ROBERTL.

ELLISON
a d COTJN G. OGDICN

Department of Environmental Sciences, l,'niversit,y of Virginh, Cliarlottesville, US-\, n n d British

Jluseum (Satural History), I,oadon, England

A Guide t o the Study and Identification of Fossil Testate Amoebae

in Quaternary Lake Sediments

X,ey words: testate airioebxe, thecamoebians, Qnii ternary, lake, sediments

Abstract

Fossil sliells of testate amoebae, coiiimoiily fourid in Quntermry scdiinonts, provide iuiot,lier key
for the intorpretntion of the recent history of lakes. Outline drnwings and det,ails of shell compo-
sition provide the micropalncontologist with n practical guide to the taxonomy of bhese organisms.

1. Introduction

Tlie value of fossil tcistatc. anioehae in the study of Quaternary lahr stdiiiic~ritsh:is
only hegrin t o be explored. FREY(1964) reviewed tlic status of palaeontological
lininology antl regarded rhizopotls as a group proiiiising valiiable insight into past
lininological conditions. Xlore recently, S C l l O N 1 ~ U B N (1973, 1984) has used testacean
:~sseuiblagesfor deciphering the trophic history of lakcs i n eastern Europe. As well
ils being trophic indicators, some testacean taxa are believed by RUZI(WA (1982) t o
reflect changes in runoff within the catchment of Krottc,nsce (Austria). PATTERSON,
c t rcl. (198.5) have iiscd forariiinifers and testate amoebae to follow thrb transition froiii
niarine t o freshwater a s higher sealevels receded from lake hasins in Sovi1 Scotia ;
furthermore, sonie species niay ref1ec.t cahangcs in the inpiit of clastic sediments
ilrtti orgnnic niutter (SCOTT and XIED~OLI, 1983a).
However, the ~iiicroj~ala~.ontologist interested i n trying to identify fossil testate
anioebae from Quaternary lake sediinrnts will find it difficult for several reasons.
The taxonomy h a s bcen done primarily by zoologists working w i t h living ni:~teriiil
rather than with fossil shclls; n i i i c 4 h of thc classification, therefore, is based on soft
parts such a s the nuclei and pseiido~iodra(see J u m , 1942 and GROSPIETSCII, 1965).
Furtherniore, a larger nuinher of studies have heen conceined with t a s a found in
mils, peat or S p h n g ~ a wantl
~ flowing M fiter than 111th those found in standing \rater.
Lastly, most of the, authoritative nork is published in French or G e r i i ~ n n ,and :IS
BAYEKS (198-1) has pointed out it is often necessary t o refer to several separate inono-
graphs when attempting identifications.
Pioneering stndies on protozoa, inclriding testpte aiiicehae, were iindertal\rn in the
inid-19th century by EHRENBERC, i n Berlin. T a t c r , I J ~ 1 i ) i(1879) asseiiibled n niono-
graph tlercribing freshwater rhizopods found in North ;\iiieiic~a,b u t thr reference still
recognized as th e standard for the taxononiy of there orgwnisins, often called the-
cnnioebians, was conipiled earlier i n this century by the Swiss zoologist YENAEI)(1902).
dlniost siniultaneously, A\-ERI~TZEV ('Russia, l906), S C H O V T E ~ E(6a N itzerliind,
1906) a n d CASH, WAILESand HOPKIKSON (England, 1905-1919) published taxonomic.
treatises o n this group. Subsequently, U E F L A ~ Dand R E DECLOITRE, in F r a n c r , arltl
Hoo~e~ltaau, in the Xetherlarids, iiiade significant contributions to the st tidies,
a n d two (krnians, GROSPIETSCH(1958) a n d HARNISPIT (1958), each published taxo-
nomic keys. An illustrated guide (OGDENa n d HEDLEY,1980) rising 8F:M photo-
tiiicrographs is the most recent coinpilation of useful information. Additional research
using niodern instrunientation has shown that fine structural featnres can be vrry
useful a n d are increasingly heing used in taxonomic desc*riptions,The existing classi-
fications are based on details of the shell structure, cytoplasmic. organization, pseudo-
podia1 displays a n d nioveinerit, with few taxa heing erected on shell niorpholop-
alone.
It is evident froin the inforiliation now heing acciiniulated that the taxonottiy of
this group will require some radical revision. Nevertheless, with the current iritcwst
in shells found in Quaternary lacustrine sediments i n Europe a n d Korth Anierica
(e.g., LAMINCER, 1972; RUZICKA, 1982: SCILOXBORX, 1975, 1984; SCOTT a n d J11cn1-
OLI, 1983a, 1983b), a guide to the tiiore coiiiiuon species found in such deposits is
needed. The present work is offered as a guide for palaeoliiiinologists a n d niicro-
palaeontologists i n the identification of testate amoebae, using readily ohservahle
features.

