, Fig. 107
Pleurotrocha elegans Zawadowsky 1926
P. altila Myers 1940, pA-5, Figs. 2:1, 2, 7,8, 11, 16. Pleurotrocha larvarum Wlastow 1956; Kutikova 1970, Fig. 109.
All are insufficiently described, and are not included in the key or figured.
Type locality: Promised Land Lake, Pike Co., PA, U.S.A.
Holotype: AMNH 1115
Genus Pleurata Nogrady & Pourriot novo gen.
Description: Body short, stout, head large, neck not separated, foot short, one joint;
toes very short, conical. RCO very prominent, duct well visible, sac crowded with It has previously been proposed by Harring & Myers (1922), Donner (1975) and
bacteroids. Mastax modified virgate: rami triangular with short alulae, curved ven- Koste et al. (1988) that Notommata tyleri Koste, Shiel & Tan, 1988 together with
trally; unci with single long tooth, manubria slender, recurved at distal tip, with small Pleurotrocha vernalis Wulfert, 1935, P. chalicodis Myers, 1933b, P. thura Myers,
basal plates. 1933b, N tithasa Harring & Myers, 1922, and N. trypeta Harring & Myers, 1922,
Total length 130 ~m, toes 7 ~m, trophus 28 ~m. could be separated from their respective genera and defined as a new genus. Our nu-
Ecology: pH 6.6, common among plants in several lakes of the area. merical analysis corroborated these recommendations, therefore we establish this
Literature: Not seen since discovery. new genus.
Code: 12,22,32,48,52.
Type: Pleurata (Notommata) trypeta CHarring & Myers) 1922
Type locality: Lac Vieux Desert, Vilas County, WI, U.S.A.
Pleurotrocha robusta (Glasscott), 1893
Fig. 274 Description: Corona ± anterior, auricles absent; eyespot (if present) cerebral; RCO
L . and subcerebral glands present; bladder can be present or absent, uncertain in two
Microcodon robustus Glas$cott 1893, p. 40, Fig. jif species. Trophus modified virgate, symmetrical, slender, showing transition to Mono-
Mikrocodides robustus Harring 1913, p. 71. mmata. Rami lyrate, thin (slender triangle in P. trypeta), without teeth; manubria
 224 225

without basal lamella; unci reduced; epipharynx present. The anterior corona and ab- 02 present
sence of auric1es distinguishes this genus from Notommata (with the exception of 03 uncertain
tithasa with a ventral corona) and the very slender rami are different from the rami
seen in Pleurotrocha as well as from those of Notommata. 06 RAMI

01 lyrate
A comparative table of these related species might be helpful: 02 triangular

Species length toes trophus bladder rc sack 07 UNCI

01 small rod
P. chalicodis 130 10 7 + 02 two small rods
N. tyleri 120-139 12-16 19 +7 + small triangular plate
03
P. thura 113-120 7-10 7 + +
08 MANUBRIA
N. trypeta H&M 150 9 16 +
P. (N.) trypeta
01 middle enlarged
(Donner 1975) 152 8-11 19 7 +
02 middle with hole ("handle")
N. tithasa 175-200 25-28 15 + + 03 middle with lamella (plate)
P. vernalis 220 15 23 + + 04 none of the above
05 undulate, no basal plates
06 ending in spoon
PLEURATA DESCRIPTORS
09 EPIPHARYNX
01 EYESPOT
01 absent
01 absent
02 large, semicircular
02 present
03 small, oval
02 TAIL
10 ECOLOGY
01 absent
01 endoparasite
02 short
02 not parasite
03 DISTAL FOOT JOINT
NUMERICAL KEY TO PLEURATA
01 with spur or cilia
02 without these
11,21,32,43,52,62,73,84,93,102 tithasa *
11,22, 32, 42, 53, 62, 71,81,82,92, 102 tyleri *
04 TOES
12, 21, 31, 41, 51, 61, 71, 81, 82, 83, 92, 102 chalicodis *
12,21,31,42,52,61,71,8~ 91,102 vernalis
01 < 10 !lm 04 straight 12,22,32,41,52,62,72,85,91,102 thura *
02 11-20 !lm 05 ventr.b. 12,22,32,41,51,53,62,73,81,82,83,92,101 trypeta
03 >21!lm 06 dors.b.

