AD-A090 680 COLD REGIONS RESEARCH AND ENGINEERING LAB HANOVER NH F/6 8/1MORPHOLOGY AND DISTRIBUTION OF THE ACANTHOECIDAE (CHOANOFLAGELL--ETC(U)JUL 80 K BUCK

UNCLASSIFIED CRREL-O-16 NL.....u..uubuli/Bill|IEND

REPORT 80-16 LEVEYItMorphology and distribution of the Acanthoecidae(Choanoflagellata) from the Weddell Sea during the

austral summer, 1977

I

C2D

...

For conversion of S1 metric units to U.S.,'Britishcustomary units of measurement consult ASTMStandard E380, Metric Practice Guide, publishedby the American Society for Testing and Materi-als, 1916 Race St., Philadelphia. Pa. 19103.

Cover: Scanning electron micrograph (approx.x5500) of Diaphanoeca multiannulatasp. nov. from pond on ice floe in WeddellSea.

CRREL Report 80-16

MNorphology and distribution of the Acanthoecidae

(Choanoflagellata) from the Weddell Sea during theaustral summer, 1977

1"'L~~r~uk Alcosion Fow/ Krt/Bck NIS GEMI

MC TABUnanouncedJustif ic t ion

Ju1¥-1980 ,Distri".

Ava 13.A',

Met.- special

Prepared tr

NATIONAL SCIENCE FOUNDATION

UNITED STATES ARMYCORPS OF ENGINEERSCOLD REGIONS RESEARCH AND ENGINEERING LABORATORYHANOVER, NEW HAMPSHIRE, U S A

Approved Dfo publl¢ relea . dlwybul on unhmied

IJA

SECURITY CLASSIFICATION OF THIS PAGE (When Date Entered)

DOCUMENTATION PAGE READ INSTRUCTIONSREPORT DBEFORE COMPLETING FORM1. REPORT NUMBER 12. GOVT ACCESSION NO. 3. RECIPIENT'S CATALOG NUMBER

CRREL Report 80-16 1-)RO9 U4. TITLE (end Subsitle) t.TYPE OF REPORT & PERIOD COVERED

MORPHOLOGY AND DISTRIBUTION OF THE ACANTHOECIDAE(CHOANOFLAGELLATA) FROM THE WEDDEP,.6 DURING THE SPERFORMING ORG. REPORT NUMBER

AUSTRAL SUMMER, 1977

7. AUTHOR(e) a. CONTRACT OR GRANT NUMBER(&)

National Scien "FoundationKurt Buck Grant DPP76-7251 and DPP76-80738

9. PERFORMING ORGANIZATION NAME AND ADDRESS i0. PROGRAM ELEMENT, PROJECT, TASK

U.S. Army Cold Regions Research and Engineering Laboratory AREA a WORK UNIT NUMBERS

Hanover, New Hampshire 03755 L

11. CONTROLLING OFFICE NAME AND ADDRESS 12. REPORT DATE

National Science Foundation Iuiy 1980

Washington, D.C. 13. NUMBER OF PAGES

14. MONITORING AGENCY NAME & ADDRESS(II different from Controlling Otfice) IS. SECURITY CLASS. (of thle report)

UnclassifiedISa. DECL ASSI FICATION/DOWNGRADING

SCHEDULE

IS. DISTRIBUTION STATEMENT (of thit Report)

Approved for public release; distribution unlimited.

17. DISTRIBUTION STATEMENT (of the abstract entered In Block 20, if different from Report)

I. SUPPLEMENTARY NOTES

It. KEY WORDS (Continue on reveree side If nececary end Identify by block number)

Antarctic regions Sea IceChoanoflagellates Taxonomy

Ocean environments

Plankton

21 ABSTACT (CmEn.e dn Pe~r80 eI* OI eweay mid idemfify by block mumb)

Eight species of loricate choanoflagellates (Acanthoccidae), Acanthoecopsis spiculifera, Bicosta spinifera, Bicostaantennigera, Caollacantha simplex, Calliacantha multispina aff., Crinolina aperta, Diaphanoeca multionnulata, and

Porvicorbicula socialis, were observed in samples obtained from the Weddell Sea during the austral summer, 1977.D. multiennulata was described for the first time from these samples; the other organisms have either been described

previously or are being described at this time. The distribution of most species within the Weddell Sea was widespread.The distributional, environmental, and morphological range of A. spiculifera, . spinifera, C. aperta, and P. socialis wasexpanded. Habitats in which choanoflagellates were found included the water column, the edges of ice floes, ponds on '

6091 1473 goD?8 OF I NOV 05 Is OL.eTE UnclassifiedSECURITY CLASSIFICATION OP THIS PAGE (When Does Entered)

Unclassified

SECURITY CLASSIFICATION OF THIS PAGE(Wmhen Data bnlend)

20. Abstract (cont'd)

ice floes, and the interiors of ice floes. The presence of choanoflagellates within the ice indicates that there may bea closely coupled trophic relationship with the other two biological components of the ice community, the ice algaeand the bacteria. The presence in the ice of seven species with both a caudal appendage and anterior projectionssuggests a positive relationship between this lorica configuration and the ice habitat. Mechanisms of variance oftransverse costal diameters between genera may be useful to the taxonomy and phylogeny of this family.

Unclassified

SECURITY CLASSIFICATION OF THIS PAGE(Whon Daa EnterOd)

PREFACE

This report was prepared by Kurt Iuck, Biological fech niian, Snow and I( eBranch. Research Division. U.S. Army Cold Regions Research and -ngineeringLaboratory. The author gratefully acknowledges the guidance. ,upport, and in-spiration of Dr. Sayed EI-Saved, Dr. Greta Frvxell, Dr Nestor ottino, and Dr

Saturo Taguchi. S. Ackley, M. Meyer, R. Warner, and the .rew and ollit ers of the

USCG Icebreaker Burton Island assisted in making the field hollec lion% 1 he

author also appreciates the assistance of H Thomsen, I Manton, 0 Moestrup, RNorris, S. Eubanks, and M. Hoban who supplied inspiration, tet hn( ial astan e,

and critical reviews during the preparation of this report The ( dlec lon programwas funded by National Science Foundation Crants DPP70-1 5251 to U S Army

Cold Regions Research and Engineering Laboratorv and DlPP7t-8071,8 to IexasA&M University. The analysis was funded by the Smithsonian Oceanographio.

Center Cooperative Systematics Program, National St ente | oundatioln ("rant

76-23979.The contents of this report are not to be used for advertising or promotional

purposes. Citation of brand names does not constitute an official endorsement or

approval of the use of such commercial products

iNo

A.

CONTENTS

PageIntroduction 1

Literature review 1Objectives

Materials and methods 3,

Results 4Observations 4Distribution and environmental tolerances 18

Discussion 19Morphology 19

Distribution 20Habitat 20

Summary 22Conclusions 22Recommendations for future work 22

Literature cited 22Appendix A Type descriptions of Acanthoecidae collected from the 1:

Weddell Sea 25

ILLUSTRATIONS

Figure1 Morphology of Acanthoecidae 22 Cruise track and pack ice limits of the Weddell Sea hiocruise. Feb-

ruary-March 1977 43 Diaphanoeca multiannulata Line drawing b

4 Diaphanoeca multiannulata. Intact lorica with caudal appendage, 4transverse and 13 longitudinal costae 6

5 Diaphanoeca multiannulata. Lorica with 5 transverse costae 66 Diaphanoeca multiannulata. Complete lorla with 5 transverse and

11 longitudinal costae 87 Diaphanoeca multiannulata. Whole lorica showing 12 longitudinal

and 4 transverse costae, anterior projections, and a caudal ap-pendage 8

8 Diaphanoeca multiannulata. Whole lori(a showing (ostal stripcomponents of both longitudinal and transverse (ostae 8

9 Diaphanoeca multiannulata. Anterior transverse costa and anteriorprole( tions 8

10 Diaphanoeca multiannulata. Partial lori( a with intact sec(ond andthird transverse Costae 9

11 Diaphanoeca multiannulata. (oales( Ing of longitudinal costae be-low the fourth transverse costa q

12 Diaphanoeca multiannufata In(omplete lori(a with (ostal stripbundle 9

I (rinoina aperta Line drawing 10

IV

F Igure Page

14. Crinolina aperta. Lorica with 2 transverse costae and 12 longitudin-nal costae and projections at the posterior and anterior ends 10

15. Crinolina aperta. Lorica showing the size difference between theanterior and posterior transverse costae 10

1b. Crinolina aperta. Section of the anterior transverse costa and alongitudinal costa showing the [-shaped transverse costal stripand the Y-shaped notch made by the longitudinal costa in whichthe transverse costa sits 10

17. Parvicorbicula socialis. Line drawing 1118. Parvicorbicula socialis. Lorica with 2 transverse and 10 longitudinal

costae and no caudal appendage 1119 Parvicorbicula socialis. Posterior portion of lorica showing the se-

cond transverse costa situated at the posterior end of the anter-ior longitudinal costal strip 11

20. Parvicorbicula socialis. Detail of the attachment of the longitud-inal costae to the anterior transverse costa showing the enlarge-ment of the anterior end of the longitudinal strips 12

21. Parvicorbicula socialis. Posterior transverse costa showing coales-cing of longitudinal costae into a stalk from which a caudal ap-pendage arises 12

22. Parvicorbicula socialis. Lorica showing longitudinal costal reduc-tion to .3 below the second transverse costa 12

23. Parvicorbicula socialis. Complete lorica with intact cell and flagel-lum projecting out of the anterior end of the lorica 12

24. Parvicorbicula socialis. Aggregations of loric as, indicating colonNf orni at ion 13

25S. Acanthoecopsis spiculifera. Line drawing 1320. Acanthoecopsis spiculifera. Complete lorica with lb longitudinal

and .3 transverse costae, anterior plrojections and a caudal ap-p)endlage 13

27. Bicosta spinifera. Line drawing 14x ~28. Bicosta spinifera. Lorica with 2 lon1gitudinal and no transverse cos-

tae and a caudal appendage 14

29. flicosta antennigera. Line drawing 1-30 flicosta antennigera. Lorica lac king the distal ends of the anterior

projections 15

.31 Ricosta antennigera. Complete lorit a with c ell, 2 longitudinaland no transverse ( ostae, anterior prole( tions and a (audal ap-pe'ndage is

12 Ca/Iiacantha simnplex. [Line drawing 10h.33; Calliacantha simnplex. tLori( a with 2' transverse and 4 longitudinal

(ostae, in( omplete anterioir iproi( t ions and a ( 01.dal appendlage lb34 (allhacanthai sirnplex. [ Ciri( J with 2 transverse anid 4 longitudinal

(ostae, ( ornplete anterior prlvr~( tion%, in1(] an in( omiplete 1atidal

appendlage lb)35 Caliacantha multispina aitt I ine drawing 1 7

V

Figure Page

36. Calliacantha multispina aff. Orientation of the lorica showing thefull extent of the caudal appendage and anterior projections... 17

37. Calliacantha multispina aff. Incomplete lorica showing the 5 longi-tudinal and 2 transverse costae ..... .... 17

38. Calliacantha multispina aff. Incomplete lorica showing 2 transverse

and 5 longitudinal costae and 6 anterior projections ........ 18

TABLES

Table1. Location, date, type of sample, surface temperature, and salinity of

the stations occupied during the Weddell Sea biocruise.