2. Method of Study

Quaternary lacustrine sediments vary in the degree of consolidation and compaction. I n the sedi-
ments we have studiad, SCHONBORN’S method (1984) of disaggregation by boiling the sample in 3 0;b
potassium hydroxide, seems to be tov severe because many, if not it11 of the testates are destroyed.
We chose t o use a gentler method of soaking the sample in warm detergent, 6 hexametaptios-
phate or ‘Calgon’ will do, for a t least 12 hours. This does not completely disaggregate all t’he claj--
organic colloids but the testates survive nearly unscathed. Following this, or a t the same time,
glutaraldehyde is added t o the silmple t o make a solution of 3 0,’”. This is a n essential st,ep to fix
iind harden the organic cement of t h e shells. The amount of sediment used will vary, depending
on the amount of clay present; in our studies we have used 0.5 t o 1.6 g (wet wt). Initially the
sediment is washed through it pair of 100- and 44-ym sieves; of the screened material, all of the
1100- pm but only part of the 44-100 pm fraction is examined for specimens. Individuals are
identified and picked from each of the samples using a pipette with a n internal diameter of 400-
500 pm, and transferred t o a watchglass. Most of the species can be identified on the basis of
general morphology, under a binocular microscope with a magnification of 50 x or 100 x ; discern-
ment of apertural details and the shapes of siliceous plates requires a greater magnification.
Picked specimens are mounted on glass slides with Naphrax (Canada balsam or Cytoseal)
diluted with xylene so t h a t it can be dropped gently onto t h e slide. Specimens are transferred t o
the slide by pipette, positioned with a single-hair (eyelash) brush, then allowed to dry under a dust
cover. For very fragile shells with so little strength t h a t they collapse 011 drying, PENAKD’S (1907)
method of transferring from wa.ter t o alcohol t o oil of cloves t o the mounting medium may be
advantageous. After the slide has dried, 4-5 drops of the rnoriritant itre placed on t h e slide. Before
adding the cover slip, fragments of cover slip are positioned t o support it a t the corners on a. levcl
above the specimens t o prevent damage. Fragments 100 ym thick are suitable for most specimens.
Enclosed bubbles will clear after a, few hours, but curing of Naphrax and Canada balsam is very
slow unless accelerated b y heating tho slide on a hot plate. Generally, slides should be stored
liorizonblly t o avoid coldflow of specimens in the mounting medium.

3. Taxonomy

The principal purpose of this paper is t o offer the researcher a practic:il guide t o
the taxonoiny of fossil testate amoebae. I n particiilar, the nine plates provide a
visual key for most of the species found preserved in Quaternary lake sediments.
 Jdcntifirat~onof Fossil l'rxtatc .\moebac 641

C h t r o p y x i d s a n d tlifflugiids are the most a h n d a n t tnxa in QuaternaIy ctqmsits,