05 BLADDER

01 absent
 226 227

DICHOTOMOUS KEY TO PLEURA TA

)
1. Eyespot absent 2.
Eyespot present 3.
2(1). Manubrium with large median handle; uncus small rod tyleri*
Manubrium without handle; toes huge tithasa *
a h 3(1 ). Rami lyrate 4.
Rami triangular; endoparasite f1ypeta
4(3). Manubria with median hole; large semicurcular epipharynx chalicodis*
Manubria undulate rods without basal plate thura *

· ·' \ Manubria ending in spoon

t
vernalis
lU~U~4
275

276
c
0
.'
....

d
::
.' ........................•.........•.. ..
Pleurata tithasa (Harring & Myers), 1924
Fig. 275

Notommata tithasa Harring & Myers 1924, p. 450-451, Figs. 23: 1-5.

Type locality: Three Lakes, Oneida Co. WI, U.S.A.

(er JOJft. 5
J)e~d.
f.J.;

Holotype: AJVINH 517 (syntype)

Description: Body elongate, spindle-shaped, slender; head small and triangular in

dorsal view, rostrum large. Foot stout and bears two characteristic long decurved and
incurved toes. Mastax a very simple virgate type, trophus very small. Rami with very
long alulae, no teeth; manubria at almost right angle to the rami. ReO rounded, clear,
277 ductless and situated at the posterior end of the elongate and pyriform ganglion. No
eyespot.
278 Total length 175-200 /lm, toes 25-28 ~m, trophus 15 ~m.
Ecology: Rare, also found in New Jersey and Wisconsin.
Literature: Not seen since description.
Code: 11,21,32,43,52,61,63,73,84,93,102.

If\, Pleurata tyleri (Koste, Shiel, & Tan), 1988

Fig. 276

Notommata tyleri Koste, Shiel & Tan, 1988, p. 125-126, Fig. 15.

279 Type locality: Arthur's Lake, Tasmania.

Holotype: South Australian Museum VA110
280
Description: Described from semi-contracted specimens. Body small, short, stou~,
Fig. 275. Pleurata tithasa dorsally, laterally, trophi dorsally, laterally and frontally. rump rounded; indistinctly segmented foot, very short recurved toes. Trophus modI-
Fig. 276. P. tyleri (a): laterally semicontracted; (b): dorsally; (c): toe laterally; (d): parthenogenetic egg;
(e,t): incus; (g): trophus ventrally; (h): manubrium laterally. Scale bars 50 flm (left) and 10 flm.
fied virgate: rami strongly convex, no denticulation; fulcrum slender, straight;
Fig. 277. P. chalicodis. manubria curved with diagnostic handle-like median structure. Epipharynx consists
Fig. 278. P. vernalis. of two large oval plated behind rami (cf. Pleurata chalicodis). Related to P. trypeta
Fig. 279. P. thura. which is an obligate endoparasite in Gomphosphaeria colonies. The trophus is, how-
Fig. 280. P. trypeta lateraly, dorsally, trophi; (a): trophus dorsally according to Donner. (275: Harring ever, very close (if not identical) to that of P. chalicodis. The very high affinity is also
& Myers 1924; 276: Koste et al. 1988; 277,279: Myers 1933b; 278: Wulfert 1935; 280: Harring &
Myers 1922, Donner 1975).
228 229

born out by the cluster analysis. Unfortunately, both species are single records, and P. Total length 220 [lm, toe 15 !-tm, trophus 23 [lm.
tylerihas never been studied live; thus further comparison and possible synonymisa- Ecology: Benthic, rare.
tion must wait. Literature: Kutikova 1970.
Total length 120-139 !-tm, toes 12-16 [lm, trophus (incus) 19 [lm, parthenogenetic Code: 12,21,31,42,52,61,62,71,86,91,102.
egg 30-45 x 50-65 ~tm.
Ecology: 8°C, pH 7.7, 17 !-tS cm-I, from shallow open water over gravel.
Literature: Not seen since discovery. Pleurata thura (Myers), 1933
Code: 11,22,32,42,53,61,63,71,81,82,92,102. Fig. 279

Pleurotrocha thura Myers 1933b, p.l-2, Fig. 1.