February-M arch 1977 ........... ........ 52. Distribution of Acanthoecidae from the Weddell Sea biocruise,

February-March 1977 ........... 73. Key for the identification of loricas of Acanthoecidae from the

W eddell Sea ................... 19

vi

t IAJ

MORPHOLOGY AND DISTRIBUTION OFTHE ACANTHOECIDAE (CHOANOFLAGELLATA)FROM THE WEDDELL SEA DURING THEAUSTRAL SUMMER, 1977

Kurt Buck

INTRODUCTION politan species, the polar reports hav., been pii-marily of arctic species

literature review As a group choanoflagellates seem to favorThe placement of choanoflagellates as a calm waters of nearshore areas for their habitat,

group in either the plant or animal kingdom has being found in the stagnant water of saltvacillated back and forth for a century. Based marshes (Leadbeater and Morton 1974a), tideupon mitochondrial and flagellar substructure pools (Norris 1965), fiords (1hrondsen 1974,(Leadbeater and Manton 1974, Hibberd 1975, Thomsen 1973, 1976) and (oastal waters1976), these organisms are now assigned to the (Throndsen 1969, Manton et al, 1975, 1976) Ob-Families Codonosigidae, Salpingoecidae and servations ill open ocean have been made onAcanthoecidae, Order Choanoflagellata, Class two spe(ies of the genus Bicosta (ThrondsenZoomastigophorea, Phylum Protozoa The cells 1970b, Reynolds 197b), Neustonic (Norris 1965),of the Family Codonosigidae are naked, the cells epiphytic (Ellis 1930, Norris 1965), and plank-of the Family Salpingoecidae are thecate tonic (Throndsen, 1974) modes of existence have(bounded by an investment without costae), and been described. Manton et al. (1975) propose anthe cells of the Family Acanthoecidae are asso( iation with sea ice as a common denomi-loricate (bounded by an investment with costae nator for a group of Acanthoe( idae observed inmade of siliceous strips) (Leadbeater 1977, Nor- the Canadian Arctic, although no specifi(ris 1965). The Acanthoecidae are further dif- reports of such an association were made. Theferentiated from the other choanoflagellates in presence of at least one species under ar(ti( seathat they are exclusively marine. This study will ice has been do( umented (Manton and Oatesconcern itself with the Acanthoecidae. 1979b). Environmental tolerances of the family

With the erection of the genera Crinolina as a whole seem to be wide; they have been(Thomsen 1976), Bicosta (Leadbeater 1978), and found in water with temperatures ranging fromCalliacantha (Leadbeater 1978), there are now -1 °C to 240 C and salinities ranging from 5, toapproximately 12 genera and 40 species which 36/,,,. Without exception, insufficient data existbelong to the FamilV Acanthoecidae. Geograph- to be able to establish habitat, environmental, orically, loricate choanoflagellates have been geographi( al ranges for any given spe(ies.found in polar (Manton et al. 1975, Throndsen In describing the morphology of the [amily,1970a,b, Reynolds 1976, Deflandre 1960, Manton Acanthoecidae the terminology of I eadbeati'rand Oates 1979b), temperate (Leadbeater 1972a, (1972a) will be followed I he hir a is ( oino)n)oe(Jb. 1973,. Manton and Oates 1979a, Moestrup of longitudinal and transverse ( ostae(0 ig 1) 1 he1979, Norris 1965, Thomsen 1973, 1976, 1977, anterior end is open (the po(sterior end in (ro-lhrondsen 1969, 1974), subtropical (G. Fryxell lina spp. is open as well) an( has the i lagelhmmpers. comm ), and tropi(al (Manton and Oates t)roje(ting from it In the I)resent tax(inmli(1979b) regions. With the exception of Deflan- s( hene Ihe lori a shape, liri'sen( v or aben(c otdre's report from Terre Adelie of Parvicorbicula anterior prolelions, nUmbers oi (ostae (bot hsocialis (MeLnier) Deflandre, a seemingly ( osmo- longitidinal and transverse), and in one ( ase thi

,,.

ik - anterior projections

anterior transverse costa

longitudinal costa

second transverse costa

transverse costal strip

~costal

strip

Pove lap

caudal appendage stalstrip

Figure 1. Morphology of Acanthoecidae.

orientation ot the costae (Acanthoeca Ellis) are costal strips, it is the non-tapered end that is re-characteristics most commonly used to differen- sponsible for joining individual costal strips andtiate genera. Costal numbers (both longitudinal connecting longitudinal and transverse costaeand transverse), presence or absence of a caudal The variation of costal substructure betweenappendage, and costal shape are the characteris- species has been largely ignored in the literature,tics most commonly used to differentiate making assessment of its taxonomic importan(especies. Each costa is made up of rod shaped difficult. The shape and size of the protoplast,subunits, called costal strips, which in some the length of the flagellum, and the number andspecies are heteropolar. The heteropolar costal length of the collar tentacles are characteristicsstrips are tapered at one end and bulbous, that are usually included in a species descrip-T-shaped, or L-shaped at the other end. Other tion, but differentiation on both the generic andcostal strips are usually tapered at both ends In sp-cific levels is based solely upon lorica mor-the species that have heteropolar transverse phology.

2

hemnv the ilom a has beent detetirmined tio I)t' sil- A( anrthovie dav. partmi ulat I\ wt Ih rtvtIw t to pretiVoris I jIt)sutt to h\11rtiliimr ) li if (II %a \ ois obslt, atiiin. is oine wi the 1nia11n oIiite ti\ (Is

heater and NMorton 1974h1) adl X-ra\ dfldl\si, ot this repcirt Another odbleitioe is ti, e\,iiti

I11 1 VI Ill',t I~) 11,1 h V' bVV liei LISt' tii ti't tis th l ii std l~ h lIU stts Ii i t Ii hIit, 1)(it ii's vN it hW\hen tt't lt k. stailin' Itri rel WTIt0 thi' sit It )t so nriiitg ilil tro ntil rust, lye Ii ,ii ittvmpiht ti%%er' uundtIn ted and itigit iNiv rt-SLI Its ilhta0ittei[ I l tli t Ie t M I ( II Itiii lliitp t I ' )I thu11 ( u1,11

he, 'ullI II eia holdui ing I h (i .mitr 111 ils till oin thn ' s. il'ta i e iic r thi rr ii' nl

JI. tit ani do. hulioin t e uai t ri i t nut i ths'pori'

if~i Iiir ti t (If I It'm ir,iiuu f9 1,1dIttittt t I i MATERIALS AND MFT HODS

h fUictullat' a'Wl diud ii10 ')I' tilit l l iaiit i ii sui. it ii'piiuu

tii utu 11, 51 Ip ( l uIt t vI I0 I it ' I, e!llr (' itulu drt It 200i it hu01 1t hif ' uno t (' til lIt It.ItilI)\itii' b III W i i lts l ' 11,901011 1 h)ld till (1' lnr ha hei.I\u o I it t q tc- nidu i ti' gI miiiht t ilithi'llhh litelr ngmid ii'tltn h'u, 1tIi, ilusir1 I! twtiil "Impi g Imts imld ht r hadlit (I(th edge'it

It I li i. 1 dI LI t 11( 1' I ( Itt i t (I i I (I Htill' it J iii ( I t I It ICI 2 Is tl Ii l i (t i Iitl t ItIIIIIIt % I ii t I 1 i t a I it ' ii it I

I t' ,iits n 1n7 I7 t tii'tu I,\t III liiitl I'I , It Ie I i i 1111t it rp \,I \Is % st I trIest'n11.it .is lit, It as khI, r(I IssIhi ,

HIMi 11M lt a J 111 lltit I,\tii tut1 1( u nt ti lt tit' li inptil mntd011 iiu't.u t e n wr l Ilil t tu tKItI

a ) 14 '.iil s It ti nli i' anuLh lIi 1,\ i t t I" iita Ihurir ini aha o tiilii'tti' nia' tus e to ti\li't

I'iueed b itlsertl lie9 iuluinule ma. i', pitt oe .i lt i ih the lih 1100 Miii oI\ han NI r i the ede (iii

altni endII ' th eum,4 tep s Ityais IItiutttsi'it If mnsvi tit i a oeateid iro iiondii nittote liloti I al

m ut " il helt lIm itlitl r oloI'iit M tin u11:,tc oi t~th iit sttmhlpe %till I s i itt t1 e v an ti eu tOif

I lepermth mo Ii m~idt itm' li, toi litevtcirea111 ttts lit' Vs1St MlCS I0i he satwoles e' thitm I(,.

1)111lo dri0 it L( (1 )stoi 1kldkHcedge or hi po snds tiid hetie %%is oith ted . lh1111,11011 d)5 III,' veit' tm (uiri shott Ii'. nix ii (1 1 11CI(ir

Ob6ei t I i[)Ies IL"InI lI I(1(le tfIldM llt l 1itin tu ( m1I neI II tha 1ini .liiixed l, Ilil-L to Ii' ovrng t-)17) t'ternttnitng I the v eigpt al htrstht arid the exi) los tttitmmn irtontenimasthn (I aped

I

60W* 55°

50o 45

TVALPARAISO, CHILE FRMSUIAG.

.. . ARV. 15 MARCH DEPART 10 FEB.0. 3 FEB

16(B) 15ORKNEY 0% (A)

17(C)

18(D) .- 40'

ANTARCTIC 19(E)

SPENINSULA **.I

20(F)65'S

21(G)

I 35'

I22(H)

23QI)

24J)

ICE EDGE--.,,

'N /70'S

25(K)

SHIP'S ROUTE

26(L)

20'

\RETURN TO B

Figure 2. Cruise track and pack ice limits of the Weddell Sea biocruise. I ehruary-.March1977. Dashed line represents the pack ice limit and the solid line represents the cruiseitrack.

away and the e'dges were sealed with tingernail RESULITSpolish The samples for the SEM were g4rounche~lto the stub with silver paint antil toate(I with Observationsgold-palladium. The light ni icroso(o)W wad, a Diaphianoeca flhIdl,I.Il~n, lp r..I I,.. 12Zeiss StandardJ K with phase optlics,; the ,( ann~ng, l),am)oi, I ()rI( , htrri' .,l plttI.ln ,v,mi(cros(copes used were Hita(:hi ISM U-I , aiil (ostae., 4 (or 'S ( omlo)'c,,, t 11 1 I oxt..al ,,Ii pxU-IS. huiho)ux l one, e,1(t. t.lIIrI ,lt i,iherl (, 1( ;1 I' (I

44

-~~ MA (P)6

Table 1. Location, date, type of sample, surface temperature,and salinity of the stations occupied during the Weddell Seabiocruise, February-March 1977.

Latitude L ongi tudel Da)1 te 'wrllei %llIjf( v, %arf,( v.

Sta 11% t 9'.- 1'7/ I I)e 1*nli ,o( 1 S11 i"

A 1O17 0' 440 4 0' 1 F 1eb Nv 1 7 4 1

IF 1'57 0' 510120.