therefore, figures of these forms and their acconipanyirig descriptions are prcsented
hefore those of the less coninionly found taxa. Some taxa sucah a s Aniyhilrevics,
AssulirLn, Hyalosphmia a n d inost species of Arcclla are not native t o the profundal
environinent, but they can occur i n profundal sediments a s a result of heing trans-
ported froiii littoral habitats. Dimensions used for drawing t h e figures are taken from
several sources, primarily PENARI) (1890, 1902), CASH, WATIM and HOPKINSON
(1905-1919), J ~ F L A N D R E (1929, 19%), C ~ R O S P l E T S C I I (1958), HARKISCFI (1958),
( ~ D E K(1979, 1983) a n d OGDI~CN and HEJ~LEY (1980). For convenience to the usc\r,
the figares in the plates all art. drawn to the same scale; uithin each sequence of
figures they are arranged in order of niininruni length; apertures are shown o n thr
plates with a stippled pattern.
Sumerous workers (e.g., OGDEN, 1980; MEDIOLT, SCOTT a n d ABUOTT,1987) have
ohserved t h a t sollie species vary widely i n their test nrorphology a n d coniposition
(depending upon the availability of particular building iiiaterials), so t h a t identifi-
cation to specific a n d sonietimes generic. level is often drfficult and can he inisleading.
Some taxa, siniilar i n shape a n d structure, may he separated only by exaiiiination
of pseudopodial displays and the ultrastructure of the test ; for example, Dj//Ziigiu
criyulostoma, B. mica a n d D.n~iiiutahave siniilar shapes and coiiiposition, and the
genera Phrygunellu and Cyclopyxis are almost idrntical. I n such instances the iise of
a single name for the species-coi~iplexis acceptable, hut should tw siiitahly qiialified
in the tclxt.

Family Centropyxidae JUNG,1912

(Plates 1 and 2 )
Shell bilaterally symmetrical, outline subcircular or ovoid; shell coinposed of
proteinaceoils matrix with agglutinated mineral grains a n d diatom frustules, niay
hc wholly organic i n the absence of agglutinated particles; aperture sub-terminal or
eccentric, may be invaginated, spinosity partly a function of habitat, spinose indi-
viduals capable of producing non-spinose progeny. One genus - Crntropysc's STEIN,
1859.
Two of the species, C . aerophiln a n d C . plutyatonw, have a shoulder-like feature
associated with t h e aperture. Two other species, C. discoides a n d C . spinosu, have
a n inner wall associated with the aperture; this appears a8 a dark band on eithcr
side of t h e aperture when viewed a t iriagnifications>lOO~.

Family Trigonopyxidae TAEBLICH and TAPYAN,

1964
(Plates 2 a n d 3)
Shell radially symmetrical, circular i n outline; coinposed of proteinneeoils matrix
with aggliitinatecl partivles; aperture central. Two genera- Tripnopyxis PENARU,
1912, in which t h e aperture is irregularly triangular, and Cyclopyxis DEFLAKDRE,
1929, in which the aperture is circular.
Faniily Plagiopyxidac BONNET,
1929
(I'late 3)
shell bilaterally symnietrical, outline subcircular or ovoid; cornposed of protein-
aceoiis inatrix with agglutinated mineral grains; aperture sub-terminal, inva ginated,
elongatt slit. Two genera-Plagiopyxis PERARL), 1910, in which the aperture coinnionly
is concealed, and Bullinulnria (€%NARD, 1907) i n which the aperture is exposed, the
anterior lip of which is perforated.
43 h t . Revue ges. Hydrobiol. 72 (1987) 5
 Fanlily Phryganellidae J ~ N I :1932
,
(Plate 3)
Shell heniisphericd, laterally ronipressed; coniposed of ngglutinw ted particles;
aperture circular, central. Differentiating fossil yltrj-gancllids fiont cyclopj s i r l s is
questionable inasmuch as the priitiary basis for classification is the nature of the
pse II tl opod ia. One gen 11s-P?i rygu n d k YENARD,1 902.
Family Difflugiidae WALLTCI~,
1864
(Plates :3-7)
Shell globose, pyriforru, aciiniinate or ovoid ; coniposed of agglutinated iiiirieral
particles and hard parts of other micro-organisms: apcrtitre circular or ovoitl. Scvcral
genera.
Bifflugiu TJCCLERC, 1815-shell ovoid, pyrifornr or acuniinate; coiiiposecl of agglii-
tiriated grains a n d diatom frustules; aperture terminal, circular or ovoid, niay he
lobed, some with rim o f organic cement.
For convenience, this genus has been divided into four main groups: A) regular
forms; U ) fortiis with collar or short neck; C) forms with distinct neck; a n d D) forms
with aboral horns or spines. Owing to the considerable variability i n niorphology
aniong difflugiids, however, this grouping most he used cautiously. For example,
regular fornis niay exhibit a slight development of a collar; conversely, collais or
necks may he obscured by large grains of adventitious niaterial.
We have adopted the existing taxononiy which for fossil material is based on iiior-
phology, texture and composition of the shell; sizes and shapes are shown in the f i -
gures, whereas synopses of the shell texture and composition are given in the follow-
ing descriptions. The texture (“smooth” or “rough”) is, of course, dependent iipon
the coniposition (angular grains of quartz, fragments of diatoiu frustules). At Iiragnt-
fications >lOOx the texture of the aperture niay be diagnostic: the apertural rini
niay be “irregular”- coniposed of unsorted grains or a mixture of grains and frustules;
it may be ,,siiiooth-“Coiiij)osed of uniformly fine-grained particles; or it niag be
formed of “organic centent”, i n which case the ceiiient also foritis a n even rint.
Unless otherwise indicated in the figures or in the text, cross-sectionsand apertures
are circular.
A. Regular forms of Difflugia (lacking collar or neck)
D. stechlinensis-scattered quartz grains i n organic matrix ( SCIV~NRORK, 1962).
D.pristis-composed of flat grains.
D.anyulostoma-coniposed mostly of diatom f r u s t ides: agertural rim irregular.
D.~ ~ ~ i c a - s u r f a csmooth,
e composed of sniall grains; apertural rint of organic
cement.
D.minutu-composed of unsorted grains; apertural rini of organic celllent.
n.lucida-laterally compressed ; surface smooth, composed of sniall grains : aper-
tural rim irregular.
D, clchloru-composed of siliall grains a n d diatom frustules; apertural rim irregular.
D.grarnrn-composed of unsorted, smooth grains ; apertural rim irregular.
D.urr~2~uZullu-siirfacesmooth, cwmposed of small grains; aperture. with slna11
collar.
D..rr~anicuta-surface rough, composed of angular grains ; aperturc surronndrtl b y
sinall grains.
D.tenuis-composed of angular grains ; apertural rim irregular.
D. glans-surface smooth, composed of unsorted grains.
D . decloitrei-surface smooth, coiiiposed of flat grains; aperture with suggestion
of sniall collar.
 Tdentif~catioiiof Fossil Testate .hioebae 643