Pleurata chalicodis (Myers), 1933
Fig. 277
Pleurotrocha chalicodis Myers 1933b, p. 3-4, Fig. 2.
Type locality: Long Lake, Mount Desert Island, ME, U.S.A.
Holotype: Not designated
Description: Body short and stout, head unusually large (about one-third total
length), trunk conical; foot short, two-jointed, toes conical with papillose tips. Corona
oblique, gastric glands large, kidney-shaped; clear, round, ductless retrocerebral sac
that encloses the eyespot. Mastax modified virgate; rami slender, lyrate, opening ter-
minally, without teeth. Fulcrum staight, enlarged at posteI10r end, manubria very slen-
der and curved, with an irregular enlargement and small hole in the terminal one-third
with a ventrally projecting lamella. Unci are tiny rods resting on the tip of the rami.
The epipharynx consists of two large, thin, oval plates behind the rami. See remarks
under P. tyleri.
Total length 130 [lm, toes 10 !-tm.
Ecology: Acidophilic, littoral/benthic, also found in New Jersey.
Literature: Not seen since discovery.
Code: 12,21,31,41,51,61,62,71,81,82,83,92, 102.
Pleurata vernalis (Wulfert), 1935
Fig. 278
Pleurotrocha vernalis Wulfert 1935, p. 594, Fig. 9.
Type locality: Pasture ditch, near Merseburg, Germany.
Holotype: Not designated
Description: Body stout, gibbous, foot conical, wrinkled, toes short. Large ganglion
with small, brown RCO. Rami small, lyrate, without teeth but with alulae, right uncus
with tiny single tooth, manubria very characteristic: the first two-thirds is straight, the
rest is bent and carries a terminal lamella that forms a spoon-shaped terminal widen-
ing. Wulfert places it very close to P. trypeta (Harring & Myers) 1922. According to
our analysis it is much closer to P. thura, although the trophi are very different.
Type locality: Mount Desert Island, ME, U.S.A.
Holotype: Not designated
Description: Body short, spindle-shaped, small tail, foot short, toes conical, pointed.
Corona oblique, RCO dark with bacteroids. Trophus virgate, simple: rami at right an-
gle to the fulcrum, lyrate with slender points and large alulae, unci two feeble diverg-
ing rods, manubria undulate without any basal plates. Fulcrum straight and long with
a thin apophysis. Swims in short jumps followed by smooth gliding.
Tota11ength 113-120 [lm, toes 7-10 ~m.
Ecology: Common in acid waters, Wisconsin, New Jersey, U.S.A.
Literature: Not seen since discovery.
Code: 12,22,32,41,52,61,62,72,85,91,102.
Pleurata trypeta (Harring & Myers), 1922
Fig. 280
Notommata trypeta Harring & Myers 1922, p. 602-603, Figs. 50:5-8.
Type locality: Lac Vieux Desert, Vilas Co., WI, D.S.A.
Holotype: AMNH 555
Description: Body short and stout; foot relatively lo?g wi~h two short,. distally
recurved and incurved toes. No auricles. Mastax a modIfied vIrgate type WIth small
trophus. Elongate triangular raIni are capped by an epip~arynx which consists of two
large, thin, triangular lamellae; unci very small; man~b~a very slender and enlarged
at their posterior end with a very small ventrally p01~t~ng. lamella: No bladder. Ac-
cording to Harring & Myers this species was paraSItIc III colomes o~ the cyano-
bacterium Gomphosphaeria, but Donner (who has seen preserved speCImens only)
mentions nothing about that.
Total length 150 ~m, toes 9 [lm, trophus 16 ~m.
Ecology: Endoparasite?
Literature: Donner 1975.
Code: 12,22,32,41,51,53,61,63, 73,81,82, 83, 92, 101.
238

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tion Zwenigorodskoy. Trudy Lab. Eksp. BioI. Moskv. Zooparka 2:261-295. (in Russian).
Bleach 1 C. dorystoma 66, 67, 82
Bupivacaine 1 C. edax 68
Cephalodella 7, 8, 18, 33 C. elegans 68
descriptors 36 C. elongata 46, 70, 105
key 40 C. (elmenteita) 75
types of trophi 34 C. (epitedia) 100
C. ablusa 46 C. euderbyi 70
C. abstrusa 46 C. euknema 71, 108
C. akrobeles 48, 120 C. eunoma 145
C. anebodica 48 C. eupoda 62, 71
C. angusta 49 C. ewynota 72
C. apocolea 49, 12 I C. eva 72, 122
C. arcuata 50 C. evabroedi 74
C. (armata) 56 C. exigua 74
C. asarcia 50 C. fluviatilis 75
C. asta 50, 61 C. fluviatilis minor 75
C. astricta 52 C. forceps 74, 75
C. auriculata 52, 67, 139 C.f01jicata 76, 92, 93, 148
C. belone 53, 108, 126 C. forficata macrura 77
C. (bertonicensis) 127 C. for:ficula 77, 127, 144
C. biungulata 53 C. friebei 77
C. boettgeri 54 C. (galbina) 131
C. (botezati) 56 C. gibba 79, 89,181
C. calosa 54, 66 C. gibba anebodica 79
C. carina 68 ~, C. gibba microdactyla 79
C. catellina 56, 137 C. gibboides 79
C. catellina major 58 C. gigantea 80
C. catellina natans 58 C. gish?ni 80
C. catellina volvocicola 59 C. glandulosa 67, 82
C. celeris 59 C. globata 83,120,122
C. clara 59 C. glypha 83
C. collactea 61, 93 C. gobio 84
C. compacta 61 C. gracilis 85, 120
C. compressa 62 C. gracilis "var." lenticulata 87
C. conjuncta 144 C. gracilis "var." sigmoides 87
C. crassipes 62, n, 137, 142 C. gracilis keta 87
C. (cuneata) 107 C. graciosa 85
C. cyclops 63, 83, 121 C. (harringi) 62
C. (deformis) 127 C. hiulca 87
C. delicata 54, 63 C. hollowdayi 89
C. delicata ':form" eudelicata 64 C. hoodi89,94, 120
C. dentata 64, 111 C. hoodi "var. " grande 90
C. derbyi 144, 145 C. (horni) 90
C. dixon-nuttalli 66 C. hyalina 90, 96
C. (donneri) 62, 142 C. incila61, 90,98, 115
C. dora 66 C. innesi 92
C. dorseyi 66 C. inquilina 93 ,
C. doryphora 67 C. intuta 52, 61, 64,92 C fJ..Ja j~·(OA·&-l.\(S
C. rrisae ww/ f/I'1e~)o.~luI.
 246 247