11 feb Nt 1 7 114

B-C S10

0' 20

1 0 17 Feb it"

B-A 610580 1' S22S S 0 17 F eb NO!

b t1058 0' 520250' 17 feb Nvt (t) II 2

(-1) b401 10' 051056 18 feb Net

D 64057 6' 50*20 2' 18 fel F e*

) b4057 6' 560262' 18 f eb Nvt I 7 14 1

)A 0501.10' s10S1 0' 19 Feb Net

t b5 021o'

510280' 19 Feb It"f (15O2 1' 51028 1' 19 feb NO! 1 7

1- ('5059 0' 5)142 V 19 Feb N,!

f ,b°45 1' 51)01015 20 Feb NetI I I 118

I (; h7°0f4 5 S()*t) 0' 20 | eb Nt

( b7°4l) 4 44007 2' 21 Feb Ne 1 08 11 1

G-il 68°4 0 I 4702811 21 Feb Nvt

It 64"21 0' 4(,04t, 0 22 Feb Net 1 7 11 r

F -I 6,4 ,1110 46',0

r 1 22 F eb Nei

I 7tN' 1r 2 4501,8 21 Feb It"

I 70°15 2 41h I,8 21Feb N,! 22 114

I 7 711°) 0'

44X) t 2 1 [Fe) N.'t

I 71025 0 450 0! 24 1el NvI I 117

I-K 7200811 410V) 24 F eb Net

K 7107!0 410440 25 Feb 1, of

K 7 1007 o 420440 25 Feb Ne. 1 11 4

K 1 740118 It 41°0810 25 feb Net

1 740I7 6 18054 8 26 feb Net I 11 4

I %l 74'41 V 1,0t2 1 26 feb NvtM 750 't! 5' 1427 0 27 Feb Net 20 4!

M--N 77027 0' 14054!! 27 Feb Net

N 77°4t I)' IS01) I)' 28 feb Ne! 2 4 1111

N-) 77°4b f0' 5°5q Fl' 28 Feb Nt( ) 7114 6' 1404 1' 1 Mar Nvt -1 I I I i

()-P 7W°49 7' 12027 4' 1 Mar NetF' 70'19 8' 280400 2 Mar Net -1 7 11 I

'I e IIh. pond or edget 1( v' Ilov ( oft,

at interval,, oF 11'2. 4, r-1,2, and 7 (ostal strips (1 abht 2) Altilough' tFlit nuiieli,'r of t lt' it It,

froin ,iterior ends of longitudinal (ost,l, (I ig. 4-f,), twelve lngitudinal and{ to11 ti,!!

longlidinal ( oitav 11 1. F ( ll ttwd t - t) ( Fntal verse ( o( ,i ( v ,s thc nIi to I igs 4, 1)) I It,( tiot

strips. bLunt at both ,itd, of slightIv stialher i1i( ro(s olV of this org,lism jetrmiltltd Fh' slitm

dtaniter than r,nverst, I otrl srips, proe( tio, tFrt' 11 tlit, ( tostl to ha' h lIts terrt( I h' tr,ins

1-12, (lost,1

strip% in length 0( 1 urrlng btuvond an- verse ( lst),t art' t ( i)m)(!t'td (fI , i111,uI (, osl~l

'rior I osl, ( aiilail |)ppevldagv I I ostil strilps i strips as there art' ltngiltudin,il tl(,t', iht, t,,t,IlIe~nglh arPing fro111 redI( lion ad( o nverg~nt(, ot strips, bulbous, Oil 011V, (Ind 111(1 blunt oil tilt,

longitudinal I os!at, o)ostertor to 4th transvers' oFhr ent, ar' aFtat III,(] to Fhe Iongitudinali is

(osti tat by their bulbous ,nds and tFo vat Ii tother b\Holotype Fiture .

Vpe lot ,ilion. 7015 2'S, 450 1b 8'W, 2 1/02/77, it v the blUnt tnd sollt'hrt altng thi aijoining

floe pond. -1 80C ( ostal strip (t ig 0) 1 hle pointof tat at hintF Ito

Loricas of this spe( ies were found in niany of the blunt enndi the adioining It st,l %tripi (i

the net tows and in four of the five ite saniplvs fers for t h ,osta; in the anttrlor transvters'

5

loI Ij 7t d!afodae taiw~

I~~~ Ig I I1 u IItIItIatik t n I ;) 1m t iil I( st( le I t , i t al

Igure Dlit 11) 11 mv 011L111111m (.

Table 2. Distribution of Acanthoecidae from the Weddell Sea biocruise, February-March 1977.(1) denotes, i(e

Acanthoecopsis Bicosta Bicosta Calliacantha (alliacantha Crnolina Diaphanoeca Par% ocorbicula

Station spiculitera antennigera spinifera multispina simplex aperfta multiannulata soc lahs

AA-B

B xB-Ct )B-C

CC-DD(I) x X xD x x x

D-E x x xW) X

F x xF-C x x

Cx x xG-H X X

H X XH-t X10I() X xxx XIl XX X XX

Ix x x

I-K X xK(1) x x x X XK X x X X

kIl X Xc Xc IKc Ic X IX

l-M xx X x

M Ic X IX

M-N X X xN I x X X

N-0 X x x XO) X Ic X Ic

()-I' X IcX XI IcFPIc X IcI Ic I

os(rtc the ,trelis ovw'rlapi eac hi other Iby 4t)- 5 0%/ oF anterior transverse costa with three or four even-their lengthI whsh Iii tn the I herd t raniserst' <( osta IV spacedl transverse costae posterior to it dis-thle o~e'rlap i, 10t- 15% (I eg,, 6. 7) SeIIc(c, the tlnguish D. multiannulata from the type speciesnuniher ot t-osthl strips is constant, this overlap of the genus, D). grandis Fllis D. multiannulataproduhc t's ttr,eiiwerse , (osie ot li hitereiit (Iliii- diiffers from D. pedicellata Leadbeater byete'rs, hem e ihe arrei shalpe ot the lori( a Ihli' possessing additional transverse costae (V.hongi ui11.1 hnlost e st ripsire' hun at hothiinids pedicellata has 3) and having anterior proiec-,erit e wvrlalp ee ,nother hy 10%, ot their httl lions constructed of more than one costalI striphe'ngt h (Fig 71 At or helo w the pot',hrior I raiis (I ig 6). he presence of costal strip) bundleswer,,e e celie, ,1 r,'diti tli iii the iiumiiiirr ci huiiigi within Ioricas was also noted (F ig 121tudinal t cest.c. uc( ur as..t they c onvw'rgi'. ,nd tunic (rnnohna .ipt'rtai (I e'acl.ihe'r) 1 hicimsein (F igsd l .(,dl 11 ,I],5)endiige' (I i/g 81) /,inlone~r a hlith 1 1-16)) 1 oint ais otI thsl'e it~' l.s wer't luun(d ,et mlost

(est ludiing the (au(Ia al~l),'n(Igze) aiic dcii,intir iit tihe southterii stlat ion,, tie ice as' ,s'11 ,is ii

at the third transverse costa we'restatist cal ly di- It' waiter ( oluilnc ( aht 21 Iht' lorica is e tn-ttert'nt hr stat0ions, I anti K 41 t I mn x 25 jm 4 strut t'd cit twelve htini.it cidiial antd two trans-2 Dim and IA __ 6 tim x 21 ±_ 2 tim, respectively vw'rse c os tat' witt t'e hciigit u~hd a t tisat' tii in

The p~resence of a caudalI appendage and tine Long hbyond t it'nc,, cit thle' Inre a (F igs 1 1. 141

0 " " " x7

,1 x: .,x , x :.a T- v ,, ; x - -II~ ~ x I x x x1 I I111 I I I I . .. I'- ... III II ..- : . . . . . . .

figure 6 [Jiaphiatioeta nitltiannila ( UfletefI I Igul ) I f)p,I~ I ) i)fdI IJ~) N )i1f Imn )I h(l 1f If

lo I ,u it hII 5 tf~il % % er %t, am lI I I h figitI I inhl 1 ( Sl m Ing~i 12 horigitiinial dfii .1 it'ln% er"I, %

me~ L N1f h~ v f1ov ;)ui d,%ation I t'ii ,litef rjv i)j liiid IOi( 111 (f.11oi ~ a I ppfl ri

J,Igf-I NI hl vie fln pndl S'Ition I

f igure 9. Dwephanov( a IIlIljti,111Iolata -ntersor

transverm, (Od a%, ndI anterior, fprole' I,())%'mf igure 8, Diaphafie'c a1 m'lu tiarinnolata VIhote Iections are I (mtl %trlpis In length I he'lorica showing costal strip) ((irlponents of both mode of iatl( /hrnerit of the', tr,en~, e'rse ct, ( (%,Ilongitudinal and transserse (o%tie %I M lve floe' strips to val: h other Ind to the', hirigilti,eneI 1opond, station f meW is I'ih' StI M ( e floe' pirid. Stitl I

( (Jdvs( Ing IU? IflflgILIdirhJ ()(a Ih'/,) i t)

IE)tj( '? f.i1

) II)II t %i'( 011(1 .11)( third 11"10 er't, if/(,% ftlfil tIII% 't'lk 4' ki h tiE,;m)( St')

osUtat. %Ii tl h( v' floe pond %tion)f I rum) 1)

figure 1.2 DidIphdflo4( a rftltiannu Idta In-complete Ioricai with (o (,i/d %trip handle Wi M~Ice floe pond. station 1)

9

I iAgur(' 14. (rinolina ap)rta Lorica vKith 2trans er(e ithe s.maller of the two is the ainteriotr

((itae and 12 longitudinal ( ostae and pro e -

trlio at the posterlor ind interior ends LM

f igure I f. (rinolifa aperta Line drdKig. 1,ater column, stdtion L

tiguP 16. (rinolina aperta s(( tion of the

anterior transverse costa ,and a longitud(tinal ( o a

showing the L-shaped ttanserse (ostl stri) and

Suigure 15. Crinolina, aperta Lorica showing the the Y-shaped notch made by the longittidmtil ( -

si/, difference between the anterior and posterior ta in whi( h the transv er e c ota %its S1 M ,i at'r

trans),erse costae. LM. Ice floe pond, station L. (olumn station L

I hli (tistal strips ot the, Iransvers ((ota, are ta, |i1ent ot the tranls.rst ((sta to thit- Imigitid

tapired at one' v'ndi ald I -shaptd at iht, other irial osla At tilt, antero r tran1sverst, e ()st,i ih(

end (Fig If)) Ihe uheptiirrl d ot end of em h ( stil ImIgitidirl ( otI torms a 'i sliapi.d 111t h In

strip) on Froth the' transverse ( iistii is cliret tedI to wir h the trainssersi, irsta is siate-d I hli Ion

wards the anterior (marrow) end ot thev loru a It Is (as rangedj it) suev trom 24 4t0 pill long. 12 If) pmi

the I shapel end that is reponsihli tfor the at In (iameter a1t the anteror endt(o id nd I i In

10

I igure IT. Parvi( Orbit uLJa sol iali L Lint' (Irdt' trig.