/,). !/(ohu/os*-coniposed of large grains a n d few diatotii f r i ~ t i i l e s ;a ~ ) e r t u r a riiii

I
irregular.
U . lernuni-coinposed of grains :inti sttiall siliceous plates, often with scattcrrng of
additional large grains.
11. I/icustri.~-cotiiQose[~ of sniall grains a n d diatoni frustirles.
D./ r w i d t ri-surface smooth, coniposed of flat grains.
D.trrleZlr~i~~i-laterallycompressed ; surface rough, conil)oseed of unsorted grains:
apertiiral rim irregular.
I ) . INIzCPOIut(X--Riirf~tc.e sniooth, coniposed of sniall, flat grains; apcrtural rini of
organic eeiiient.
D. lohostonlo-surface suiooth, coniposed of unsorted grains; ape1triral rim irregiiliir.
I ) . terw-coniposed of unsorted grains, often with one or t m o largc, angular grains
at ahoral end.
I). / i s r/ir,,.e?Lsis-coiiiposetl of angri Iar grains.
1). rotunda-conil~osetl of nnsortetl grainsand diatoni f r list iiles, ofteri with additional
large, whole diatoiiis; apertural rini surooth.
D.I~imrthogii-laterally coinpi essed ; coniposed of angular grains anteriorly, 14 it h
I I I O ~ regular
C arrangement of unsorted grains posteriorly; apertural riiir irrt,gitIar.
D.i~iscirlwl//-composed of unsorted grains; apertural riiii siiiooth.
D.2rbrs-composed of angular grains; apertiirre of rotire M i t h qliort collar.
B. Forms of Difflugiu with collar or short neck
f). hrr7licollri-surface rough, c ~ o i r i l ~ oof~ dmineral grains a n d diatoni frnstitlcq;
apertural rini irregular.
U . inicru.stumrc-coinposed of niineral grains a n d tliat ont fr iistules; apertural ritti
sitiooth.
D. lirri~~cticcn-c~otri~ioscd
chiefly of tliatoin frustules.
I). rZiffiri/i~~-coniposedof angular grains; apcrture with sinall collar.
D.I’etri~colo-eotiiposrd of unsorted grains arid few dtatoni frristules; collar oft en
obscured by large particles ; apertural rim smooth.
I>. lithoi,/iilu-eoniposed of unsorted grains a n d diatoni frustules; apertural rini
i r reg1i 1 a r.
1). vrceoluctm~-irregular aboral protuberances ; coinposed of angular grains; apertural
ri ti1 snioot h.
n./trh,‘osri-siirfacesniooth, coiiiposecl of angular grains niixcd with stnaller grains;
apert tire with recessed collar.
C Forms of Difflugiu with neck
I). pulex-surface rough, coniposed of unsorted grains and tliatoni friistulrs;
apertural rini irregular.
I ) . puurdli-surface sniooth, coiiiposcd of sniall grains a n d flat diatoni f r ustriles;
apertiiral rini sniooth.
I). ruhescew-surface sniooth or rough, coniposed of unsorted grains a n d diatoni
frustules; apertural rim of organic caenient.
D.hucillrrrl’rnrurn-Iliany with aboral horn; outline w a y he concealed heneath
construction of diatoni frustiiles; :tpcrtursl rim irregular.
D.~ms.sorvnA.~i-srirfacerough, roniposed of angular grains; apertural rini smooth.
D. br~~o/~hi/o-c~oiiiposetl of angular grains, often v i t h additional large grains;
ape r t Lira 1 ri i n i rregu 1a r .
I). lir~~~uris-cotitposcd chiefly of diiitoni frustules; a ~ i c ~ r t u r ~
a li i i sniooth.
i
D./ j b ~ f ~ ~ / ~ / ~ l - ( ~ ~ ) of
~ l l unsorted
~ J O ~ ~ t l grains and few tliatoni frustules; apertural
rirn sitiooth.
U . ,)ioleetr/-composed of unsorted grains anti diatoni friist riles; apert irral ri 111 irregular.
43*
644 R. T,. ELLISOX
a d (I. G. OCUES