C. Uakubskii) 62 C. rotunda blJ"ophila 61, 64, 126 Eothinia 8, 152 N. apochaeta 171
C. labiosa 84, 144 C. (sabulosa) 131 key to 153 N. auri/a 171,207,208
C. laisi 94 C. (sagitta) 85 E. argus 155 N. avena 173
C. lepida 94, 109 C. speciosa 126 E. carogaensis 153 N. bennetchi 174
C. (leptodac(vla) 33 C. stenroosi 127 E. elongata 157 N. (brachiata) 195
C. licinia 96, 120 C. stenroosi austriaca 127 E. (elongata macra) 157 N. brachyota 209
C. limosa 61, 98, 124, 140 C. sterea 122, 127 E. eukolpa 155 N. (carpatica) 181
C. lindama:l/a 98 C. sterea dentata 129 E. (lamellata) 157 N. cerberus 174, 188
C. lipara 99 C. sterea exoculis 129 E. (lasiobiotica) 154 N. cerberus "form" parvisa 174
C. macrodac(l/la 48, 99 C. sterea minor 129 E. poitera 157 N. cerberus "var. " longinus) 190
C. marina 100 C. sterea mutata 129 E. triphaea 154 N. cherada 176
C. megalocephala 100, 104, 117 C. sterea serrata 129 Enteroplea lacustris 7, 11 N. codonella 176, 207
C. megalocephala ':/brm" compressa 101 C. (strepta) 48, 99 Glossary 238 N. codonella galena 178
C. megalocephala ':!'orm" rotunda 101 C. (strigosa) 89 Itura 7, 8 N. collaris 178, 185, 194
C. megalotrocha, 10 L 142 C. subsecunda 18, 121, 129 Manfredium 230 N. contorta 169, 178
C. melia 101, 108 C. tachyphora 131, 144, 147 Metadiaschiza trigona 33 N. copeus 181
C. mineri 103 C. tantilIa 131, 132 (Microcodides robustus) 223 N. cyrtopus 174, 181
C. minora 103 C. tantilloides 132 Monommata 8, 14 N. cyrtopus 'form" te/mata 183
C. mira 104 C. tecta 89, 132 descriptors 15 N. cyrtopus paracyrtopus 183, 210
C. mira nidicola 104 C. tempesta 132 dichotomous key 17 N. diasema 183
C. misgurnus 104, 109, 111, 117,118 C. tenuior 61, 93, 98, 124, 125, 134, 140, 142 numerical key 16 N. doneta 184
C. montana 105, 134 C. tenuior "var." pigmentata 96, 134 M. actices 18 N. (epaxia) 173, 207
C. mucosa 105 C. tenuis 135 M. aequalis 18, 22 N. endoxa 184, 185
C. mucronata 106 C. tenuiseta 80, 118, 135 M. aeschyna 20 N. falcinella 174, 185
C. mus 106, 118 C. tenuiseta americana 109, 137 M. appendiculata 20 N. falcinella europea 185
C. (myersi) 56 C. tenuiseta simplex 137 M. arndti 21, 22 N. fasciola 185, 204
C. nana 53, 71, 107, 126 C. theodora 137, 142 M. astia 22 N. (jorcipata) 202
C. nelitis 103, 108 C. tinca 94, 137 M. caeca 22 N. (galena) 178, 190
C. (nodosa) 111 C. tinca "var." conspicua 139 M. caudata 24 N. gisIeni 210
C. obvia 89, 109, 132 C. tincaformis 139 M. dentata 24 N. glyphura 173, 188, 190
C. oxydactyla 54, 96, 109 C. unguitata 146 M. diaphora 24 N. groenlandica 181,209
C. pachydactyla 64, Ill, 117 C. vacuna 52, 146 M. dissimile 29 N. haueri 190
C. pachyodon 85, 105, 109, 111 C. ventripes 83, 96, 137, 139, 144 M. enedra 24 N. lenis 190
C. paggia 89, 112 C. ventripes angustior 140 M. grandis 25 N. longina 190
C. panarista 80, 112 C. vitella 140 M. hyalina 25 N. mera 192
C. papillosa 114 C. vittata 140 M. longiseta 27 N. omentata 194
C. parasitica 101, 114 C. wrighti 142 M. maculata 27 N. (pachyura) 170
C. paxi 92, 115 C. xenica 63, 73, 137, 142 M.phoxa 28 N. pachyura 185, 194
C. paxilIa 115 C. zeteta 105, 144 M. pseudophoxa 28 N. pachyura 'form" triangulata 195
C. pentaplax 105, 111,116 Code 2 M. (robusta) 25 N. pachyura spinata 197
C. pheloma 117 Drilophaga 7, 8 M. (sphagnicola) 22 N. (parvida) 174
C. physalis 67, 83, 96, 118, 120 D. bucephalus 9 M. viridis 29 N. peridia 197
C. planera 118 D. delagei 11 NaOCl1 N. (pilarius) 207
C. plicata 119 D.judayi 9 Narcotic N. placida 197
C. poitera 120 Dorystoma caudata 32 bupivacaine 1 N. prodota 199
C. praelonga 48, 120 D. 'form" furcata 32 procaine 1 N. pseudocerberus 185, 200
C. psammophila 121 Eosphora 8, 158 tricaine 1 N. pygmaea 200
C. pseudeva 121 key to 158 Narcotization 1 N. (quinquelobatus) 170
C. reimanni 63, 83, 122 E. anthadis 159, 162 Notommata 7,8, 164 N. rugosa 202
C. (remanei) 89 E. ehrenbergi 159, 162 descriptors 165 N. saccigera 202
C. retusa 145 E. najas 161 keys 166, 167 N. silpha 169, 202
C. rigida 98, 122 E. therina 161 N. aethis 169 N. silphoides 204
C. rostrum 124 E. thoa 162 N. allantois 170 N. stitista 204
C. rotunda 124 E. thoMes 162 N. angusta 171 N. (spinata) 197
248

