EARLY CAMBRIAN LINGULATE BRACHIOPODS FROM GLACIAL ERRATICS ... · early cambrian lingulate...

EARLY CAMBRIAN LINGULATE BRACHIOPODSFROM GLACIAL ERRATICS OF KING GEORGE ISLAND

(SOUTH SHETLAND ISLANDS), ANTARCTICA

LARS E. HOLM ER, LEONID E. POPOV and RYSZARD WRONA

Holmer, L.E.. Popov, L.E . and Wrona, R. 1996. Early Cambrian lingulate brachiopods fromglacial errat ics of King George Island (So uth Shet land Island s), Antarctica . In: A. Gazdzicki (ed.) Palaeontological Result s of the Polish Antarctic Expedi tions . Part 11. - Palaeontologia Polonica 55, 37-50.

Erratic bould ers of Early Carnbrian age , occ urr ing withi n the Ear ly Miocene glac io-rnarineCape Melvi lle Formation on King George Island (So uth Shetland Island s), Antarctica, haveyielded lingul ate brachiopods including Eoobo lus aff. elatus (Pe lman, 1986), Karathelenapuru (Kruse, 1990) and Vanda lotreta djagoran (Kruse , 1990). It is the first EarlyCa mbrian brac hiopo d ass emb lage described from Antarctica . The faun a is closely similarto that fro m the late Early Ca mbrian (Toyo nia n) Wirrealp a and Aroo na Creek Limestones,Flinders Ranges, South Austra lia. The new family Eoobolidae is prop osed and the typespec ies of Eoobolus, E. triparilis (Matthew, 19(2) is re-figured. The evo lution of thebotsfordiid s and acro thelids is disc ussed in the light of the new materi al of Karathelenapuru, which is intermediate in morph ology between Botsfordia and Eothe le. The superfam ily Acrot heloidea , is referred to the order Lingulida because of the similarities in musclesys tem, shell structure and deve lopment of pseudointerareas.

K e y wo r d s : Brachiopoda, Lingulata , taxonomy, Cambrian, Antarctica, South ShetlandIsland s.

Lars E. Holme r, Inst itute ofEarth Scien ces, Department ofHistorical Geo logy and Palaeon tology, Norbyvdgen 22, S-752 36 Uppsa la, Sw eden.Leonid E. POpOI', VSEGEI, Srednij Pr. 74, 199026 St. Petersbu rg, Russia.Rys zard Wrona, Instytut Paleob iologii PAN, Aleja Zwirki i Wigury 93, 02-089 Warsw wa,Poland.

Received 27 A pril 1995. ac cepted 15 Octob er 1995

38 LARS E. HOLMER, LEONID E. POPOY and RYSZARD WRONA

CONTENTS

Int roduction

Acknowledgements

Material .

Systematic pale ont ology

Class Ungul ata GROJIINSKY et Po r o v . 1985

Order Lingulida W AIIGEN, 1885 .. . . .

Superfamily Lingul oidea M ENKE, 1828

Family Eoobo lidae fam. n.

Genus Eoobolus MIITrH EW, 190 2

Superfam ily Acrothe loidea WIILCOTr et SCHUCIIERT, 190 8

Family Botsfordiidae SCHINDEWOLF, 1955

Genus Karathele KONEV11, 1986 .

Order Acrotretida KUIIN, 194 9 .

Superfamily Ac ro treto idea SCHUCH ERT, 1893

Fam ily Acrotretidae SCH UCHERT, 1893

Genus Vandalotreta MERGL, 1988

References .

INTRODUCTION

38

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44

44

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46

46

46

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48

Cambrian brachi opods fro m Antarctic a are poorl y known ; only five paper s have been published to date(SHERGOLD et al. 1976; Ro wELL et al. 198 3; POPo v and SOLOVJEV 1981 ; POPOV 1984 ; HENDERSON 1992),describing M iddl e and Late Ca mbrian asse mblages. The Early Cambrian brachi op ods described belo wwere firs t recorded by W RONA ( 1989) from erratic boulders of limestone wi thin the Earl y Mi oceneglacio- rnarine Cape Melvill e Formation of Kin g George Island , So uth Shetl and Island s (Text-fig. I).Earlier, BRoc K and COOPER (1993) described a simi lar asse mblage, including Eoobo lus aff. elatus (PELMAN, 1986 ), Karath ele napuru (KRUSE, 1990) and vandalotreta djagoran (KRUS E, 1990) from the lateEar ly Cambrian (Toyonian) Wirrealp a and Aroona Cree k limestones, Flinders Ranges, South Au strali a.

The poorl y known type spec ies of Eoobo lus, E. tripa rili s (MATTH EW, 190 2) is re-figured to drawco mpariso n with Eoobolus aff. elatus (PELMAN, 1986). Previously, most Early and Middle Ca mbrianlingulides of this type have been referred usuall y to "Lingulella" or "O be lus" , In co ntrast to these ge nera,Eoobolus has a pitt ed larval she ll, much like that of the acrotretoids , in addi tio n to a distinctive granulatedpos t-larva l she ll. Below we propose the new fami ly Eoo bo lidae to include th is type of linguIides.

Th e evo lutio n of the botsfordiids and ac rothe lids is discu ssed in the ligh t of the new materi al ofKarathele napuru ; it is possibl e to trace a gradua l tra nsition from botsfordi ids, such as Karath ele, toacrothe lides , such as Eothele. Th e mu scle syste m, she ll struc ture , and pseud ointerareas of the bot sfordiidsare very similar to those of Earl y Palaeozoi c Ob olidae (WALCOTT, 191 2; Po POv, 1992 ), and the superfamily Ac ro the loidea, is here referred to the Order Lingul ida.

Acknowledgements. - Frederick J. COLLIER, Jann THOMPSON, Rex DOESCII ER, and the late RichardGRANT (Washing ton, D.e.) ass isted by arranging fac ilities to study mater ial store d in the U.S . NationalMu seum. Janet WADD INGTON (Toronto) arranged the loan s from collec tions in the Royal Ont ario Mu seum.This work has been supported by grants (to L. HOLMER) from the Swedi sh Natural Science ResearchCouncil (NFR). Leonid Po POV gratefully ackn owl edges receipt of a one year NFR visiting scientis t grantand a 10 month visiting sc ientist grant from the Royal Swedish Acade my of Scien ces (KVA) that haveenabled him to wo rk ex tens ively at the Institute of Earth Scien ces, Department of Historic al Geology andPaleontology, Uppsala Uni versity, as well as two Visiting Scientist grants from the Smithsonian Institution .

CAMBRIAN BRACHIOPOD S FROM ANTARCTICA 39

59' W 58' W

E

--:;;:=-~.----j 62' S

O'

King George Island

¥o.o.