D.~ ~ ~ ~ i ~ Z ~ ~ - c of
o iflat ~ o s e [apertural
~ i igrains; ~ riiii sniooth.
D.brrrl'llr'feru-roiii~)ose~chiefly of diatom frustules which ilia. 01
apertural rim slightly irregular.
IT). pcrrcrt-surface rough, coniposed of unsorted grains; aprrt iiral rini snioot ti.
D.ctrf)r"oZccta-snrface rough, miiiposed of unsorted grairls. apertr~ralrltir irrp-
gu la r.
1). io.l/rifor))22:S-surface rough, coiiiposed of unsorted, angular grains; apertiin~lrinl
s I 1 1oot 11.
D.r~orlosrc-at)oralrcgion often coniprcssed to form distinct lateral wings: srlrface
roiigh, composed of itnsorted grains; apertriral rini snrooth.
D. Horned forins of Difflicgfa
D.elecpns-composed of angulwr grains and diatom frustules ; apertural riiii
irregular.
D.unrirms-composed of angular grains and diatom frustules ; apertural riin
irregular.
L). ~~sni~~illurrs-surface sniootli, coniposeed of flat grainq; npertural rinr stiioot 11.
1). acuminata-spine distinct projection froiii 1)ocly; coiiiposeed of angiilar gratt~\;
aperttiral rirn of organic cement.
D.NIt//)ILr)r(~lis-s11rfaclesniooth, coniposed of unsorted grains: apertural rini irre-
gular.
D.tr.icor/rrk-siirface rough, coiiipohed of unsorted, angular grains: apert iiral ritii
irregular.
D.cororzcc-surface smooth, coniposed of unsorted grains; although toothed. the
apcrtural rini is smooth.
D.curuicnuli.s-spine coniiiionly appears as extension of body ; surface snioot 11,
composed of unsorted grains; apertural rim sniooth.
D.fe/~u.~trr-cotiiposed of unsorted grains and dintoiii frusttilcs; apcrtural ~ I I I I
slnootll.
D.z~e?itricoscc-surface rough, coinposed of unsorted grains and diatonr frustiilvh;
a p r t riral rim irregular.
I). bicrurl's-surface rough, coinposed of iirisortetl, angiilzr grainq: apertunil r i i i i
irregular.
I).rl;.sterL~~n-coiiiposed of unsorted grains, some with a few diatoiii frustiiles;
apertural rim smooth.
1).bfclcns-conipored of flat particles which appew t o h e arrmgccl in several
rO\CS ( H A R N I S C I I , 1958).
L). I7thr)i)lites-coiiiposed of flat grains closely packed it1 rows (HAKKISCII, 195s).
D.clfl1,iforiiizk-srirfac.e sinoot h, composed of nnsortecl grains: apertnral r1in
stlloot 11,
P o n t i p d u s i c ~R I I U I V ~ R L E R i895-,ihell
, pyriforiii, with a constriction to foriii a ~ i r c k :
internal cliaphragni or bridge on level with constriction: composed of :igglutin:tted
iiiineral grains.
CucurbitPZlu PENARD, 1902-shell ovoid, with four-lohed apert lira1 collar; ~ ~ I I I J X M ~
of agglrrtinatcd mineral grains arid siliceous idiosonies: aperture circular, recesed.
,i7etzel;a OGDEN, 1979-shell ovoid; coiiiposed of siliceous idiosoiiies, often 11 itli
agglutinated ininernl grains :tnd tliatoni friistulcs; apcrtrrrc tilay have rim of organic
cenient.
Fatiiily Hyalospheniidae SCHVLZE,
1S ' i i
(Plates 8 a n d 9)
Shell ovoid, elongate ovoid, or pyriforin, laterally coniprcsscd: coiiiposcd of 1iro-
 Identification of P o s d Testate ;hnoebae ti45