I igure 118 Ilari, it orbit Lla Sol whs% ori( a1t4 Ith tt$() tran%'.erw adl ten IiongituialI((I Otat' dirId nio id I adi IppendaLg'Nothe attd?( hflhfe't iv4 the (hiIdI end of theIiingititiial I (i%tdt to the mhidpo)inrt otthe anterior trainsItrst I)%tal %trii)% %I Nti ater I lun dIxiwi &~

tiof of ric %hn~fl theecod tafl~t'(e ( ~ t(1ittItt atl tht e~t litt wnl .11 cgtig 1,1 k li4;.1

gttll(IlridI ~~ ~ II ((IId )1tip logiudna ,, Imi Ieii Iiteah IIIt .eitt tr:i\ I'dIr t.1 t ostd (I it1

lionPro te 1( tut' Ot it~ u'hi* te ~ on PI mi that till ui1tttditi,I it ( '11ii tit t.

tra~te~i'( i',d bfor unitng t apuuul 'A(N tiatilt's ILitt'In aitthit tiilit IittI titldIIi

SI 1IIc t ' w ll- I l I(' '1tn a t II II

I igurv 21) I'm, o rbli, iiI ri 11%1)ea,

4 the .Ittat him-fr c It tht. 1( mgctuiti'lI/

(i iS~t It , Oile Interuit tIf11I% st(( is t'/laf'I t igu i ih LIII tli %m whi fi~ii ft.,)

%/icbt mg the' In,,g rce f i ll' .1iftiri if %'. cow9 II%f. Olt its Ing~ i).Ilis Ing) 'f 1( mg~.iti I

1 11( if - t IN Iitgitlib .II Stips %I NI I 'l t IIto' .1 Stalk Inif )11 11( 1 .It 1, 1 11# q//i I)

F 'ih''e 22 Patrv i orbit tiict w~ hd% I timi I 1 191sre P,9I' it uit tila % g,1g (ah wiuitqt lint

%hom,ing lmngitu~dinal ( ()%I re(im tion toc ( a ls crit m.Ii f ( vI/ andI( 1.1gelh/imrrt .Ir i e I

I twitm~ the' -it, Iofl ttdl%i.er%E' (osft I M 'fA. ig i m it f'.tit(t, nerifiren (t)I ith, Ificrs .1 1

Kajter ( ttIumn sftto I I,1,ite'I (Itiflf11 statimi) It I

I.I 44L

FI

figure 24. Parvicorbicula social is. Aggregaitionsof loricas. indicating colony formation. L M. I igure 2 5. A( aflt ht o)pl ,~" I( u litqjr I If)(Water column, station fl-/. ~,s injg

I t~(),I'll til 1,pIi .1,tid i 21 -6

I li( IM)i p tnit ' ui)Iltt' I ngrittai ini't

(?,1 1?(' r tti 1 inl titim' Ol t, (.s (mit't o t

(I sl (I 241 8 Il 21 2 1 1lc lt ild

pendagltit' x iji'l' 0 .l ti it'r 211 B-1h ill tit)~

lorca ith16 ongtudnaland tra.venf'co O(d theill uiial Iii'lig' iiip

tae. anterio proiection~and a cau(n a10n1g \\,is 14 -~ li-t'pill Itt'. ii Alit it g I i'i ithe ~I

B J d1 JJV dl0 11tr~ 01d 1111 W

transverste c ostae (t ig 20) P ro Iv( tions beyond junction between the longitudinal c ostae andthe anterior trainsverse (osta were observed an(] the interior projet tions ( an he seen as thi( ken-the' ii1ngitodInal t adi ( onverged p)osterio~rly to ing in the ( ostae (see arrow, Fig 28) The size and]give rise to the ( ddd appendage No) observai- gross morphology of the loricas from the Wed-tiois oil the Subunit strut tUre of the c ostat "as dell Sea fit well wit hin the original des( ript ionpossible duet totite st arc ity, of the organism in made with the LM id( the subsequent EM studythis material I here IN little doubt that this organ- by Reynolds (1976) (Appendix A).sin is thet spet les des( ribed by Norris (1965) (Apl- Bicosta aneennigera Moestrup (figs 29-30)

pendix A) with the light mit rosc ope and exam- Lori( as of this spec ies were observed at stationsinedi by I hoisen (1977) with the elet tro r(i ro- I and K in the ice and stat ions 0-P and P in thest ope Dist reliant ies in uoric a siie, the number water column (Table 2). 1 he lurk a of this speciesot longitudinal (ostae. and the arrangement of is turmed by two short longitudinal costae fromtilt- tran~sverset' ostae dto exist hot it is ttt that which spring the sweeping anterior proict tionshese cut tereil es ret let t add itionalI nlirlhologi- (two) and the c audal appendlage. T he anteriorali %ariabil ts' not noted by Norris in the Pm.if it protet tions seem to be composed of two costal

samples with Mm hitI he worked (1 horust'n 1977) strips eac h (Fig 30) while the (audal appendageBicosta spinhera I hrondst'n (F igs, 27-28) and the longitudinal (ostaie appear to be unit

Lori( as, b)oth wshole aind partial, werte founld at a strut tures. 1 he base of the (eit is situated at thenum111ber oI sout hero stat ions in the waiter ( OiUmnn tunttion tof the longitudinal ( ostae and the an-

anld in the it v' at tOUur stat ions (Table 2) 1 ht tenior propec.t ions (Fig 31) No) transverse (ostae

strut tre ot the lori a is simpJle, Icicking traris- "~erte observed T he total length of the Iric a, in-verse t ostat' and potsse'ssiing but two ltongituidinal (loing111 the (auddl appendaige, was 25 pm, the

(o~tdt' (F ig 28) 1 he longi1tudinal (ta tct r ss ciistant r between the ends ot the anterior pro

each other about two-thirds the length down the let tioils was also 25 pill The lori a riiorphtiligv

lorit a and give rise, to two sweeping interior prti- of the organisms t ollet ted from the Weddell Sea

le( tions aind to) the auia appendage ptister- agrees well wAith the type diest riptitel (Moestrop.

itirly I he length ot thet lori a troin its base to the 1979) (Appendix Al although tlhe arte somewhat

top) ot the anterior propet titois is to Pni I he smaller in si/t' [)tterentiation between B. minor

Figure 28. Bicosta spinifera Lorica with 2longitudinal and no transverse costae and a cau-dal appendage. The junction of the anterior pro-jection and the longitudinal costa is denoted by athickening (arrow). LM. Water column, station

Figure 27. Bit osta spinifera. Line drawing -M.

14

I ig'ire 30. Bi( n.fa Mitimigera torica 1,ickinp theistal ends of fill, ifrior prow( tin' \ote num

tion', of the mttriiir prow tefion and the hirigiliid

I itare 29 Ilit t'.ta ,tuitt'nigt'ra Line drao,. tig. niii (ii'.f a (,irro% /. LM\. 0. Ater coilun statttion ()

HKe ,iiitt'iIiii ai',.i t n I i N hu it ' min i t) ,'

ifil '. i t li i it o Ok ( t" , t1 iii 1il flt' f ((S~

Nntf tft t at'l up ilt , rif li (tlit' tt ti t'. i.t

Figur 1. H~ O~t dfltflOI~i'rd(nntpt'qe oric fli t1,Vl'dt ;iitt ftt1 tI'll 13t I14 ttu 1t1t Fl li''#ivth cll. ltmngtiidttalannino ta~i~er% ' (C~t~i'.flit, t'tintai' t r' ()I tl 'll ( .l im c l't fill. t'ii)

.ift~io P~f~(t!mfl dl~ a(al~didJ~efd~ f1'I cfit'ftft il'ilfttiI 'tii'itt liatft tt

( Iihi i pe M il lmi l.

I> _4 pl wi n iiuNtodto, et )

I igure 32. (all taantha simaplex L ine drawinig

Figure ;i~. Call Iaantha, simjple x I ora %ith 2

transverse (arrovs denotes Positio lolt the WI(on

trans verse costa) and 4 longihnil11,1 ( o.te. in-complete anterior projections and a (,miii/ q)ippendage. L M. Ice floe pond, (a (tion I.

ot ( nai.tans (( onfk'~edt I tibithr the c

thireet antetrior prolei tirns ,indlsi Iriiuiiidiiil

'10d I emauiet 19781) 1 he liihl itut ioguriphi (i

Ihe raplms t l st, i0' %\lls I, iltee 11o limr~ to

gah ' I thet riIriri vt Ireh It tt9oi sho%\li trt at

I,ir ( onlitious ( il thet' 1 o gi i a ')79a

(a/fiar antha nriltispiri ,111rrttn 'Iiind ( )tv'

Figure 34. Calliacantha simple'x. Lorica twith 2 foutr southerrn statirtil' I I nut' 2)1 Ihe shape. of thet

transverse (arrows denotes the position of the se- htrt a (r ormi ahl. thei prrnrof nithI ,rit-tion

cond transverse costa) and 4 longitudinal costae. pio( (i tons arnd a ( ,nidl i alipe'n(Lirge.,t an thecomplete anterior projections. and an incomnplete) nrrnrhebr ot Innigi I urir r',I\i) a'rnd Ira'1. nt si (Iv'.oh

caudal appendage. LMN. ice floe pond, station 1. ( rusta dl'resnt ( Iiatrat t'r i/v th is' or ganirsmi a nd

16

Figure 35. Call aca ntha in Lilt ispina afit. Line(Irai ing.

Fig.,ure M6. (ailiat anthia 11mi1ti-siiina at f. Orientation oi the lot-ica shovwing the full ex tent or thecaudal appendage and anteriorprojections. Note t'.s st in) thelongitudinal costae larroi andthe s irnutaneous longituidinalcosta/ reductions (arros ). L Al.Water column, station 0-P.

IIIa( c it ssIthiilie I enu0)LI ( 111111 ai I~ lIe 1i)[191

rLOHMI F tat,-t~~ jmvg t%%() i(t(rlrion (,c lstr~

After the No'( ()till re(Iu ti1M) III, Iimlli uldlliml

1I M ) l ~ ~ 'I I ' ( I t t t l ' I p Idl t l g Vl l M IS t t I ) I l '

Urs N'IIiiL1111tlillti ton eal ii ~Id ()I the u(rn( Itsil resfiel t II flte (hit,til e trlllil tih'e ist) t

Figure .37. Calliacantha nmulti- tra,vtist' I ti'ta is SitiaiteI qlhpiOl\hiwftl-l\ WIC11

showing 5 longitudinal and 2 tenior transverse ( o'Nta I lie lengt.thiIt flth l IItransverse costae. The anterior e'\( lIlllltni till- t ,tua pnLIc *lh'tit isI' 20 pill, Owtcosta is thicker than the second i anieter at theta iteritir t ransverse it osti a, I) Ptilltransverse costa. Note the twvist diiii till' I ,~tIdl appendagei IN I ' 1 111i longfin the longitudinal costae farrow) I it'st dimensl'iostS art' %%s titin tt range~l gi eni t it

and the simultaneous longitud- tilet, ;I I'de rtpttttr lAppleutlls Al lMantt i tlI .nal costal reduction (arrow). L M. ( )a es~1 9f7 1 . Nimtm tttt 1(] ( ),t tt' 1 979h) til ItIIa