W

o,

~~~\>-

f--------_+_ Q -------,-=-"""""----

Fig . ILocation map of King George Island (arrowed) in Antarctica, and the outcrops of glac io-marine strata (asterisked)

of the Ear ly Miocene Cape Melville Form ation .

Lars HOLMER'S fie ld work in So uth Australia was fina nced by NF R, and arrange d by Bru ce RUNNEGAR(Los Angeles) . R. WRONA is deepl y indebted to Stefan BENGTSON (Uppsala) for the arra ng ing of a onemonth visi ting gra nt from the Royal Swedish Acade my of Sciences, for his immeasu rable help during thecourse of this wo rk which was extended to three mo nths in Uppsala, and for the use of laborat ory andSEM fac ili tes at the Depar tment of Palaeontology, Uppsala Un iversity. WRONA'S field work in Antarc t icawas undertaken as par t of the Fifth (1980-8 1) and Tenth (1985-86) Poli sh An tarctic Expeditions organizedand finance d by the Polish Acade my of Sc iences. The investigatio n of Antarc tic material in the Instituteof Paleobiology, Polish Academy of Sc ience s, was supported by Project MR. I. 29. The manuscript hasbenefited from comments on both the language and the sc ientific content by Michael G . BASSETT(Cardiff),Gertruda BIERNAT (Warszawa), Peter D. KRUSE (Darwi n), John S. PEEL (Uppsala), and by Anthony D.WRIGIIT (Be lfast) . All SEM micrograph s were taken at the Electron Microscop y Laboratory of theDep artment of Zoology, Uppsala Uni versity. Thi s work is a co ntribution to the IGCP Proj ect 366.

MATERIAL

The Antarctic brachiopod mater ia l described in this paper co mes fro m the res idue of lim estone erraticboul ders etc hed with 10% acetic acid . The bo ulders are scattered as iceberg-rafted drop stones within theEarly Mioce ne glacio-marine sedi me nts of the Ca pe Me lville Formation, ex posed in the eastern most partof King George Island, So uth Shetland Islands (Tex t-fig. I). The for matio n is represented by a 200 msequence of highl y fossili ferou s sha les with siltstone, marl and sands tone intercalat ions (BIRKENMAJER etat. 1983). The indigenou s fossi l assemblage of the Ca pe Mel ville Fo rma tion co mprises of diatoms,chrysomonad cysts, si licoflagellates, ca lcareous and are naceous fo raminifera , so lita ry corals, polych aetes,bryozoans, gastro pods, bi valves, decapods, echinoi ds, asteroids and fish rema ins (for references see

40 LARS E. HOLMER, LEONID E. POPOV and RYSZARD WRONA

GAZDZICKI ed. 1987 ). The glac ial charac te r of the sediments is ev ide nce d by the presence of numerous,poorly rounded or sharp-edged boulders of rocks exotic for King George Island, with glacially-generatedfeatures (BIRKENMAJER et al. 1983 ; BIRKENM AJER and B UTKI EWI CZ 1988; W RONA 1989 ). Th ese strata alsoyield allochtonous Cretaceous calc areou s nann opl ankton (D UDZIAK 1984 ) and belemnites (B IRKENM AJ ERet al. 1987) dropped into the sedime nt fro m melting icebergs. Th e Early Mi ocene age of the formation ,and co nsequently of the Melvill e Glaciati on , was determined by biostrati graphic data (BIERNAT et al. 1985;GAZDZICKI ed. 1987) as we ll as by rad iometri c dating (cf. BIRKENMAJER 1992 ).

The limestone erra tics were co llec ted fro m the erosiona l surface of the Mel vill e Penin sul a and arehou sed at the In stitute of Paleobiology, Poli sh Ac ademy of Sc iences in Warszawa (abbrev iated as AElMe).This coll ecti on co ntains severa l hundred boulders grouped int o distinct lith ological typ es (W RONA 1989 ;WRONA and ZHURAVLEV 1996 thi s vo lume).

Very characte ris tic boulders of the light co loure d calcimic robial-a rchaeocyatha n frameston e (type 11 ;WRONA 1989) co ntain rare she lls of Eoobolus aff. elatus and an abunda nt blue-green algae Gordonophyton-Renalcis and arc haeocyathan assemblage of the sa me age as the uppermost Botom an units withSyringocnema favu s Bed s distingui shed in South Au strali a (ZHURAVLEV and GRAVESTOCK 1994; WRONAand ZHURAVLEV 1996 thi s volume) ; they ca n be also co rre lated with faci ally si milar beds of the Sh ackletonLimeston e ex posed in the Transant arcti c Mountains (DEBRANNE and KRUSE 1986, 1987 ; REES et al. 1989;ZHURAVL EV and GRAVESTOCK 1994).

Lin gul ate bra chi op ods identified as Eoobolus aff. elatus are abundant and well preserved in black ,some times slightly bituminous pack stone and wackeston e (types 12-14; W RONA 1989). These kind s oflim estone erra tics also co ntain arc haeocya ths, spo nge spic ules , gas tro pods and hyoliths as we ll as a di verseassemblage of problem atica (sma ll shelly fossil s), amo ng them numerous sclerites of the tommot iidDailyatia, which are being describ ed separately (BENGTSONand WRONA, in prep aration). Th is sma ll she llyfossi l assemblage ca n be co mpared with a simi lar fauna and sediments of the Parara Limestone (Bo toman)in South Australia (BENGTSON et al. 1990 ; ZHURAVLEV and GRAV ESTOCK 1994), but on the othe r hand itcan also be corre lated with an autoch tono us Antarc tic ken nardi id asse mblage recorded fro m the Shac kleto nLimeston e in the Transant arctic Mo untai ns (EVANS and ROWELL 1990; EVANS 1992 ).

Extre me ly ric h and we ll preserved shells of larval as we ll as ad ult Karathele napuru, Vanda lotretadjagoran and Eoobolus aff. elatus orginate exc lusive ly from one erratic boulder: AE/Me52. This com pletely etched sma ll boul der, size: 4 cm x 5 cm and I cm thic k, co mpr ise d a brachi op od coquino idlimestone; it was the most fossi liferous samp le stud ied and also co nta ined chancello ri id and spo ngespic ules, gas tro pods, hyoliths and phosphatize d trilobite carapaces. Th e age of th is erratic ca n be es ta blished as late Ea rly Cambrian (Toyonian) by the brachiopo d asse mb lage described below, which is closelysimilar to the brachi op od fauna fro m the late Ea rly Ca mbrian Wirrealpa and Aroona Creek lim eston es,Flinders Ranges, South Aus tra lia. Equiva lent stra ta co ntai ning these brach iopod co quino id lim estones hasnot ye t been discovered on the Antarctic landmass. However, the general lith o logical co mpos ition of thewhole spec trum of erratics fro m the Ca pe Me lvi lle Forma tion clea rly suggests that outcrops of Low erCa mbrian roc ks aro und the Wedde ll Sea (in particul ar the Arge ntina Range) and north ern Transant arcticMountain s acted as so urce areas for the glac ia l boulders (WRONA 1987; BIRKENMAJER and BUTKIEWICZ1988; W RONA 1989).

New materi al of Eoobo lus aff. elatus and Karathele napuru fro m the Wirrealpa Limeston e, FlindersRanges (Ten Mile or Mount Bill y Creek sec tion within the Wilkawillina Go rge; see KRUSE (19 91 a) forlocalit y da ta) is illustrated below for co mparison with the Antarc tic material. Karathele napu ru andVandalotreta djagoran were descr ibed origina lly by KRUSE ( 1990) fro m the ea rly Middle Cambrian TindallLimeston e (Ordian) of the Dal y Basin, Northern Territory, Aust ra lia.

SYSTEMATIC PALEONTOLOGY