teinaceous iiiaterial and xenogenous siliceous shell-plates or agglutinated tuincral

grains; aperture terniinal, oval, circular, or slit. Several genera.
H?yu~oSpherLiu STEIN, 1859-shell elongate ovoid ; coniposed of proteitiaceous
material; aperture oval.
Hdeopera LEIDY,1879-shell truncate ovoid, wedge-shaped in lateral view; ante-
rior part forrned of siliceous plates, posterior part mostly aggiritiriated; apcrt urc a
n a r r o w slit.
NebrZcL LEIDY, 1875-shcll pyrifortii to suhcircular ; coinliosetl of ovtrl, rod-like,
su~)-cii~"drangrilar arid irregular siliceous plates; aper tiire most Iy oval, some nit11 t hiri
collar.
Qundrz~lcllu, COCKERELL, 19OS/PnrnllucndrulN I ) m r , m i ) R E , 1932-shell ovoid o r
pyriforni ; coniposed of regularly arranged, quadrangular shell plates: aperture oval.
Specinien.: of t hcw two genera are difficult to separate iriasniuch a s the priniary clif-
fertJnce IS t h e coinyosition of the shell plates; Quadrulelltr has siliceous, whereas f'u-
r(4(yu(drdu has calcareous plates.
Lesyucreusk SCIILCN~LEILU ER, 1843-slicll co t i 1 I iia-sliapd : c o tiilmhecl c.Iiic.fly of
curved, siliceous rods, with agglutinated tiiineral grains.

Patriily h i p hi treniatidae P'c)cTTT~:,

1913
(Plate 9)
Sht.ll elongate, laterally compressed; composed of proteinaceous material, mnie
with agglutinated particles; apertures are located a t each pole of shell. One genus-
Amphitrema ARCHER,1867.
Fainily Euglyphidae WALLICH,
1864
(Plate 9)
Shell ovoid or circular; composed of regularly arranged oval or circular, autogenous,
siliceous plates; some with siliceous spines, spinositp partly a function of hahitat,
with spinose spccies i n littoral zonc htwiming less spiriose i n profundal zone (Sc3110~-
B O R N , 1983); aperture circular, ovoid or lenticular, surrounded bp either denticulate
apertural plates or rim of organic ceinent. Several genera.
fic;u{jlypl~nDUJARDIN, 1840-she11 ovoid, in cross-section either circular or ovoid ;
apertiirt. hordered by denticulate apertural plates.
Assulr'w EIIRENBERG, 1872-shell ovoid, laterally cotiipressed; aperture Icnticu-
Jar, with a slightly dcnticulated rim of orgmic cenient.
Plncocista LEIDY, 1879-shell ovoid, laterally coiiipressed ; spinose and non-
spinose forms habitat. dependent.
Sphenoderia SCHLUMBERGER, 1854-shell ovoid or circular; aperture lenticular,
v i t h small collar.
Y'rucholruglypha L)CFLANDIW, 1928-shell ovoid: aperture with a denticulatecl rini
of organic cement.