17

sampling, and environmental data Cenera Itrends in the distribution in( ludi' an inc re,se In(diversity (as imeasuredl by the numbers ot speC iesptresent) f romn the nurt here stat ions to t hesouthern statitons and t he present v ot I tessspe( tes, niotably P. sociafis anrd 1) muluiannid/aaat mnost or all of the stat ions where (hoanof lag-ellates were re( orcied As metntioniedin 1 the~ oh-serva tions, sonme spe( ies A spic ibiera anrd (3,antc'nnigera, wvere absent I rom all but a tiv sta-tions. Ice station,, I and~ K hadJ not oniR morespec ies than the niait( hing %v,itur ,olunstnbut nmore' than anv other station, sampl~redl

Tecniji('rdttres and s,ihnirties tor the ss ,ter c ol-UmeII at the station% %,,here, these measduri'inc'ntswere taken (Taible I) a n he approximrated h\ t hesujrtac~e temperatures and Sal inities (JIbe to thewell i mixed nature of the Uipper water i okimnfrom whit I the sampiles were taken (I I-Sayedand 1 aguc hi 1979) 1 he temlperat ire range ot-2.40 to 1 70C represents a maximum differenceof 4.110C while the salinitv range of 33 2', to

Figure .38. Calliacantha multi- 14, 3',, represents a maximum differen( e ofspina aff. Incom p/ete lorica 1 1, The relative uniformity of the two envi-showing 2 trans verse and .5 longi- ronmental parameters studied prevents explana-tudinal costae and 6 anterior pro- tion of the distribution of loricate choanoflagel-jections. Note the longitudinal lates front the water column based upon salinitytwist (arrow) and the sirnultane- andc temperature aloneous costal reductions (arrow). Measurements of salinity and temperature ofLM. fce floe pond, station F. the ice floe ponds were not obtained in the field

The temperature can be approximated to be be-tween the air temperature (about 00 C) and thewater temperature, placing it within the range of

the posterior end (Manton, pers. comm.). The7 temperatures observed at the surface ot thedescription of the type of the genus, C. natans, water ( olunin. Ponds that forni on sea ice can doallows tor the reduc tion of the six longitudinal so in two different ways, either through solar in-( ostca to as few as three before Ioales( ence ait duced melting or subsidence of the floe and sub-the p~osterior (end. If it were riot for this disc rep- sequent flooding. In the case of the ponds en-dinc v and the piosterio~r twist, the organismns in Countered in the Weddell Sea there is strong evi-l igores 36-38l Would be very similar to (. natans. dence to suggest the latter case All the pond,.1Until further observations (an be made, the or- encountered were near dletormational featuresganismis iii Figures 36-38l will be designated C. such ais ridges where the topside loading wouldmtItispina aff deflect the flat plate floe so that the surtace

The key in Table .3 was prepared to assist in would be below sea level, allowing it to beIdentifying the loric~as of the spec ies of Acan- flooded through (ra ks or connections to thethcw idde found in samples fromn the Weddell open water (S. Ac kley, pers. comnm.) ThereforeSea. the salinity of the ponds should approxiniate

that of sea water or be somewhat less if snowwas incorporated into the water of the pond I hi'

Distribution and environmental ice vcore sample was not acc(ompanied by tem-tolerances perature observations, although the assumption

that th(' temperature, partic ularly of the brineThe distribution of loricate c hoanoflagellates poc kets and f issures, must be at or below tre,-

in the Wedcoell Sea is summari/ed in Table 2 by Ing (-1 80C for sea water at 361 ) would seemi tostation, Fable 1 presents pertinent geograpihic a, be valid Salinities of 10-cm11 sectlions of the core

18

Table 3. Key for the identification of loricas of Acanthoecidae from theWeddell Sea.

A L or io a possessi ng projet ions ot the Iloigi tudinaI( ostIade beyond the t ra nsverse t usta (e) at oneor both e'nds (it yes then 11)

AA Lomna not p~ossessing prolet tion, ot longitui nal ostae e tyond the transverse CoStd (e)Pars icorbicula '.ucialis

Ii LongitudimalI projections of tost de pre'sent on Is at anterior end of tori a, posterior end(harat teried by a i oa)i'sr Oer pOf the luirgitudiiAi ( Ostae arnd a adudat appendage

(It yes then (C)BB Longitudinal prulei: tions of (ostai present at bot h ends ofi torit a Crinolina apertaC Lorica ucomrposed ot tb or fewer longitudinal (ostae aod 2 or tewer transverse kostdt'

(it yes then 1)CC Lorit a ( om posed of 1ll or morire longi tUdind ira 051d aid (I or more transverse (ostae

(tt yes then D)1 Loric a coniposed soltely oh 2 longi tudinail i ost at' (it Iyes then A)V Loric a conmposed of foiur to six longitudinrat anil two transverse u stae ( it yes then 2)a Longitudinali Ostap spiraled or twisted about One a not her Bicusta spiniferaaa Longitudinal tostae not Spiraled or twisted about one another Bicosta antennigera2 Anterior projet tions 4. i oniitInuous with loingituinal (istae Ca))iacantha simp/ex2 ' Anterior prolin I ions br. (ont intiOUs with longi tui nal (ostae Cafllacantha multispiria

1) Longitudinal coustae' 14 or more. tranis'.ersv u i~tae i, largest diamnettr of the torica at the,interiuor transverse osta 'canthoecopsis spiculifera

DD Longitudiural c ostae 1 I or tewe'r. Itranisv.erse u ust at 4 or nmore. largest diameter of lori( a at thirdor fourth transverse i ota !Jiaphanoecj molitannulata

were, deterinined, and shtrWed a oisnutOf harao. tt'rst ( I (hit' slapt' Of thli ge1LIuS IS

b', dt ut deptfh of 1 2010 In)o below I tit srti( t' aR hireved b, hiaving a posterior ftrawo. t'rse ( Oslo

(ice tovt'fhiok ness wits 1495 m1) (At k lt' et al (t ht'r are hut two t ratn%\ers' ( istat') tor Ialer1 97%) 1 his, however. do~es not rvprt'seit th l i d iameiter't thaini the anterior t tat s~ t'rst tostd

linity of the brine pot kts and fisSttrts whiere' it is(Mdnton e't ail 1976, 1 horsto 1970) 1 lit numbetrhyvpothesitt'd that the ( hoaiiotlagellates exist In oft (oiiifttlt't ( otiSld strips is le'ss in ft'e posteriorthe, brine po ktfs and] tissurt's, sahI ii t' vatrie's (tLI t han the anItt'rilr transverse tiosta, "%hilt' thetto the' variable degree' oft frteing throughout IIt' length an d degree of t sa I strip o\,t'rlap; is alt-floe. In somie nof tht' smialIler pot Istts (W here the Iiroiititl eqiual Iii Plt'trasiga rev tI0I(si

great test amouint nf t rve' tg od b~rimi wtere I irondsen, I a hrrel Shape is itt iieved ii, havitogfound) Salinity extt'eds t hat of tht' watter Itron dii driteior transve'rse ( osta vvitli a largterth -whit h the it Ie wus frmned eft'r t han ft'e posterior t ransvterse' rIsti (tlis

slpe( res also lids (lV two transverse' nos Ide) I hisvarial tion iii osta I i aiieft'r is it hievt't Iby hiai -

DISCUSSION in anterior tostal strips ot greafer lengthI t hanoIhtist, of fte posteritor train%%t'rst t osta 1 ht'eor

Morphology lx'rs and dlegr't' tofl' theI stal Strip overlap art'I ht' use tot SE M as a toiol in) the' tiorphilrhio atl totisant I(I htrnist'n 197 0i) 1 n ( rioina isefhorden-

anld tdxtononi it aSpt't ts Of tis Study lifs rt'- ',is, th li k irt-shapt'd lorit a is at liitvt't by hiaviigv'iilt't that thte miainner in whi, If t ransvterse two transvterste t otat' ni ditfteret'n dianitttrs Iii(ostae( within ao organismt vary theitr dtiitn'ters is this I ase, howvever, a t tnttbtoattoti of t ostal Strip)

a ( liarat ter that is p~rttbably Imtportant in ft'e (s- length and degree of overlap is used to achievefahl ishnit'nt of a spt't it's n tint tpt for thet At an- tlls tONte t (I honlist'n 11970) Iii Vlaphanoeo( a pedi-thoet: idat'. I ht' vartaf ion tof ft'e t ransvterste nrstat, cellata I hrondst'n. a barrt'l-hapt'd lorit a is,Utiliately dettermi nes lori a shape, i witd'- Mt htt'vtd hy hatving two t ransv'rst' ostae withISprea~d (harmt terust i LJS't iii (lt's rihing gtetitra the s,1n11 ott nbt'r of transvvrs' n osta I st rips but

In Parvicorhictila socialis, P. quadritostata with a lesster t'gree tot overlap in ft'e St tinI hrondsen, and P. circularis T honisen, the ftransverste usta. T rainsvers' n otisal st rip lengthI is

constant (T homsen 1970). D. nialtiarnolata sent a signific ant extenision of t lt' geogratihi alshares this method of lown.a shape (ontrol rdinge of the touLr spc'cns. 13, spinifera h1is heen'ialthough it is somewhat more elaborate be( ause reported tfrum its type Ioc il it y thel Nurss giall

I tur (ostae are involved. I he exac trnes% of depu- Arc t i (1 hrondse'i 1 970a), tfruim tjords ot Noi% iN,~

sition of c ompoflent c ostal %trips is indic ,ited by (Leadbeater 1972a1), and tloim t jrds of t -niiil,

the differences il si/e .ind nmorp)hology nut only (1 hornsen 197 1) C apc'nta has bieein repo(rt ed onii

between transverse and longitu1dinal ( ustae hUt front its type loc d.ity at I. ornsallis Island in thn

between individual transverse c ostae in suome Caniadian Ar( tli (Mntuii tet il 1475) A. %pit Wuit-

ca1ses (Calliacantha simplex). I he cliftfereoc es in era has been reported tromi its t ypeI i '11I (alit

the manner of loric a shape deternmination be- thel Washington (oast (Norris 14015) and t rinl 1te

tween the genera (investigated to (late by elec - (oast ot inland (I hoiisen 1977) 1P. so( wihs hlastroll rim r0ot y) and the exac tness ot ( ostal been reportedi trcin I erre Adelie (l)t't l1indnt'

strip de'position indi( ate thait this ( hiaron ter miay 1900) inl the Ait it~i ti s wsell as tromi N iai~oln'ibe potentially implortant from a taxononc Ioi a Ilit ies throuM91hout tl' 11(b (lrld (MaIini etl iI

standpoint, both in assigning newv spec it's to a 1970) 1 his and other stuics itucic ate Otli( hitgenus or in reassigning spec ies des( rihed wit h- anu1flaigellates plrefer temnperate and] polar ie

out attenltIOl to this devtail More investigations 9101S .IlthlOugll sulbtr(Ili al and tiopic ~Id lvgionswith the elec tro in miros( ope will he needed he- needl to be InvestigatedI hetoi' tisl' c an bt n (I

fore the taxooioni potential (If this c hara(ter t inned

(.111 be fujlly apprec iateci, bot the results oit this I he spaitial diistrihutioii (t il(,ilotIagelI~ites

StulyI in( ate that the manner in wil Ii the Ioni- within the Weddcell Sea usevr a .i m ifuise ka Is lt