Measurements (Tables 1-6; in millimetres if not stated othe rw ise) are as foll ows: W, L, T = width,length, height of va lve ; Iw, 11 = width, len gth of dorsal pseudointer area; Pw = width of medi an groove ;Cw, Cl =width, len gth of ca rdina l muscl e field ; VI =length of visceral area; SI = len gth of dorsal medianrid ge. S =standa rd deviation, N = number of measurements, MA X = maximum value, MIN = minimumvalue .

CAMBRIAN BRACHIOPODS FROM ANTARCTICA 41

Th e terminology of the lingul ate brachi opods used here mainly fo llo ws that of Ro wELL (1965); KONEVA(1986); and HOLM ER (1989). Th e term median tong ue was introduce d by PoPOV and HOLMER (1994) toden ote the narrow ant eri or ex tension of the dorsal viscera l field of man y lingul ates (Text- figs 2-3).

The illu str ated and/o r discussed material is dep osited in the Insti tut e of Paleobi ology, Pol ish Academ yof Sciences, War szawa (Z PAL), So uth Australia n Mu seum, Adelaid e (SA M), Northern Territory Mu seumof Arts and Sciences, Dar win (P) . Un ited States Nation al Mu seum, Was hington D.e. (USNM), Swedi shMu seum of Natural History (RM), Stockho lm, and the Royal Ontar io Mu seum, Toronto (ROM) .

Class Lingulata GROJANSKY et Porov, 1985Order Lingulida WAAGEN, 1885

Superfamily Linguloidea M ENKE, 1828Fa mi ly Eoobolidae fam . novo

Diagnosis. - Sh ell dors ibicon vex, so mewha t inequiva lve d, e longate oval to subtr iangular; larval she llwe ll defined, with pitted micro-ornam en tation; postlarval she ll fine ly pustulose ; ve ntra l pseudointerareaelevated above va lve floo r, with deep pedi cl e groove and we ll-developed flex ure lin es; dorsal pseudointerarea d ivide d, raised above va lve floor ; mu scle sys tem wit h paired umbon al mu scl e scars bisected byv-sha ped impression of pedicle nerve ; dorsal visceral fie ld wi th we ll-developed medi an ton gue ex tend ingto mid-len gth ; mantle canals bac ulate with we ll-develo ped vas cu la medi a.

Gen era included. - Eoobolus MATTH EW, 1902 (= Clivosilingu la USH ATINSKAYA, 1993); VassilkoviaPoPOV et KHAZANOV ITCH (in Po POV et al. 1989).

Discussion. - The Eoobolidae show a co mbination of charac ters , that are un iqu e wi thin the Lin guloid ea. There are no other lingul oids with a fine ly pi tted larval she ll and pustulose pos t-larval she ll. Othertypes of pitted micro- ornamentat ion are distributed wide ly with in the superfamily, but usu all y invo lveboth the larval and post- larval she ll as in the Zhanate llidae, Elka niidae, Dysorist idae, and the Paterul idae.It is possible that the eoobo lids are re lated so mehow to the Botsfordiidae, the ea rliest members of whic hare also characterised by pi tted larval and pustulose post-l arval she lls . Th e detailed morphology of thelarval she ll of vassi lkovia PoPOV et KHAZANOV ITCH (in Porov et al . 1989), is no t known, bu t the pustulosepost-larval she ll is closely simi lar to th at of the Eoobolidae. The fa mily is firs t recorded from the Botoman(USIIATINSKAYA 1993) and ranges to the Late Cambrian (Porov et al. 1989).

Ge nus Eoobolus MATTH EW, 1902

1902. Obolus (Eoo bolus) MAlTIIEW. p. 97.

1903 . Obolus (Eoobolus) MAlTHEW. p. 135.

1993. Clivosilingula USH ATINSKAYA. p. 125 [Type species: Lingulella cli vosa PELM AN. 1983 ; Midd le Cambrian (Amg ian).Siberi a].

Type specie s: Obolus triparilis MATrIlEW. 1902 (se lected by ROWELL 1965. p. H263).

Diagnosis. - She ll do rsibiconvex, e longate suboval to elongate subtriang ular; ventral va lve slight lyacuminate with high , ort hoc line pseudo interarea ; ped icl e groove deep, narro w, wi th subparalle l lat eralmargins ; do rsa l pseudointerarea wit h broad, shallow med ian groove; propareas of both va lves with flexurelines; ventra l viscera l fie ld not extend ing to mid-valve ; paired ve ntra l um bon al muscle scars bisect ed byv-s haped imp ression of pedic le nerve ; ventra l vascula lateralia sub marg ina l, straight, slightly d ivergentproximall y; dorsal visceral area with median tongu e usu all y bisected by med ian ridge and pair of subme dian ridges ; dorsal vascula lateral ia marginal, arcuate; vascula media long, s lightly divergent.

Other species included. - Ling ule lla wanniecki REDLlCH, 1899; Ea rly Ca mbrian, Pak istan (Sa ltRan ge). Ling ulella clivosa PELMAN, 1983; Middle Cambrian (A mgian), Siberia. Ling ulella elata PELMAN,1986 (= Clivosiling ula dilata ta USHATINSKAYA, 1993); Middle Cambria n (Amgian), Siberi a. Eoobolus aff.elatus (PELMAN, 1986); Early Cambrian (Toyonian), So uth Austra lia, West Antarcti ca (King GeorgeIsland ).

Discussion. - Eoobo lus has been regard ed as a junior sy no nym of both Lingul ella (WALCOTT, 1912),Obolus, and Ung ula (ROWELL, 1965; the figure d va lves prob abl y represent Ungula) . However, it see msthat und oubted species of Lingulella and Ungula are not kn own earlier th an fro m the Late Cambrian ; onlyLingulella is int ern all y similar to Eoo bolus, but the former has poorly developed dorsal flexure lines andpse udo interareas that are not stro ng ly ra ised above the va lve floor (Porov and HOLMER 1994). Obolus


-----w- ---- _ .,

- --- -- - - ------ --- - --~-------~

A

---- Iw---~

anteriormuscle field

umbonal

: r-posterolateralmuscle field

Iimpression ofgastroparietal?ands I

L

median tongue anterior lateral BFig. 2

Eoobolus aff, elatus (PELMAN, 1986), reconstru ction of vent ral (A) and dorsal interi or (B) . Location of measurementsindicated : W = width, Iw = width of pseud ointerarea ; L = length, VI = length of ventral imp ression of pedicle nerve , 11 =

length of ven tra l pseudointera rea.

appears to be endemic to Balto scandia and is not similar internall y to Eoobolus (Porov et al. , 1989).Moreover, our brief re-study of the type species Eoobolus triparilis (M ATTH EW, 1902) revealed a distinctive pustu lose ornamentation on the post-larval she ll, of a type which is not known from any other obolidgenera with the exception of Vassl lkovia Porov et KHAZANOVITCH (in Por ov et al., 1989 ) and Clivosilingu la USHATINSKAYA, 1993. E. triparilis also has a characteri st ic lon g median ridge and a pair of lateralridges in the dorsal valve . A similar type of morphology was recorded in the typ e species of Clivosilingula,C. clivosa by PELM AN ( 1983 , p. 125) and the pseudointerareas as well as most other morphologicalcharacters are closely similar; Clivosi lingula is here regarded as a junior sy no nym of Eoob olu s.

According to USIIATINSKAYA ( 1993) Lingulella viridis COBBOLD, 1nl , Lingulella rotunda PELMAN,1977 , and Lingul ella variabilis PELMAN, 1977 , may be also referr ed to Clivo si lingula , but the det ailedmorphology of the se taxa is poorly known.

Lingulella wanniecki REDLICH, 1899 , fro m the Early Cambrian of Paki stan has a we ll defined medianridge and a pair of later al ridge s in the dorsal val ve as well as a di stinctive pu stulose ornamentation(SCHINDEWO LF and SEILACHER 1955); it is referred here to Eoo bolus.

Eoobo lus triparilis (MATTHEW, 1902 )(PI. 10: 1-9)

1902. Obo lus trip arilis MATrl IEW, p. 94, pI. I : 2a-c .

1912. Lingul ella tripa ril is (MATIHEW . 190 2): WALCOTI . p. 536 . pI. 45: 2. 2a-L text- fig. 44 ( CUIll s)"n.).