N. (le/mata) 181 P. romanica 13

N. thopica 205 P. similis 13
N. tripus 174, 207 Resticula 210
N. venusta 207 descriptors 210
N. voigti 208 keys to 211
N. weberi 209 R. anceps 215
Notommatidae 1 R. gelida 213
key to subfamilies 1 R. lestes 212
Notommatinae R. melandocus 213
descriptors of genera 5 R. nyssa 215, 216
dichotomous key to genera 7 R. plicata 215
numerical key to genera 7 R. vermieulus 216
Observation techniques 1 Repauliana dieerea 5
Paracephalodella lat~rulcrum 147 Rousseletia corniculata 31
Pleurata 223 Scanning electron microscope
descriptors 224 Scaridiidae 230
keys to 225, 227 key to 232
P. chalieodis 227, 228 Scaridium 230
P. thura 229 S. bostjani 234
P. 1.ithasa 227 S. elegans 236
P. trypeta 227, 229 S. grandis 232
P. tyleri 227 S. longicaudum 232
P. vernalis 228 S. longicaudum maculatum 232
Pleurotrocha 217 S. mon/anum 237
descriptors 217 SEM see Scanning Electron Microscope
keys to 219 trophi preparation for 1
P. altila 222 Sphyrias lofuana 14
P. atlanUca 219 Taphrocampa 150
P. aurea 223 key to 150
P. channa 220 T. annulosa 150
P. elegans 223 T. clavigera 152
P. hyalina 222 T. lemurensis 152
P. larval'um 223 T. levinseni 152
P. (parasitica) 114 T. selenura 150
P. petromyzon 220 Tetrasiphon hydroeora 3
P. robusta 222 Tetrasiphoninae 3
Pleurotrochopsis anebodiea 147, 149 Tricaine 1
Procaine 1 Trophi 1,34
Pseudoharringia 13 Tylotroeha monopus 29

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