(a shape is determined may he a ( unservative overecl apIpromimlitely 1t0tti riatii if niiles inui

haram teristi( . .ites that lom it t hoainot I~igellaites %%ci' geii

Another niorphol(Igi( .11 ( (lnsidicraiticln that de'- 111vl proent. ailthbough in simalil (Ipianiiititns tw

serves further investigationi with rt'slpt( t tol its some spt c ns. 1it the tint' (It samp~linlg (t .ible 2)

tontribution to the taxonomy aInd phlogenv ott I he pa1(1ty it of n (licrg~iilisilis ina 11iidi Iie tie4

this groulp is the sue ranige (It a splc( it's I he (lit- truic p lai~tioni, il1i1\ inifir equ1 ipmein't sthtiIn,

hic lty of determiniing diianetc'rs and lengths" ot ity wlierehy soon' ot the( siriill'r oig~mihsis dlip

two-d imensionalI represcit.itlns cit thre't-(linltn- I hrucgil, (Ir iiiay e l t' iis'd NI(l II ton ( Is m(il (Ii t

sionfl) strut fuii'5 otwittllading, the siit'o (htet the less robiust ttlils Jis a resutlt o)t inl' ptrst'rs a

loric as cif a Spec it's seemls toi h~ive ani inverse rt'- lio Iii nd prepairat((int' le imittocs Additi na

ment in whic h they are tound (Manton t't 11l (ket'ping in mind thc' vtt( ts (It c ollic tiltit tiv%'

1976, Manton in(] I t',iwt'.tir 1978) 1tIns, is t'rvaiticl, andic iprt'tIiititoi tt' liniques) (t)I (hu m

spicalifera. whic h has ai ttlemt'ratt' typet ht ilm p'tt' ilc tlurt' (it thet lhiigt'(tgIi~iphv ssll begin it)

was c onsidt'rably lairge'r thain the' type s 1t' ietmerge Iihe pre'se'nt' (Ioft hoaint laigv'lkites ((,.i

men; however, 11 spiera in(] C j;perta welte ai vast, ailbetit relaitivel'y uiilin .it'i iiidi itl's

toisieltrbly sriialler thain the type spIme illhtn~n i is'(r~riil l~Vlt sltjtl

I he diiscretian(: ics betwee'n till-' si/t' rainge's tit ilt' taii, i ertaIiily 1). soc ia/i is uliclcilows It stst

two bipolar spec ies are difficult to comprehend. thtert' is a imit v (It r'eport s in %,%Itt li( h ( ilt t lkig

Seasonal fwinter-sunllnn'r) thris oft Pairsi(o(I t'latts .irt' pret-t'it .is part tlt thet tlmi Stpl iid

hictila sotialis, with clistinc t and diittert'nt silt' smillpling tt'o ilIitcei' 11incl (mn istees oilit'

rjng's havt' be'en obse'rvedc in tht'- Art toi (Ma1ntotn 1wl tt rt'sc.n hers ()t tivt' t'Sst'ii v 0 It ( ict.iilo

t't all. 197b), and it is piossiblte thait this sornt (it f1li'llts wollead It ) .Icll I ltiitn tsiil ini tit11lt'

plhe'nomeltnon is re'spocnsible' for tih' cist ea ni le~ ts stiilt'snotted here It is sulggested that1 hld a si/i', .il-

thouigh an easily dis ernil"t' haratt'nsi with'the light mic( rosc ope, nlot be citiliit'ci tt dittt'ntii- Habitattuatt' spc'c n's, at least ciil i more0 ( uiiiitin'l'i The assoc: iation of c hoanollagellates posse's%-

sive inc lure t'mergt's ing caudal appendages and] anterior prociec t ionswith tive st'a itt' has be'tn stiggt'st'd iI Maintoii

Distribution t't ai[ (11)71)) It is tliitting tot .irvt ait thet saimet

IThe rteports cof B. %sinderat, (.ape'TltI A. s/i( il. ( oiii liisitti n trlt' Wevtddell Sea~ whet'i it is iltlt'(

ijera, and C. simrplex from tilt We'clcitll Se'a retire that six (It thet st've'n spet( it's rt'titld( tili this

2(0

area possess these appendages. If this (OMi l1.- ev~apora tion and pretijiitat ion All w~idein e

sion is valid, then the present. e of (hoanotlagel- troni fiVld 't~idie' iidit ate's t hat, as a gr cup lit) hlates in the water Column could he thle result of anoflIcgel .ites have g.te'a tii e iroiimeota liI dc

seeding f rom the ice. Breakup oit sea it. e along di(LsWork With Iitrga 111511 ii (Lritift' (oul1(the open oceano/pack ice interface d~oes O~C Ur nf iriirnot only ge'neralI pa, terns lint .it( lo(I ler-

and is responsible for the release of d~ense algal ain(tV le'vels teir id1% Idual Spit- les

populations into the water columin (A(k Hey et al On the other hand tit', ad~ anitages ot iinhabiit1979). Presumably this seeding (ou Id he a iog flte it e art' twotold I he it ev prirdtes a spai

method of introducing choanooflagellates from Jiijy stable envirnniient l ike that inI %hit Ii li eiii

the sea ice into the water coIlumn1. Quan~titative' oflagelIa tes have bieeii totiitdII~ iofeit t'ii11iit

studies of ice algae and water columin algae presumably pireter and III whu 1i food Is thouightfrom the same areas iniit ate that there are spt'- to he abundant I he it. t' oJi1nniooitv, ;ire(oilcies which not only prefer the it. e environmjent, riantly lieniiatt' di'ittnns, (hia riot lagel lat''%. andbut even flourish in it (Ackley et al, 1478) No bacteria, is trapped within the ice until, in theconiparable quantitative studies were done withI ase of the Weddell Sea (niui 0r11MMN iysiVI a I orchoanoflagellates, largely bet.ause t t o(rate, imechanical breakup wrought hy the winds,niethods for enumeration have niot beeii devel- waves, anid t ides rt'l'ases t heii Not onl., detsoped. This limits the extent of our abil ity to cle- thit tI rovidet a substrate III %%h Ii t flit,' the almfine preferred habitats or ( OW ludt' whether the f lagel lates a n grow hUt t11 lie'alga staiidiiig ( 1 0i

Acanthoecidae observed in the water ( oltii is onst'rvetl within th lit i t(iut il -tit Ii re'le'ase otare autoc hthonous or were introduc ed froni the u trs, predormiinant ly along thle i( edtge') ri'sulItiiigic e. Choanoiflagellates in this study art' de'fiinitely in high standing t rops ot algaeV (in11XisInIinlI t h114)-ic e-dwellers, as there is rio tither iiie( hanisni to oplVI a Values raniget'roni tI1 to 4 4 migoi frallow for their transport from flte water to tflie' t host' (ores examuined) (At kle It' t 19I~78) Al-iRt' (with the except ion of init ial f reezing) Quan- thbough lit tl ottr no work has bet'i tdont' on bmi

itative studies of saniples otainred fron tranOs- tt'riil populations withi() the' it t', givenf the', ighects perpendlit ular to the Ie edge WoulId Vt'tab- toncentrat it)1i tt algae and thlit re'sti Italit dislish whether the c hoanoflagellates found iii the soilvedl orgaflit matter, It woo Id Sei saIte' t0 tdas

water column were native or riot. sumie that they aire high (hoanitt lagel laiti's te't

Ice is an et ologric il interestinig habitat for n bat. tt'ria (tevadbeater and Mortoii I1 74,u, I Ilbs

choanoflagellates for st'veral reasons. I he (hisad- 1930) aind iiay lit able to tise Ihe dissnlve't or-vantages usually aisso( iatetl with an I( vt'nv iron- gaim matter dhire't tly (Gtld ttaI 1970) Siebti thment include freez ing or sUbt reez ig teripera- et ~It (1478) plate t' hoanot lagt'llate's in ft, ve o-tures and osniotic and nietahol it stresses riot logk ~iI tate'gory tot prtitt/ oplanik ton, a group~normally encouintered in the tope'n ot tan. Th li' tefined as being unit el tlar and( possessing atemperature in the ice Cduring the austral winter, si/t', nietabhli at t ivity, and] div isitin rate simiilara period of time during whi( h (hoanoflaigelat's to phytoplank ton althiough they art' iiie ltile' iiihave to survive if not grow, ( arn he at( ulated hmy thlit arinmal k ingdom Preito otiplarik In art' t oniassuming a linear ( hange be'tweenr the watt'r sidered ht'ritt wil to the( phyvtoolank tor be'( atise'underlying the I(-e (about -1 80C) anid tht' air they re yt It' the not rilit s storet iiiflthe bat teriai.tt'mperature, whi( h during the wiriter is, we'll whit [I livte otf the disstlved tirgait matte'r aintbelow freez ing Assumiing that the itt' was 2 iii dead]( tells of thle algae' C hoameitlage'lla tes prot-thick, water temperature was -1 70C, air te'mpt'r- vide for a rapid t ormv'rsitul oit bat ter iail liiniassature was -200C, and the heat (onduI t ivity t0 tt nitt ie'its A tightly e oupled tropim se lie'ie'it.e is constant, the'n thme tempe'rature and salinity t'sists in tlie' inte'rstitial po~cke'tso cit e ut'ihm30 cm from the bottom of the Ie' floe' are the I(e', at least uintil re'le'ast' to thit' o V'ari ott Lifs

-4.45 0 C and 44.7"/,. (Meguro t't il 1907) 1 lit A mitre t lassit il tuephie s( he'oie etwe'e'n batmechanism with whit h theo hoantdlagellatt's, fe'ria aid( larger nme'ta/tmnplarkton is poestuilatedtand ice algae for that miat ter, (opt' withk the's' for th lie htiariot lagel la tes in thle waiter e e ltiniii Itlow tempetratur-s arid osrimoti stresses riluis Jit~jii',rs that the', disaidvanrtaige% ot bie'iig sse( Itunknown However, t idet pool orgaisris, IM( ILitI- Mierl with an e'nvironmmie'rt wit h vxe'sIeis eel tei'i

Ing those' (hoanotlagella tes ft'(o(rd('( by Norri% p'ra tirt' aind salinity sire (uteelt ighedt he, theii po's

(1965), must t ope with gre'ater extre'me's iii saliii- e'no e ot a substrate, aot k nt preeatots. ~inel theity and temperature nitdiatedr by period% of availability cd food %eetrt v's I his It v' %Sfstiri W itti

21

its three, nin ( oilcni 'flts deserves miore atteri- morphological and phyletic relationships, Al-ion than it has hithevrto riieived, keeping in though the advent of electron microscopy has

in,,n( the i'lm(idactioii ct ibove-invnitionied ri'ia- greatly expanded the morphological base by in-iun)Ships) and t heir ouanlt itl iiat ion ( reaising resolution, depth of field, ind the abili

ty to dits ern suitace detail, it is hoped that moreemlphasis onl ( onservative feaitures swj h as ( (is-

SUMMARY tal Strip shape, relative size of the (ostal %trips.anld method% o) ( ont rol ot loriw a Shape will play

Conclusions greater roles in establishing t his morphologmi alI choanoitlagellates were wi'lv (list riliutecl base Howev-er, the need tor light mu( ros( oly~ to

throughouit the region oi i t he sea I( v i'd go oif t he ide ciY 11v ad enu ni 'rate heild ciamples shiu inot0)