Lec totype (se lected here): ROM 5 10CM (A), dorsa l valve (PI. 2: 7).

Type horizon and locali ty: Middl e Cambrian (Bo urinot Group), Ca nada (Ca pe Breton ).

Diagnosis. - Sh ell dorsibiconvex , slightly elon gate subtriangular, with maximum width somewhatanterior to mid-length ; ventral valve slightly acumina te ; ventral pseudointerarea high , tri angular; dorsalinterior with lon g median ridge and pair of submedian rid ges bounding narrow median ton gue .

Rema r ks. - The micro-ornam entation of the larval shell could not be determined on any of theavai lable specimens. The fragmentary pre servation of the material at hand made it difficult obtain goodmeasurements .

Materia l. - Figured. Ventral valves : USNM 51 855 a, 570l3c. Dor sa l valves: USNM 5701 3a, 57013d,57013e; ROM 5lOCM (A) , 5 lOB, 678CB (C), 678CM (E) .

Occur rence. - Type locality only.


Eoobolus aff. elatus (PELMAN, 1986)(PI. 9: 1-15; Text-fig. 2)

aff. 1986. Lingulella elata PELM AN, p. 138, pI. 12: 1-4.aff. 1993. Clivosilingula elata (PELMAN); U SHATINSKAYA, p. 133, fig. 1: 10-11.

1993. Lingulella sp.; B ROCK and C OOPER, p. 780, fig. 14: 14-15.

Material. - Figured from Antarctica. Ventral valves: ZPAL Bp.XXXIII/33S6, 33S9, 61S6. Dorsalvalves: ZPAL Bp.xXXIII/50S1 , 61S2. Figured from Au stralia. Complete juvenile shell: SAM P35112.Dorsal valves: SAM P3510 1, P35104, P35105 , P3511 1. Ventral valves SAM P35102, P35103, P35109,P35110, P35114.

Measurements: See Tables 1- 2.

Table I. Eoobolus aff. elatus (PELMA N) , average dimensions of ventral valves.

L W 11 Iw VI LIW IVL IwlW VVL

N 27 27 26 26 17 27 26 26 17

X 2.39 1.79 0.62 1.17 1.29 135% 27% 68% 50%

S 1.117 0.882 0.264 0.513 0.545 7.46 3.1 5.9 6.1

MIN 1.04 0.60 0.28 0.52 0.64 120% 20% 58% 40%

MAX 5.28 4.28 1.40 2.56 2.36 150% 32% 79% 63%

Table 2. Eoobolus aff. elatus (PEI.MAN), average dimen sions of dorsal valves.

L W Iw LIW IVL IwlW VIIL

N 20 20 17 27 26 26 17

X 2.27 1.81 1.12 135% 27% 68% 50%

S 0.860 0.700 0.464 7.46 3.1 5.9 6.1

MIN 1.06 0.84 0.52 120% 20% 58% 40%

MAX 4.32 3.48 2.40 150% 32% 79% 63%

Description. - Shell slightly dorsibiconvex , elongate suboval, on average 135% as long as wide, withmaximum width somewhat anterior to mid-length . Ventral va lve gently and evenly convex, slightlyacuminate; ventral pseudointerarea high triangular, orthocline, on average 27% as long as wide , occupying68% of valve width; ventra l propareas slightly rai sed above valve floor, with well defined flexure line s;pedicle groove narrow, with steep, subparalle l lateral margin s. Dorsal valve gently convex; dorsal pseudointerarea moderately high , orthocline, occupying on average 66 % of valve width; medi an groove broad,sha llow, poorly defined laterally ; dorsal propareas narrow with flexure line s. Larval shell small, close tocircular, around 0.10-0.15 mm across , ornamented by fine circular pits, around I m acro ss. Post-larvalshell co vered by fine , densely spaced pustules, around 5-6 m acro ss.

Ventral viscera l area somewhat thickened, extending to mid-length; ventra l umb onal muscle scarpaired , bisected by v-shaped impression of pedicle nerv e; vent ral vascula lateralia subperipheral, slightlyarcuate. Dorsal visceral area, with narrow median ton gue ex tending somew hat anterior to mid -val ve;dorsal median ridge weakl y developed; dorsal vasc ula lateralia margin al, arcuate; vasc ula media weaklyimpressed.

Discussion. - The specime ns fro m Antarctica are similar to Eoobolus elatus (PELMAN) from theKuonamka Formation of Siberi a (USHATINSKAYA 1993, figs 1-11 ) in size, general outline, and in havin ga weakl y de veloped dor sal medi an ridge, as well as in the absence of dor sal submedian ridges; the onl ydifferen ce appears to be that the ventra l viscera l area is more thickened in the Siberian specimens .However, the detailed morphology of the Siberian spec ies is still poorl y known and it cannot be comparedin detail with the Antarctic material. A second species, Eoobolus dilitatus described by USHATINSKAYA(1993, fig. 1: 5-9) from the same horizon, differs from Eoobolus etatus in bein g transversely oval andwider, but the se features are known to be very variable in lingul ids and moreover, change significantlydur ing ontogeny; thus, E. dil itatus may be a junior synonym of E. etatus .

The Antarct ic material of E. aff. elatus is also clo sely similar to Lingulella sp. described by BROCKand COOPER (1993) from the Early Cambrian Wirrealpa and Ramsay limestones of the Flinders Rangesand Yorke Peninsula, South Australia . Additional material from the Wirrealpa Limestone illu strated here

44 LARS E. HOLM ER. LEONID E. POPOV and RYSZARD WRONA

(PI. 9: 3-4, 6-8, 10, 13, 15) indi cates that they may be con specifi c, as the Australian specimens have anidentical type of external and internal morphology.

Occurrence. - Early Cambrian (Toyonian), South Australia ; King George Island erratics , Ant arctica;Middle Cambrian (Amgian), Siberia.

Superfamil y Acrotheloidea WALCOlT et SCHUCHERT, 1908

Diagnosis. - Shell with short, con vex posterior margin ; postlarval orn ament of fine, evenly distributedgranules or pustul es; larval shell pitt ed , with apica l spines or tubercles; ventral valve convex or lowconical; ventral pseud ointerarea ves tigial or lacking; pedicl e emerging through delthyrium or for amenposterior to apex ; dorsal pseudointerarea vestigial or lacking; muscle sys tem linguloidean-like, with pairedumb onal muscle; mantl e canal sys tem of both valves baculate; vascula media well-deve loped; shellstruc ture bacul ate.

Discussion. - The close relationship between the Botsfordiidae and Acrothelidae has long beenrecognized (ROWELL 1965). As noted by ROWELL (196 5) it is probable that the botsfordiides were ancestralto the acrothelides; moreover, it now see ms possibl e to trace an almost continuous gradual transiti on inmorphology from Botsfo rdia (PI. 10: 10-15 ; Text-fig. 3A-B) through the intermed iate Karathele, with adeep, but unrestricted delth yrial pedicle opening and rud imentary ventral pseudointerarea (Text-fig. 3C- D)to ear ly acrothelides, like Eothele (PI. 12: 6-8) with a low, subconical ventra l valve, a reduced pseudoin terarea that is gradually transformed into the pedi cle tube, and with an elonga te, suboval ped icle fora me nthat is formed post-l arvally (PI. 12: 6- 8).

The muscle sys tem of botsfordiides is very similar to that of Early Palaeozoic Obolidae (WALCOTT1912; POPOV 1992); the earliest known genera, Botsfordia and Edreja have sca rs that appea r to match afull set of obolide muscles (Text-fig. 3A-B), whils t the numb er of muscles in most acrothelides wasprobabl y reduc ed . The scars of a linguloid type of v-shaped pedi cle nerve in Botsfo rdia and Edreja(Text-fig. 3A) also may indicate a close relat ionship between the two gro ups.

The shell structure of botsfordiides has not yet been studied adequate ly, but includes bacul ate laminae(HOLMER 1989; POPOV and HOLl'vt ER 1994).