Family Trineniatiidae HOOGENRAAD a n d J ~ E G R O O T , 19-10

(Plate 9)
Shell ovoid, with apertural rcgion compressed; aperture sul~terniinal,circrrlar,
invaginaf eci, with riiii bordered 1)y small denticulate plateh. One genu-Trr'ttemn
DLJARDIPI', 1841.
Fanlily C~yphoderiidaeDE SAEDELEER, 1934
(Plate 9)
Shell retort-shaped; aperture circular. Fossil siwciiiiens of thc t n o genera, Cypho-
 C U S 1877, arc difficwlt t o separate
rlrrl’ri S c F r L u M m R G m , 1845, a n d C N ? ~ / ~ I BLmny,
hecansr the characteristic thin, disc#-likeapertrrral collar of t h e living COIN)JUSCU.S is
lost.
Cyphodoriu is n coninion coniponent of modern testate coniniiinities in t h e sedinient
of oligotrophic. lakes ( S C I ~ O N H ~ R r.t Nu,/ . , 1965; R ~ Z I C K A ,
1982), hut its shell is
fragile a n d 1 5 rarely preserved.
Paniily Arcellidae EHREKBERG,
1843
(Plate 9)
Shell circular or round; c o i i i p s e d of proteinaceous illaterial; surface siiiooth or
p i t t e d ; aperture circular, central ; rarely preserved i n lake sedinient s. One genus.
Arrcdlu EIIRENHERO, 18:XkhelI profile hetnispheric.al or plano-convex ; s r i n i n i i t
may he depressed; basal border c.oniirionly present laterally; surface niay he inoldetl
to form angular facets or lateral spines; aperture n i t h sniall collar: iiiay he siir-
rounded hy pores.

Rote: since this manuscaript was subm itted for publ~r,itionI’ontiguk(\io c p w t n b t l i 9 ,ind Y.roiu-
essn h:ive been transferred t o t h e genus 2 tvkowicicr OCDEY. 1987.

CENTROPYXIS ~

Plate 1 . Speries of C‘Pnfrop!jxi,s,

 Tdcritifivirtion of Fossil Testate Amochac 1347

Plate 2. Species of Cmtropyxis (cont.), Trigonopyxis and Cyclopyxis.

nidulus elisa spectabills romprwa

Plate 3. Species of Cyclopyxis (cont.),Bullinularia, Phryganella, Plagiopyxis and Pontigulctsin.

ii4s R. L. ELLISON
and C . U. OGDEN

_.
.....

P h t e 4. Species of Uifflugin.
 Identificaation of Fossil Testate dinoebae 649

Plate 7 . Species of Uifflugitr (csont.), Setzelitr and Cucu&tella.

650 R.L. ELLISOX
and ('. C. OGDEN

penardiana rnarginala cartilala CJaledld lubulosa

Plate 8. Species of Heleopera, Lesguereusia and Nebela.

100

P
lO1lllllld acinthupora slrigosa Lotnprrssa inuscoium ~caiiiltiidv1~3

p'omc)gl ~~~~~~
cii51n1a

SPHENODERIA TRINEMA
PLACOCISTA 0 -- 0 -
i\biio\lris Ienld lmearr +ti1 hplys
TRACHEL- AMPHI- QUADRUgLLA

50

spinosa denfafa flavum slnpulld CLJnQIL13 PIecjaiiS ) apl\\C \ynlniel!iLa

Plate 9. Species of Arcella, Campascus, Euglypha, Assulina, Sphenoderia. Trinema, Plucocista,

Twacheleuglypha, Amphitrema, Cyphoderia, Hyalosphenia and Quadrulella.
 Idcntificntion of Fossil Testate Amoebae 651

4. References

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Piof. Dr. ROBERT L. ELLISOT

Dcpartnient of Geolopic'il Sciences
tTnircrsitg of Virginia
USA
Dr. C. G. OGDES
British Museum (Satur,il History)
Department of Zoolozy
Cromwell Road
London SIT7 SBD, LT.I<.

Manuscript accepted: April lst, 198i