Wedde'll Sea I Ight spwi ii's, Im hiding oni' de's- he forgotten

ribi'd tor the( first time. were observed fit wditiroltinin and lo v theo sallis2 I1 ilirmii t of the low temperatures and LITERATURE CITED

high salliitic's, ( aram teriotio )I the Weddell Sea( e ( nimutl'. h all1 eight spi' le's has beenl A( kl' 5 , SI t S I19I mtIll ai~nd K Bt k 11978) Pfimajr produutiv-

's tabIi shled itV i sea ie o' ot hi' Wedde'll re'gion U S ArmV Cold

1 heit gei grai lii( al ra ,iii go I i ye siw w s ot Re'gionsi' Resva rc ti an d I nginec'ricrg t chrbocrv. CR RI I

irit hi ii( id a e (( rif1olina~ aperm. Px0 V orbiti In) epr 78-119, 17 1)A( kl's. St I K K t111(k andii S Icigur hi 11979) Algal standing

Ntii [is. f(i (isa Ic i 11ter& Ha ((05 9,1 i tiigrlg I ( r of) thei %e'a iiie oft tih' Weiddell region Deep Sea Re-

ci 11(1 A ( lo~tht-~ii i)J., Spa1 uidera) hacs beenl sigmtf I- secio-h. Vol IV no 26A, 1) 26,9-281.iiit Is i'\iciiiili'(lt tLire (; t9o) Stir Ia reset v' de' Palrvoorbic Wa soc whis

4 I hei nii'c ha nist i s hap ii ont nl ot the lo r t- (enli' fj (fai'r% t, cr5ii ccirton (le- I Antari tiquc'e clrr'ni'\ h ( fI'r~jlVandpoW I'liv isetil1%Ade')it') Rvc's io lgi lique N S ,voil 9. 18 18'8;

ci~~~~~~~~~~~~~~~~~ iHcis, ni'vmierdpoittclc i'it~ i b W N (1910) Heetr i'iinisici( li' on the ( hanrftcgi'llacasuoi(ic l cii o liircu ter I(( redurciiidinecs) (t I'shwm (r and marine) An-

5I hie s/i' cit oi as is iSJuidgied [loti toi bei ai ali' de li So iiec' r toologiquei de' He/giqui'. Vol 60it

goodr~ taimiiiuiiii licrcii ter bi'i risi' cit thel crii-i 49-88

liilitv' s's thin spi'i is re'torte'd fotrii difterent I I Sa~qvd. S / rail S tlcgui hi 11977) Pliyioptanhion st1dri's intihe ssatir r nrlui amnd ptii k ire v o the We'dde'll Sea

airv'cs Wt the ssiithi Antar it Iiirnrmi of the' I I S .vcit 12. p) 15; If,rvxsill, (, A amnd. (; R ftli'l (1970)) tOratorn r ti'ing andi

RecommendatIIions for future work moritirrtcg for light rind le lo ion muri r iy tlvransau -

It ,IiaN i' i coi luiuh'il trurm ( (iniiiliiii iif licon, oi the' Arnu'rcan Mir ncsoicaI Soti cety, sol 84 1)

videm vl'n froiim thei pre'sent swo rk imnd other 4#0ri474

stuit.li' thcit 1I citlrigvI'lit('S o th i' I ciivi (,old Ks ,H NiIl'iwrsl nod V H 1 I noon (1970i) Asi'nic t rtiliation and) r'lc' loon mii ros iipN ci two, p( it., of r hrccn

Ai an~thiii' idaii ar', in giieiral. iismiiin~oilitcti, mligilicift' Jouirnali in )'nuniciooloi1 , voli 17, )ci'(irhriliiii indi i'iirsi'thIi'niiii ciid dwec'ill iii iiici 2110212

ma~crinei hcmhiitcts hBiog~eograiphN. spcitiail iistriii- Itihherd. I) 1 119751 Obsi'rvaions cn thi' ulttastrut ure oi ihi'

ii ion, enV i roii nin 'it i I toi ili vilis, r5Si ii itfn ((ri hod notcidi')laieV ( odos ga hotrtis 11 hr I Savi lle' K ent

us ith Ii' toi u4 tint. (olflii irMoriiolimgV rIid Iiii-with spec iaiI rc'iere'cue to thi' Itigc'tlac sirtj ur jun ouronivof he ( inhou, iavall ddrsse tonalof(e( i'(e. vol 17,1p 14121ll

iin~ict lii'Ac ~iitlcii' dci', ,ll ~ddmcssed iiiihhird, 1) 1 (1976) the U~ttliratuori' and taxonoiu of thi'sririii etefnt i tisl, ri'pcort, ri'v'd Itirthir iiyisti I.hrVsriphVc eat' and t'ryni'sriophv v'at.' (Hlaptopiv

gaili l)i'vi'loireiitt utI a imiethod for iiiniiri i'ai'l A tjrrve'V with somne 1cw cOhSe1 Vdtcrns on the' cil

tion iii ciii i reasi'd cwi's's Ivitcs% onf hi' liar) of crastnirhire cit the ( htvscrjh~tve Hotanr( a) journalri's'~in hi's ott~n'pni'i'ti ic i) ihii~iiiit lcgo' 01e hi') ocein 'Sot i'ft si) 72. , 961180

rf-e~ri:hes I tw recluv f (Il,11oligllI i'.idtiiaiiir If S ( ti72a I in ro tuftra) chsi'rs afiircs inIcites, anid wi'esrea'dc srmnrpliiig Inll 11geccgral)ii O[iM.i 1r1111cni' A tiIirrleit~t trn theii c rit ofi Nric'gicins jrjl sijsns ,ci(l mv~ini v' muir uiid'r- %s,,V I)iccnici ot thc' Mainei Hooluogcual Arsiton of

stanmdiiig ot thi' biiigi'graphii il &%itribilitiiii id (hei' te liiikfnhrdin krl ',,1 1) t7 79

this grip A miurei ( o tiri'ln'nsivi' 'amntiocm I i'ciitii'ti'f If S ( )tri72lIl Utitcitruc Itirl rc~o~tii'rorcs n

cit tfill' cliporitmin'is o t 14 t 1( ve ( umlniiulit' " Ith su01M.i rcacccci i frccccig'lie' crom thi uast cof tDe'nciark 11nls h Mh ir iligri a) joinnacl. srrl 7. p) 19% 211

thle goall ii uinide'r'taniiinig its trerthiiil'sicimiii s is I ea'rrleti', tt11 ' 17 t stivrnal mcirphcrlogv (it some

All totuimertiili' itist hei hicsi' illl p Iii tint stia i hstn ho P rnolistcninlic. Vcol 115, pc 2 14 2S

Ilttci iifi Ihai kgrccmnld thait is Ilselt icise i'l i I oce'ciati' It' S (14i7S) A iii( rcisil riir a str ci th' nianine

22

I onoII Iiiiit IM latei s% II (I m mic ropo (Nor ris) I omnb *I' iii411 ftill fure Iivvft~wIiplii iimiphrtIiieit, ((I

ment fin s naoot a anl~ d S teptihdnie(ad iip/oco~tdt trdi tin l nlg a uiih.i.nd Otaiouug. 1 h% %01 2 1 pI hII I'titoi'la~i~ %il It I p 111 2012 9 21,4 1 ti

I idilledtiq, It S ( 14771 ()bsersations ilit lt ie thistifr dnd I homiirs, 11 A ( 111i Stuijh, in mdliiiii iitIiiI~t~

ii,'ia 5iit5)l I IIs Journadl oit the Marulie HioougitdI (pheibu %o 12 p, 1 26i.\,ioi t.ilui'ru O~f fill Uitedj kiungdory. s-oh 57. p 285- 101 Ihm n IiiIisii A (11q7blStoivie iiiirleihirii detts

I t-ii.jItat. It S, ( 1197811 tReniidiing ot iijjafplfgoet- senl5 I lilt. 'flit, lord1 i,,.i tiais l n, ui it oe ItI ed ( Iii iiii

I .,iuot1,i 01u:iii uth m'atruine HisihogucaI A'iso w~tin thligei'tvii fromuu 'st old (Deirnmark) fi( ludiunu flt- de,'

ot the tiniftongdot \,f~I~fi of 581) ' It 515i 1, option ot tw)u iu's spul ii's \iiFn'gwn Iiiouti of HotI i'.iiiii'tiV 1; S ( adi I Mhimitiii ( 1474) F'retiinarN ohsrs- .ii I i 2 ;1 11 iI

hlls oni flit, I hi'iiltri anid bIltug ii thii .m ai .Ia I iuu wiii'i i A 11077) St'itl' 1-i1 miilli t Ihiaiot Idl'lat'I, otlirii tlagigi'hI.ts vpeh.luul' .1 diluco%t~ita I II, IIll vni'i' thin tilt( ri, ls itii tjl v Sill i'S iii i gillus

Iiiuurfidl it th,''t. iirtrii' thuuhigcai Asoit ioiin ilf the A( anlthoiu' of)SIS -Ai( 111 flritst 'ikol ui -Ik 11 il I111 p -

Imniteu kinguliur \O Si 4, 1) 26u9

276 811"1'

I i'diwai'ter. It S, ( jIdii I. Motuiin I 1'74a A light dnd e'Iu trni I iindni, I I 11'-mI I Ijlgi'Iliti' it Niiuvuigujjuri otji ,is t .t

IVM u-4 OIWi stLuu' Of 11 thu O,iitfIfI9VIIdti'S ACuIu N it flia~g.suin ?(i hoiiti(k %0 it, 1ii p 1,1 2lt

thiut i spv' tabdo, I IsI ainid A tits ptiul I Ili, Art hi Iionulsin 1 I I1471ta %.u/prgoe' i Sphhirweri ,, iii% j lit'."

flit fttukniIii'Iigui %ot 95. If 279-2921 pla~nk ti'li spi', ws Ofi flit, r,t"iO'iutiil i'mu rvi iil'i

stuuuli 01i .uilitrin.,' ,ii , ii' t ( iiilisigau lamei,s( laurkit'tl

hw i oiiru liai'eli I tI li. 1,1i' dlreturirr I toifile Ingi's I hruinds.'ir I t 1470tb) Madrie lldktii Al dtnii t va'.n,

it% uiIi b 1 147 Wtor Htni kk oI~ 17 1) 10 1 111

r,jllti. I .111( it su ( I ividlieater I 111) 'ioilel t Wiaal .1/(11. I Iifiwiii' I ( 19741 I'laniktiiiii , Ilunurnit uiIIati's triur Nuirtl

or ( .,I,.u .iithai Ivi'uutiuetr I( iiuidiil IgiIlttli Prio

Ii'uding, iof flit, Ri'iu/ Sun #(-f ot I unihmn It . %il 21)11 p

411 57

Mntoiun, I jd( Ks ( )dlet-( 111711.0 1 kifthur- Oihsers11nins (ii ( lilt

.iiiut tdgfi'Ites fi the gino', C alwuinuu~uthau I i'.dhi'dter

f lit spvc Ih n'ti'reu iv toi ( fhiltisnaih~ 'u) Nio% ftoii

Souuthi Altoi, Ii adilini juuuialu o the Mie luo/ugiI Il -1,%oi iairu ot the I rugged kinguhi %oIi 54 1)

2017 It1Ma~uritu I unit Is ( Litus I (1174l I Lather iiiuuers-,timuis Off ( .I-

rIti'ri'iii 1'. 10 ( ui~~' IIpe fruOM nudui'iN 1wit5 ifth %%,)F'illi

Ploti veiuig, it ft(e !Ruiiai %ou( uti uf IiuuuiiiuuR. sii 204.