The earl iest evo lutio n of the group is st ill poorly known . Although the botsfordi ides and acrothelideshave some characte rs in common with the acro tretoids, such as a pitted larval she ll and a pedi cle foramen(in acroth elid s), it is clear that they also exhibit many lingul id fea tures , such as bacul ate she ll str ucture .They might be related close ly to the lingulid family Eoo bolidae, which is also charac terized by a pittedlarval shell and a pustul ose-granul ar post-larval orn amentation . WILLlAMS and ROWELL (1965) prop osedthat the acrothelides probably were ances tra l to the discinides, but in view of the conside rable differencesin their ontoge nies, this is unlikely (CHUANG 1971 ; HOLMER 1989).

Family Botsfordiidae SCHINDEWOLF, 1955

Diagnosis. - Shell biconvex; ventra l pseudointerarea ves tigia l, divided by deep pedicle groove forming triangul ar delth yrium; dorsal pseud oint erarea ves tig ial, divided by median groove; muscle sys temconsisting of paired umbonal, transmedian, outs ide lateral, internal lateral, anter ior lateral, and centralmuscles; vasc ula lateral ia straight, submedian, divergent in both valves ; larval she ll with one to threeapica l tubercl es in ventra l valve and two in dorsal valve .

Genu s Karathele KONEvA, 1986

Type spec ies: Karath ele coronata KONEVA. 1986 .

Type horizo n and localit y: Midd le Cambri an (Peronopsis? ultimus and Ptychagnostus atavu s Zo nes ). Kazakh stan (MalyiKaratau ).

Diagnosis. - Shell subequibico nvex, subcircular ; ventra l pseudointerarea narrow triangul ar, proclineto catacline, with vest igial prop areas; pedicle groove deep, with stee p lateral slopes; dorsal valve slightlysulcate ; dor sal pseudoin terarea ves tigial, with poorl y defined median groove; ventral larval shell withthree tubercles and one or two pairs of dorsal tubercl es; ventra l visceral field small, short, onl y slightlythickened ; dorsal visce ral field small, slightly thickened ; median ton gue narrow, extending to mid-valve,bisected by median ridge ; mantle ca nal sys tem bacul ate ; both valves with straig ht, submedian, divergentvascula lateralia ; vasc ula media long, divergent.


pediclegroove proparea umbonal

transmedianimpression ofgastropa rietalbands

vascula mediamediantongue

impression ofpedicle nerve A B

mediantongue

Dvascula mediac

umbonal

transmediananteriorlateraloutsidelateral

middle ----''------lateral

-...r--- - ....L central

vasculateterelie -t-- - .

proparea

pediclegroove

Fig . 3Reconstru ct ion of vent ral (A) and dorsal interior (B) of a ge neralised Botsfordiidae and ventral (C)

and dorsal interior (D) of Karathele napuru (KRUSE, 1990 ).

Other species included . - Eotlzele napuru KR USE, 1990; Early Cambrian (Toyonian)-early Middl eCambrian, Australia, Antarctic a.

Discussion . - Karath ele was referred originally to the Acrothelidae by K ONEVA (1986 ); howe ver, ithas an open delthyrium throughout onto geny, with a ventr al pseudointerarea and ped icle groove muchlike those of Botsfordia and Edreja (Text-fig. 3). Karathele differs from both Botsfordia (PI. 10: 10-15)and Edreja mainly in having a procline to catacline ventral pseudointerarea with vestigial propareas, small ,densely spaced post-larval pustule s, one pair of ventral larval tubercle s, as well as a short ventral visceralarea that does not extend anteriorly to mid-v alve. Externally Karathele is closely similar to Eoth ele (PI. 12:6-8), but the delth yrium in Karathele remains open through out ontogeny. The juvenile shells of Eoth elespurri are similar to Karath ele in the morphology of the larval shell , delthyrium and rudimentary pseudointerareas, but unlike Karathele, a regular pedicle foramen is formed in adults of Eothele spurri(PI. 12: 7) and the ventral propareas are completely lost. The interior of Eotlzele is poorly known andcannot be compared directly.

Karath ele napuru (KR USE, 1990)(PI. 11: 1-8; PI. 12: 1-5 ; Text-fig. 3C-D)

1990. Eothele napuru KRUSE, p. 3 1, pI. 12A-K, text -fig. 16.

199 Ib. Eoth ele napuru KRUSE, p. 178, fig. 7A-E.

1993. Eoth ele napuru KRUSE; BRoc K and COOPER, p. 782, fig . IS : 1-14.

Holotype : Ventral valve SAM P85206.

Type horizon and locality : Middle Cambrian , Tind all Limestone (bore -hol e NTGS 83/3), Daly Basin , Northern Territory,Australia.

46 LARS E. HO LMER. LEON ID E. I'OP O V and RYSZAR D WRONA

Material. - Figured fro m Antarctica. co mplete she lls: ZPAL Bp.XXXIII/29S8, 38S4 ; ventr al va lves :ZPAL Bp.XXXI1I/53S2, 5340U, 54S 3; do rsa l va lves: ZPAL Bp.XXXIII/29 S5 , 34S3 . 50S2, 50S 3, 53S4.Figured from Australi a. Ventr al valves : SAM P35100. SAM P35 107, SAM P35 113.

Measu remen ts: See Tabl es 3--4 .

Tab le 3 . Karathele Ila p llnt (KRUSE). av erage d imens ions of 5 vent ral val ves.

11 11 I. I W I Iw I I' w 1 VI Vw 1 LlW Iw/W 1 VI/I. I11 X 11 1.12 1 1.18 .1 0.33 1 0.16 1 0.38 0.43 I 979c 279c 1 349c 1

S 0.242 I 0.331 0.130 I 0.038 0.060 0.103 8.6 5.9I

3.8 II I

i\ IIN 0.82 I 0.76 0. 16I

0. 10 I 0.32 0.32 84'le 2 1'le I 299c !!I

11 MAX I I Ii -

II 1.40 1.46 1.34 0.20 I OAO I 0.50 I I089c I 349c I 39%11

Table 4 . Karath ele napuru (KRlISE). av erage d imens ions o f 6 dorsa l va lves.

I I. W Iw SI I LlW Iw/W SilL

X I A I 1.59 0.381 '~ _~

89'7e 24% 60%

S 0.334 00409 0.083 0.28 2.5 3.5 604

MI N 1.02 1.1 6 0.28 0.58 85% 20% 52%

MAX 2. 14 2.14 1.22 1.22 92% 29% 69%

Diagnosis. - Pedi cle groove deep, forming narrow slit in mature specimens; dorsal larval she ll withtwo tubercles.

Descript ion of materia l fro m Anta rct ica : She ll s lightly ventribiconvex, subcircular. about 97 % as longas wide. Ventral valve low subconica l with maximum hei ght at umbo; ven tra l pseudoi nterarea narrowtria ngular. proc line to slightly catac line, occupying on avera ge 27 % of valve width: pedicle groove verydeep . forming high tria ngular to scmioval delthyr ial opening: posterior part of delthyrium forming narrowslit in mature specime ns ; ventra l propareas vesti gial. Dorsal valve ge ntly con vex with maximum heig htat about umbo: dor sal pse udointer area vesti gial. occupy ing abo ut 24% of valve width. Ventral larval she llwith high medi an tubercle above delt hyri um and pair of tubercl es in postero lateral part. Dorsal larval shellwith two high tube rc les posterolatera lly.

Ventral visce ral area small. occupying about 34% of valve length and 38 % of valve width: vasculalateralia of bot h va lves submedian. straight. di vergen t. Dorsal interior with slightly raised viscera l areaocc upying about 60 % of valve lengt h: dorsal median tongue bisected by long ridge.

Discu ssion. - Karathele napuru differs from the typ e species , K. co ronata (KONEV,\ , 1986 ). in havin gon ly two tubercles o n the dorsal larval she ll, and a much deeper pedi cle groove .

There are virtually no di fferences between our material and the material described by KRUSE ( 1990 ,199 Ib). However, the maxi mum recorded size from the Antarctic (max L = 2. 14) is much smaller thanthat recorded by KRUSE (1990: max L = 7 .6) , and the delthyri um remains ope n thro ughout ontogeny inthe Antarctic spec imens, whilst the larger va lves described by KRlJSE ( 1990) have a fine slit or suture. Asimilar type of delthyrium is also found in the larger specimen s from the Wirrea lpa Lim estone, Fli ndersRanges (PI. 12: 4-5 ), but in all know n specime ns of K. napuru the delth yria l margin s rem ain separated(eve n thou gh they mig ht "to uch" one another) and do not join completely as in mature she lls of Eothele(PI. 12: 6-8).

Occurrence. - Ear ly Cambrian (Toyonianj-early Middle Ca mbrian. Au stral ia (Nothern Territory).King George Island erratics, Antarctica.

Order Acrotretida K UHN, 1949Superfami ly Acro treto idca SCIIUCII ERT. 1893

Fami ly Acrot retidae SCH UCII ERT. 1893Genus vandatotreta MERGL. 1988

1988. vandalot reta M ER(;L, p. 292 .

1990. Luh otreta MER(;!. and S!.EIIOI'ERo vA. p. 95 (Type spec ies : L. pompeck ji M ER(;L et SU :1I0I'ERo vA. 1990).

Ty pe species : vanda lotreta vafra MEJ{(;L. 1988.Typ e hori zon and localit y: Mi ddl e C ambrian (A mg ia n). High Atlas (Yag o ur inli e r). M orocco.


Diagnosis. - Sh ell tran sversel y ova l to subcircular with narrow, convex posterior margin; ventralval ve wide conical ; ventra l pseudointerarea procline to ca tacl ine, poorly defined laterall y with intertrough :foram en not enc losed within larval she ll; dorsal valve weakly convex; dorsal pseudointer ar ea shor t withbroad ly triang ular medi an groove; apica l process for mi ng bo ss-like thi ck ening ante rior to internal for ame n; dorsal visceral fie ld with narro w median ton gue , usu all y bisected by ves tig ial med ian ridge ; dorsalmedi an buttress develop ed .

Other species included. - Acrotre ta pompeckji M ERGL et SLEIIOFERovA, 1990; Middle Cambrian ,Jin ce Formation , Ce ntra l Bohemia (Czech Republic). Hadrot reta dja goran KRUSE, 1990; Early Cambri an(Toyonian)-early M iddle Ca mbr ian. A ustralia, West A ntarctica. Acrotreta lim oensis WIMAN, 1903; earl yMiddle Ca mbr iant") , Sou th Bothnian Sea (Limo n) , Sw ed en.

Discussion. - Luh ot reta M ERGL et SLI~HOFERovA , 1990, from the ea rly Middle Ca mbrian of Bohemiadi ffer s only s lig htly from vandalotreta in detail s o f orna me ntatio n and the complete abse nce of a dorsalmedi an ridge ; the ge nus is regard ed here as ajunior sy nony m. Vandulot reta is mo st similar to HadrotretaROWELL, 1966, but differ s mainly in havin g a boss-shaped apical proc ess that do es not fill the entire apexas in Hadrotreta ; moreo ver, the dorsal medi an ridge is vestigial to co mplete ly ab sent in vandalotreta andthe dorsal va lve lack s a medi an sulcus. vandalotreta is also so me wha t s imilar to the ceratre tide Bozshakolia HOLMER et USHATINSKAYA, 1994 , in the poor development of a dor sal median ridge and having afo ra me n that is not enc losed with in the larval she ll, but di ffer s in the lack of a rid ge-like apical processenc los ing the ped icle tube. Th e poorl y known Acrotreta lim oen sis WIMAN, 1903, from early MiddleCa mbrian(?) g lac ia l errat ics on the Island of Lim on in the South Bothn ian Sea can also be referred tovandalotreta (HOLMER and MERGL in prep arati on ).

vanda la treta djagoran (KRUSE, 1990)(PI. 13 : 1-9; Text-fi g. 4)

1990 . Had ro treta djagoran KRUSE. p. 29 . fig . 5. pl. II A-N .

1991 b. Had ro treta djagoran KRUSE. p. In. jig. 6H-L.

1993. Hadrotreta primaevu (W ALCOn): BROCK and COOPER. p. n2. fig . 14 : 1-1 3.

Holotype: Ventral va lve SAM 1'851 38.

Type hor izon and local ity: Midd le Cambrian, Tinda ll Limestone (bo re-ho le NTGS 83/3) , Daly Bas in, Northern Territ ory.Austra lia .

Material. - Figured from Antarctica ; complet e she lls: ZPAL Bp .XXXIII/29S I, 39S8, 54 S6 , 54S 7;ventra l va lves: ZPAL Bp .XXXIII/54S5, 6 1S80, 5337U ; dor sal va lves : ZPAL Bp .XXXIII/5 333U.

Measuremen ts: See Tab les 5-6.

Tab le 5. vando lotreta dja goran (KRlISE). average dimensions of ventra l valves.

L W T LIW T/L

N 8 8 8 8 8

X 0.55 0.65 0 .26 85 % 48 %-

S 0.148 0.239 0 .118 1.82 1.61

MI N 0.30 0 .34 0 . 10 83 % 33 %

MAX 0.80 0 .94 0.46 88% 80 %

Tab le 6. vanda iotreta djagoran (KR USE). average dimensions of dorsa l va lves .

L W 11 Iw PI Cl Cw SI LIW IwlW CI/L C wlW SUL

N 5 5 4 4 4 4 4 4 5 4 4 4 4

X 0.85 0.99 0 .055 0.54 0 .21 0 .23 0 .64 0 .66 86 % 45% 26% 56 % 60 %

S 0.439 0.503 0 .025 0.230 0 .099 0 .079 0 .189 0 .388 4.5 6.9 3. 1 6. 1 6.4

MIN 0.30 0 .36 0.02 0 .24 0 .08 0 . 14 20% 0.12 8 1% 39 % 23% 49 % 52%

M AX 1.30 1.56 0 .08 0 .76 0.32 0 .32 32 % 0.98 90% 65 % 30% 63 % 69 %

Diagnosis. - See KRUSE ( 1990, p. 29).


cmedian ridge

___________________~_o__,__~_ _ -+-_-

.:~ Iw--:

11 : ' ':--~- t ::::j:--: :~: -- - ~ -----j------- -- T--: i~lrQJ): 91

: -- SI

L

cardinalmuscle

field

vascutatateralia

umbonal(apical pits)

apical process B(muscle platform)

A

Fig . 4Vandalotreta djagoran (KRUSE, 1990), lateral view of the ventral valve exterior (A), reconstruction of ventral (B) and dorsalinterior (C) . Location of measurements indicated : W - width of valve, Cw - width of cardinal muscle field , Iw - width

of dorsal pseudointerarea; L - length of valve, Il - length of dorsal pseudointerarea; T - height of valve.

Description of material from Antarctica: Shell ventribiconvex, transversely suboval, on average 85%as long as wide, and 43% as high as long . Ventral valve moderately convex with maximum height nearumbo ; pedicle foramen subcircular, situated immediately posterior to apex , not enclosed within larvalshell; ventral pseudointerarea moderately high, procline to slightly catacline, divided by broad intertrough.Dorsal valve gently convex; dorsal pseudointerarea low, with broad, shallow median groove, and narrowpropareas.

Ventral interior with apical process forming low boss-shaped projection with median depressionanterior to foramen; internal pedicle tube short; apical pits closely spaced on either side of apical process.Dorsal median ridge thin, low, occupying about 60% of valve length, and originating directly anterior tolow, triangular median buttress; dorsal cardinal muscle fields relatively small, somewhat thickened,extending anteriorly for about 26% of valve length; dorsal central muscle scars weakly impressed.

Discussion. - There are no significant differences between the specimens from Antarctica and Australia. Vandalotreta djagoran is closely similar to the type species, V. vafra, differing only in having amore transversely suboval outline, and a more well defined dorsal median ridge, as well as less welldefined dorsal central muscle scars ; it may prove to be a junior synonym of the type species.

Occurrence. - Early Cambrian (Toyonian)-early Middle Cambrian, Australia (Northern Territory),King George Island erratics, Antarctica.

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Porov, L.E., KHAZANOVITCH, K.K ., BOROVKO, N.G ., S ERGEEVA, S.P., and SOBOLEVSKAYA, R.F . (Flortou, Jl .E ., Xa3aHOnl1lJ,K.K. , Boposxo, H.L, Cep reeaa, e. n ., C060JlCIICKa5l, P.<D.) 1989. Onopuue paspesu 11 crparnrparpna KCM6po0PtlOBl1KCKOH cjJOCcjJOPIITOIIOCIIOH o60JlOnOH TOJlIl\I1 Ha ceeepo-sanane PYCCKOH nnarrpopnu. - AH CCCP, MUHU cmepcmeo r eOJlOCUU CCCP, Me xeeooucmeenuua cmpamuepatpuuecsuu «ouumem CCCP, TpYObt 18, 1- 222.Hayxa , Jleanurpan.