\tdihiuiu I I Suit1laodiru .uuii It (I vi',iai'itr ( 1117S) 1 uuit

iii'i* %p-' w% 01 iiuuoiiut IagviIIatv%' tlIi Fo tr it1) 111I'ldl

Prt'ii eviuigs ot fhei Ro%iuI%,.1 if ot I iiniluu H %oi 119.

Ntianiomu I I Soluli'riiut andii It SI( I i'iuh.titr ( I9 7

6) 1 i

Owti ,I)'u II, .i''u iuit orii ' t wiv 01 mint hI u

Mi1.1 pg i".I ,iuih( u ac-I'liduiu , it iitin ut 4. n

14 2 It'G,Mt'.-gur(. 11 I fOli .11iu) It Itiluiuhirul llm 117) Botitioim ti t'

ptituiih, t I,, it flit-' Alint (),va i'ui Aitarun lii A'i .

suitM 21 I 22W7 2-71

uiiir-iii figiuu~it .i romtt Nvis /va'.i ill lii iiig flit,'

d.ii Hiut.il sit fo Ii. l 1it it- v%/S11djun

Niurris R I 1 t114,5l N.'itoiuii mriivi ( r.,stetiiriinauI,

hiiiifidl iot Pinhtii/miliigi sui 12. 1i 58'l112

Nii in lds N 114'76ii I Ilisns lihiS off %dIliintltiie .19)(1 itute

I alulds' . u % niiniu j) s o% (i i rad phi ( Vlaeii'i

Pritish) Ilimilogin dl jriurn,11 %,Of 11, Ii 1 1 17S wboriith I N. VSitii

meli n I eft/I 1 0781 Ijt23

23IL

Mi

APPENDIX A. TYPE DESCRIPTIONS OF Cell Solitary, r ollarlt'%S, egg S11.iptedl t) I Iimu

ACANTHOECIDAF COLLECTED FROM THE (,Illy ovoid, with single rollar. tiigt'Iltili mig~t.,

WEDDLL EA.about 4 timers the length ot the' fell, inu ho'. a

terror; lori( a exrtrunir'ly liyalli'., nn[ less

g.obl4t-hapd)( Withi 2 [or( isloll,l115 i Illiiti

Crinolina aperta runnring longitudinal ridlges sirndiig slighilNls it

Original description: Leadbeatr'r, 19795 rally onl thll outer surtme Mid11( I (IItuInnllr .11)14

Lorica barrel-shaped, open at b ot h ends, Witlli ciily a% slenider ( Urved sp~ile'., lwtc11114411. t14

a long fringe of slender projectin~g 4 osie around Ig to make I long ;ntsii() jrp' 11. II

each opening, Overall length 45-50 l.111, width at length 6-9 pi (lialnetir 14 pm ,11 ,i Itr-igihI

the anterior end 17-25 jim, at the po(sterior end Withi anterior Ili(] postir t.t piI4 wit, 'itim

25-35 gym. Number of longitudinal (ostae 12, pill Habitat. the sea In tile. 1( 11111 i. lit-.1t

each composed of 9 or 10 costal stritps wit h over- kIandl, 74030)'N, 2501 (11 50(1

lapping ends and with a firmly ( onst rU tid (0'sti

located at a distance of 2 (ostal st ripls f rom Ihc Bicosta antennigera

anterior end of the lorir a, and another hx r(le I Original description: Moestrup, 1979

or 4 costal strips from the posterir r end] I reins- Solitary, protoplast c o t, orlng. wit a .ingl(.

verse costal strips L-shaped, taperedl at ()i)(en i'i~ nterior flagellum, 16-18 g..n lonig I lie (t-1l I(,

and sharply upturned at the other, those fron rated Ii a lorir . (ompo~l~e( it of Vii seven i

the anterior ring thicker than those frm i(.e ns strips. Iwo r ostal strips, (,11( Ii 8-( lo ng. ttliiit

tenior ring but both tli ker than the longitudinal the lorir ai ( hamiber wil Ii I ottiinis iII(' tbnltl4

costae; all upturned tips Ii bo1th transverse I os- plast Anteriorly eam h of lite strip'. loins %%Ilia

tae directed towards the anterioir e'nd ot the lInn long anterior spine, 12-17 guni jlng i11( inI'nle (it

ca and each individually atta( hed to the I onet- two ( ostal strips 1 hese ditter it, hlngth, lite 1)1)\

spondling longitudinal r os14 FProtnplat hoi ,ife'd inial being 8-il) pill jlng, ti4' dis'll, Mlho 1r

flagellum ca. 40 prm long aind a ring oif aibonut 501 ni Posteriorly thel strips ot tdiw norIi a I ilailillel

tentacles, eal h 01) to 8 pilm long. I yp, Im alit\' overlap aid( toin witl, i poi(4'.rIlll 51114', 1 25 pn

Resolute Bay, Cornwallis lslaiir (lit 74140'N. long, madde r4 a single r It st, lti t.1'irgli

long. 95 000d'W). nof the' lrri( a ( . 0 piltype mm rrigriii. igrirt' 41 tftoint kiktomi.

Acanthoecopsis spiculifera New Zealandl (42 0 25'S, 1730421h) 1 1 tNtw lifedw

Original description: Norris, 1965 1974

Cells with a form typir al for I lialla~tigt'l

lates Cell enm losed by a brim a ( I oipls trrti (alliacanthla simplex

manly of 14 longitudinal I ostae, I osn'ly env'l- Original dles ription Maioll iI()i'.

oping cell bo~dy, spreadling slightly at I t llir ft 1 9791)

form a broad r hamnben emf Inserl by widt'lN, I ori: a Wilit tour ,inteninr spiniies.,i .14 titii

Spd( ed ( rostae. Tips of I osta' free tromtili li 0 otus with a lon~gitulidnal I (45tl, ill' l~ii'i I f~illlj

other, two on three trarisvi'rsi' I olavil cm1 ir lilng Ing without mnnnm al luhnge if, ft(ll (iat IIt .

the longitudlinial ( ostae at lit- i' nt of I iipit'd li,, 4 hiin11ber Bhll transverst' I t~tlit,t lwth 144 111(

(0's anterior end; two) or lilt('-(,tolr traiisvv'lse pllultliniate n ostail strillso llt hitItllgiilliil.1

costar, en( irr ling lrrit a just abolveI' Il llast 4ota and,11 ir not rlirv' tly Itt if,' ftiiiii tillp,

Base of lorir a attenuate, longitudlinal 4ttiwo' forminrg till' siirl' e'nds "\itv~tii Itligill tttIl,

(onverging 1t) form a stalk up to 50 pinItlig ante'rior spun's 15 jini S' pill, till' 4.i'14

Lorica 1S-18l pm long. lori a odyl 8-9) mm long, spint it) pm 4 4 pil ( )vr'rill hin. a ligtlr 411111

lori( a neck (spines) 8-1(0 i long, Ill-( k dia lilt it) mlonly 15) piml 1( Pill, Wil11li II I lijiillbitI t12 tint1

10) Mm, cell dia 4-5 pim Dia (it stalk ,ipprttxi I~ piml but all1 parts 1 .11) heiitl'llrgt'l ill(IltI

mately 0.3 PiM am tin (Inditin 1115rm %iiillt 1,m il 1111 0111

Type locality. I I* pool at I orui Is , Salt I lan tphe'ri aindrl alaipigtts lsiiolsd). sp. ilill11litt lilt'

County, Wash latte-r type, Wilit .,r1 dvV'19fgt tts.'ral lirigll "1t 211

Biicosta spinifera I he rniuber of tf-l'nmi I'% ,ilIMsIJlI'IVl Ili I laIg

Original de'scription I hrornls'n, 111701,1 elar lungth vairiaible btut I Ii 444'11 1's1te 1t1

25

i-*iZS i, PA M S A.&ama- 1 1u"

ot the -pine"

' idt, in tidtal waters IN pt PI pe( iientf i ii

let ted 1') June, 1475 in South Alaska troi the seasturlte in mid-channel about a third ot thei %%abetwAeen hoiiie and St Augu1.tin1 Islandl. % atel

%terN lean Also re(otrdeci in other samnples Irion)the saint, area at depths trom the surtau t, to 1t) inMtid at %va teniperattres 0 10 0(., also in Nm lhAlaskd IMI uidr't'J ii.e (temperature 00C), in art tit

Canada (temperature - I0(j) in Britain (teipt'iture 1b 51Cj), Galapagos Island(l% tenipu'rtIre'

220C an South Ain.i a

O)riginal dest riptin. iMvantoii and O)ates 1971)tI ri(. a 19- 14 p.Uf long, loit a thainihe (c',

iluthing s.pines) 8-15 Pin long Spines (oin(Ioiil 1

hut sonietint's 4, 0) or em\ e'ttioitliN 7 1 ongilti

randtiin dtipli.ation% oir tielet ions attu' tingsingle 1o( i) Anterior trans\xtre ( (ota lirmNiI at-tat hed to the spines inear thieii lIo%%r ends, the'

posterior trainsverse (osta thinner mnd no it'

lightly attat hed near the m)iddle oIt the' longitudi-nal i. otiavi The ( impoilent strips~ ot bo1th trans

verse, ostae, e\attlN bridging thet tlisf,imi v' it,tWt'VfIi longi1tudindl t ostavt Ntiiiit'it ilrt'dIm tionamon01g lon1gituinal111 ( ristat of ( urring at the' hin1dend ot the thamht'r as in) (. iatans. IDistriiit ion,temnperate' toI~ttl waIters near t it, stirt,it v' ii

both the northern (Potrtsnmouth) ,it ind t Suliiheiiphere(('ape I ti) nII ~p spet lint'n Sptil111en)in I-igut4 tollet( t) INo\emht'r i972 littwet'n Robben Island andI Capt' li,~n to idet pt h, sea t enipera t Uri 100CO

26

A facsimile catalog card in Library of CongressMARC format is reproduced below.

Buck, KurtMorphology and distribution of the Acanthoecidae

(Choanoflagellata) from the Weddell Sea during theaustral summer, 1977 / by Kurt Buck. Hanover, N.H.:U.S. Cold Regions Research and Engineering Laboratory;Springfield, Va.: Available from National TechnicalInformation Service, 1980.vi, 26 p., illus.; 28 cm. (CRREL Report 80-16.)

Prepared for National Science Foundation, by Corps ofEngineers, U.S. Army Cold Regions Research and Engineer-ing Laboratory, under Grant DPP7b-15251 and DPP76-80738.

Bibliography: p. 22.1. Antarctic regions. 2. Choanoflagellates. 3. Ocean

environments. 4. Plankton. 5. Sea ice. 6. Taxonomy.I. United States. Army. Corps of Engineers. II. ArmyCold Regions Research and Engineering Laboratory, Hanover,N.H. III. Series: CRREL Report 80-16.

i ra . o . . . ..