Porov, L.E. and SOLOVJEV, LA . (Il onos , Jl .E ., COJlOBhCB, l1.A.) 1981. CPCtlHCKCM6pl1iicKIIC 6C33aMKOllhlC 6paxllonOtlbl,Xa Hl\e JlJIOplIlI 11 Tpl1J106l1Tbl 3allatlllOH Anrapxrn nu, XPC6Thl W eKJlTOII 11 Apnaceuruna. - Aumapxmuxa 20 , 64- 72.

REES, M .N ., PRATT, B .R., and Row ELL, A .J . 1989. Early Cambri an reefs complexes, and associated lithofacies of theShackleton Limestone, Transantarctic Mountains. - Sedimentology 36,341 -361.

REDLlCII, K.A . 1899. The Cambrian fau na of the eastern Salt-Range. - Memoirs of the Geological Sur vey of India .Palaeonto logia lndica 1, 1-13 .

ROBERTS, J ., JELl.. P.A 1990 . Ea rly Middle Cambrian (Ordian) brachiopods of the Coonigan Formation , Western New So uthWales. - Alcheringa 14 , 257- 309.

RowELL, A.J . 1965. Inarticulata. In: R.e. Moore (ed .) Treatis e on Invertebrate Palaeontology, Part H. Brachiopoda 1(2) ,H260- H296. Ge o logi ca l Society of America and University of Kansas Pre ss, Boulder, Co lorado and Lawrence , Kansas .

ROWELL, A.J . 1966. Revision of some Cambrian and Ordovi cian inarticulate brachiopods. - The University of KansasPaleontologica l Contributions 7, 1- 35.

ROWELL, AJ. 1980. Inart icu lat e brachiopods of the Lower and Middle Cambrian Pioche Shale of the Pioche Dis trict, Nevada.- University of Kansa s Paleontological Contributions, Paper~, 1-34 .

ROWELL, A.J ., COOPER, R.A ., J AGO, 1.B ., and BRADDOCK, F. 1984. Paleontology of the Lower Pal eozo ic of Northern VictoriaLand: brachiopods with Au stralian and New Zealand affinities. - Antarctic Journal of the United Stat es 18 , 18- 20 .

SCHlND EWOl.F, O.H. 1955 . Uber einige kambrische Ga ttung inartik ulater Brachiopoden. - Neu es Jahrbuch fur Mineralogie ,Geolo gie und Palaontologie 1, 1- 15.

SCHI ND EWOLF, O. H. and SEILACHER, A. 1955 . Beitrage zur Kenntnis des Kambriums in der Salt Range (Pakistan). Akademi e del' Wissenschaften und del' Litera III1', Abhandlungen dermathematisch-natunl'issenschaftlichen Klasse 10.1- 341.

SCHUCHERT. e. 1893. A classification of the Brac hiopoda, - The Am erican Geologist 11. 14 1- 167.SHERGOLD. J .H ., COOPER. R.A ., M AcKINNON. D.1. and YOCHELSON. E.L. 1976 . Late Cambrian Brachiop oda, Mo llusca. and

Trilobita fro m Northern Victoria Land. Ant arctica. - Palaeontology 19, 247- 29 1.USHATINSKAY A, G .T. (YIII3TllI1 CKa5l , L T .) 1993. Panuc- 11 CPCtlII CK C~16pllii c KII C nI lll rYJIII/\bl CII611pCKOii n JI3TcjJOpMbl.

-r-Ilaneonmo.noeuwecxua »cypnan 2 (1993). 133-136.VORONOVA. L.G.. DROZDOVA. N.A .. ESAKOV A. N.V .. Z IIEGALLO. E.A .. ZIIURAVLEV, A.Yu.. ROZANOV, A.Yu., SAYUTI NA,

T .A . and USHATI NSKAYA, G.T. (Bopouona , Jl .L, [lp03t10Ba, H.A. , Eca xo aa, H.B. , )\(e raJIJlO, E.A., )\(ypanJlc n, AJO.,P0 3aHoB, A.lO., Ca ioruna, T.A., YwaTIIIICKa5l , LT.) 1987. Hcxonacuuc HlI)KH e rO KeM6pll51 rop MaKKcH311 (Kau ana) .- Tpyobt Ila neoum onoeu uecxoeo uucmumyma AH CCCP 22 4, 1- 88.

WAAC,EN, W . 1885 . Salt Range fossils. I. Produ ctus-limestone fossil s 4 (5 ). - Memoirs of the Geological Sur vey of India.Palaeontologia lndica 13 . 729-770.

WALCOTT, e. D. 1912. Cambrian Bra chiopoda. - Mono graph of the U.S. Geological Survey 51,1 - 872.WILLlAMS, A. and ROWEI.I ., A. 1965 . Morphology. In: R.e. Moore (ed .) Treatise on Invertebra te Paleontology, Part H,

Brachiopoda 1(2) , H57-H 138. Geo logical Society of America and University of Kan sas Press. Boulder, C olorado andLawrence , Kansas .

WIMAN, e. 1903 . St udien uber das Nordbalti sche Silurgebiet. I. OIe ne llu ssandstein , Obolussandstein und Ceratopygeschiefer.- Bullet in of the Geological Institutions of the University of Uppsala 6. 12- 76 .

WRONA, R. 1987. C ambrian microfossil Hadimopan el la Gedik from glacial erratics in West Antarctica. In: A Gaidzicki(ed .) Palaeontological Resu lts of the Poli sh Antarctic Expeditions. Part I. - Palaeontologia Polonica 49 , 37-48.

WRONA, R. 1989 . Cambrian limestone erratics in the Tertiary glacio-marine sediments of Kin g George Island , WestAntarctica. - Polish Polar Research 10. 533-553 .

WRONA, R. and Z IIUR AVI.EV, A.Yu . 1996. Early Cambrian archaeocyaths from glacial errat ics of King George Island (SouthShetland Island ), Antarctica . In: A. Ga idzicki (ed .) Palaeontologi ca l Results of the Po lish Antarctic Expeditions . PartH. - Palaeont ologia Polonica 55. 9-36 .

ZHURAVLEV. A .Yu. and G RAV ESTOCK, DJ. 1994 . Archaeocyaths from Yorke Peninsula, So uth Austra lia and arc haeocyathanEa rly Cambrian zonation . - Alcheringa 18 , I- 54 .

Palaeontologia Polonica, No. 55, 1996

L.E. HOLMER , L.E. POPOV and R. WRONA: EARLY CAMBRIAN UNGULATE BRACHIOPODSFROM GLACIAL ERRATICS OF KING GEORG E ISLAND (SOUTH SHETLAND ISLANDS), ANTARCTICA

PLATE 9

Eoobolus aff. elatus (PELM AN, 1986) 43

Lingulate brachiopods from the Cambrian glacial erratics (dropstones) found in the Cape Me lvilleFormation (Lower Miocene) at Melville Peninsula, King George Island, West Antarctica (Figs 1-2,5,9, 14), except for specimens of Figs 3-4, 6-8 and Figs 10, 13, 15 which are from Wirrealpa Limestone(samp le no . 13, L.E . HOLMER 1993), Flinders Ranges, Austra lia .

Fig. I. Ventral valve exterior, ZPAL Bp.xXXIII/33S9, erratic boulder Me66 , x 15.Fig. 2. Dorsal valve exterior, ZPAL Bp.xXXIIII6IS2, errati c boulder Me52, x 5.5.Fig. 3. Dorsal valve exterior, SAM P35111, x 12.Fig. 4. Ventral valve exterior, SAM P35109, x 40.Fig. 5. Ventral pseudointerare a, ZPAL Bp.XXXIII/6IS6, erratic boulder Me52, x 39.Fig. 6. Dorsal valve exter ior, SAM P3510 I, x 18.Fig. 7. a - ventral valve interior, SAM P35103, x 11 ; b - oblique posterior view of ventral valve interior and

pseudointe rarea, x 30.Fig. 8. Ventral pseudointerarea, SAM P35110, x 2 1.5.Fig. 9. Dorsal valve interior, ZPAL Bp.XXXIII/50S I, erratic boulder Me66, x 16.5.Fig. 10. Dorsal valve interior, pseudointerare a, SAM P35105, x 37.5.Fig. 11. a - ventral valve exterior, SAM P35102, x 10; b - oblique posterior view, x 50.5.Fig. 12. Ventral valve interior, SAM P35114 , x 19.5.Fig. 13. a - dorsal valve interior, SAM P35104 , x 15; b - oblique lateral view, x 30.Fig. 14. Ventral valve exterior, ZPAL Bp.xXXIIII33S6, erratic boulder Me66 ; a - oblique lateral view, x 25;

b - ventral view, x 22,5; c - pustulose ornamentation of post-larval shell, x 65; d - magnified fragment ofthe same surface as c, x 375.

Fig. 15. Complete juvenile shell, SAM P35 112; a - dorsal view x 55; b - oblique lateral view, x 55; c - obliqueposterior view of larval shell, x 195; d - pitted micro-ornamentati on of dorsal larval shell, x 760.

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FROM GLACIAL ERRATICS OF KING GEORGE ISLAND (SOUTH SHETLAND ISLANDS), ANTARCTICA


L.E. HOLMER, L.E. POPOV and R. WRONA: EARLY CAMBRIAN U NGULATE BRACHIOPODSFROM GLACIAL ERRATICS OF KING GEORGE ISLAND (SOUTH SHETLAND ISLANDS), ANTARCTICA

PLATE 10

Eoobolus triparilis (MATTHEW, 1902) . . . . . . . . . . . . . . . . . . 42

Specimens on Figs 1-9 are from the Middle Cambri an, Cape Breton, Nova Scot ia, Canada.

Fig. I. Dorsal internal mould, USNM '57013d, x 4.4.Fig. 2. Dorsal internal mould, USNM 570 13a, x 5.5.Fig. 3. Dorsal internal mould , ROM 678CM (E), x 5.Fig. 4. Dorsal valve exterior, USNM 57013e, x 5.7.Fig. 5. Dorsal internal mould, ROM SlOB, x 4.Fig. 6. Ventral internal mould, USNM 570 13c, x 4.3.Fig. 7. Lectotype, dorsal valve, ROM 510CM (A); a - exterio r, X 5.3; b - pustulose micro-ornamentation

of postlarval shell, x 7.5.Fig. 8. Dorsal internal mould, ROM 678CB (C), x 4.Fig. 9. Ventral valve exterior, USNM 51855a, x 5.

Botsfordia caelata (HALL, 1847)

Specimens on Figs 10 and 12 are from Lower Cambri an, northc entral Siberia, River Kotuj basin,Medve zhja River, (sample no. 182/3 of A.I. GROZDILOV 1963), except for the specimens of Figs 11and 13 which are from the Lower Cambri an, Troy (USNM loc. no. 2b), USA (New York State).

Fig. 10. Dorsal valve interior, RM Brl 36457; a - x 18; b - oblique anterior view, x 32.5.Fig. 11. Oblique anterior view of ventral pseudointerarea, USNM 34617 , x 21.5.Fig. 12. Dorsal larval shell with two tubercles, RM Brl 36456, x 19.5.Fig. 13. Posterior view of delthyrium and ventral pseudointerarea, USNM 366 17, x IS.

Botsf ordia pulchra (MATTHEW, 1889)

Specimen s of Figs 14-15 are from the Lower Cambrian, Hanford Brook Form ation (USNM loc. no.308d), Catons Island, New Brunswick , Canad a.

Fig. 14. Ventral valve exterior, ROM 164CM (A): a - x 4; b - oblique lateral view of umbonal area, x 8.Fig. 15. Dorsal valve exterior, ROM 164CM (B), x 4.

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L.E. HOLMER, L.E. POPOV and R. WRONA : EARLY CAMBRIAN UNGULATE BRACHIOPODS

FROM GLACIAL ERRATICS OF KING GEORGE ISLAND (SOUTH SHETLAND ISLANDS) , ANTARCTICA


L.E. HOLMER , L.E. POPOV and R. WRONA : EARLY CAMBRIAN UNGULATE BRACHIOPODSFROM GLACIAL ERRATICS OF KING GEORGE ISLAND (SOUTH SHETLAND ISLANDS ), ANTARCTICA

PLATE 11

Karathele napuru (KRUSE, 1990) 45

Specimens at Figs 1-5,7-8 are from the same Cambrian glacial erratic (dropstone) no. AEIMe52 foundin the Cape Melville Formation (Lower Miocene) at Melville Penin sula, King George Island, WestAntarctica; specimen of Fig. 6 is from the Wirrealpa Lime stone (sample no. 13, L.E. HOLM ER 1993),Flinders Ranges, Australia.

Fig. I. Complete juvenile shell, ZPAL Bp.XXXIII/2 9S8; a - oblique lateral view, x 55.5; b - oblique lateral viewof umbonal area, x lOO; c - posterior view, x 55.5; d - ventral view, x 45 ; e - umbonal area, x 80.

Fig. 2. a - dorsal valve interior, ZPAL Bp.xXXIIU53S4, x 27.5; b - oblique lateral view, x 32.5; c - obliqueanterior view, x 29.5.

Fig. 3. Dorsal valve interior, ZPAL Bp.XXXIII/34S3, x 30.Fig. 4. Dorsal valve interior, ZPAL Bp.XXXIII/50S2, x 19.5.Fig. 5. Ventral valve interior. oblique anterior view, ZPAL Bp.XXXIII/53S2, x 37.5.Fig. 6. Umbonal area of ventral valve, SAM P35113, x 80.5.Fig. 7. Umbonal area of dorsal valve, ZPAL Bp.XXXIIU50S3; a - oblique posterior view, x 100; b - dorsal view,

x 75.5; c - pitted micro-ornament ation of larval shell, x 215.Fig. 8. a - ventral valve interior, ZPAL Bp.xXXIIU5340U, x 10; b - detail of delthyrium, x 21.5; c - oblique

lateral view of delthyrium, x 50.

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L.E. HOLMER , L.E. POPOV and R. WRONA : EARLY CAMBRIAN UNGULATE BRACHIOPODS



L.E. HOLMER, L.E. POPOV and R. WRONA : EARLY CAM BRIAN U NGULATE BRACHIOPODSFROM GLACIAL ERRATICS OF KING GEORGE ISLAND (SOUTH SHETLAND ISLANDS), ANTARCTICA

PLATE 12

Karathele napuru (KRUSE, 1990) 45

Specimens of Figs 1-3 are from the same Lower Cambri an glacial erratic (dro pstone) no. AEfMe52found in the Cape Mel ville Formation (Lowe r Miocene) at Melville Penin sula, King George Island,West Ant arctic a; spec imens of Figs 4 and 5 are from Wirrealpa Limestone (sample no, 13, L.E . HOLMER1993), FIinders Ranges, Australia .

Fig. I . a - complete j uvenile shell, ZPAL Bp.xXXIIII38S4, dorsal view, x 60.5; b - lateral view, x 65.5;c - posterior view, x 90.5; d - oblique lateral view at ventral valve, x 75.

Fig. 2. Pitted microoma ment of larval shell, ZPAL Bp.XXXIII/29S5, x 1360.Fig. 3. a - ventral valve exterior, ZPAL Bp.xXXIII/54S3, X 11 ; b - umbonal area, ventra l view, x 40; c - pos terior

view, x 40 .Fig. 4. UmbonaI area of ventral valve, SAM P35 107; a - posterior view, x 40 ; b - obl ique lateral view, x 32.5.Fig. 5. Ventra l valve, pedicle opening, SAM P35100; x 50.5 .

Eotlzele spurri (WALCOTf, 1908)

Specimens of Figs 6-8 are from the Lower Cambrian tBonnia-Olenellus Zone), Pioche Shell (Co mbined Metals Memb er), north of Highway 93, west of Oak Spring Summit (locality no. 70-f-24;ROWELL 1980), Nevada (Linco ln County), USA.

Fig. 6. a - dorsa l valve exterior, USNM 459703c, x 19.5; b - umbonal area, x 90.5 .Fig. 7. Ventral valve, USNM 459703a; a - lateral view, x 32.5; b - posterior view, x 11; c - oblique poster ior view

showing pedicle foramen, x 30; d - lateral view of umbonal area , x 32.5 ; e - larval shell and pedicleforamen, x 75.5.

Fig. 8. Dorsal valve interior, oblique anterior view, USNM 459703b; x 40.

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L.E. HOLMER, L.E. POPOV and R. WRONA: EARLY CAMBRIAN U NGULATE BRACHIOPODSFROM GLACIAL ERRATICS OF KING GEORGE ISLAND (SOUTH SHETLAND ISLANDS), ANTARCTIC A

PLATE 13

Va ndalotreta djagoran (KR USE, 1990) . . . . . . . . . . . . . . . . . . .. 47

All specimens are from the same Cambrian glacia l erratic (dropstone) no. AE/M e52 found in the CapeMelville Formation (Lower Miocene) at Melville Penin sula, King George Island, West Antarctica(Figs 1- 9).

Fig. I . Complete shell, ZPAL Bp.XXXIII/54 S6; a - dorsal valve exterior, x 60; b - posterior view, x 80; c - obliquelateral view, x 80; d - oblique posterior view, x 150.

Fig. 2. Vent ral valve exterior, ZPAL Bp.xXXIIU54S5, x 32.5.Fig. 3. Complete shell, ventral valve, posterior view, ZPAL Bp.xXXIIU39S8, x 90.Fig. 4. Complete shell, oblique lateral view, ZPAL Bp.xXXIIU54S7, x 13.5.Fig. 5. a - complete shell, ventral valve exterior, ZPAL Bp.xXXIIU29S I, x 40 ; b - posterior view of comp lete

shell, x 37.5; c - posterior view of commissure and dorsal larval shell, x 110.Fig. 6. Ventral valve, ZPAL Bp.xXXIII/54S5; a - oblique posterior view, x 40; b - posterior view, x 80.5;

c - posterior view of ventral larval shell and pedicle foramen. x 150; d - pitted micro-ornamentation oflarval shell, posterior view, x 10 IO.

Fig. 7. Ventral valve, ZPAL Bp.xXXIIU5337U ; a - interior, X 40; b - apical process, x 60.Fig. 8. Dorsal valve, ZPAL Bp.xXXIIU5333U; a - interior, X 23.5; b - interior, x 50.5; c - pseudointerarea and

median buttress, x 100.Fig. 9. Ventral valve interior, ZPAL Bp.XXXIIU6 IS8, x 75.

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L.E. HOLMER , L.E. POPOV and R. WRONA : EARLY CAMBRIAN UNGULATE BRACHIOPODS


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