Cambrian brachiopod assemblage described from Antarctica. ... Cambrian brachiopods from Antarctica are poorly known; only five papers have been published ...
EARLY CAMBRIAN LINGULATE BRACHIOPODS FROM GLACIAL ERRATICS OF KING GEORGE ISLAND (SOUTH SHETLAND ISLANDS), ANTARCTICA
LARS E. HOLM ER, LEONID E. POPOV and RYSZARD WRONA
Holm er, L.E.. Popov, L.E . and Wrona, R. 1996. Early Ca mbria n lingul ate brachi opod s fro m g lacial errat ics of King George Island (So uth Shet land Island s), Antarctica . In: A. Gazdzick i (ed.) Palaeontological Result s of the Polish A ntarctic Ex pedi tions . Part 11. - Palaeontologia Polonica 55, 37-50. Erratic bould ers of Early Carnbrian age , occ urring withi n the Ear ly Mi ocene glac io-rnarine Ca pe Melvi lle Formation on King George Island (So uth Shetland Island s), Antarctica, have yielded lingul ate brachiopods including Eoo bo lus aff. elatus (Pe lman, 1986), Ka rathele napuru (Kruse, 1990) and Vanda lotreta djagoran (Kruse , 1990 ). It is the first Early Ca mbrian brac hiopo d ass emb lage described from Antarctica . Th e faun a is closely similar to that fro m the late Early Ca mbria n (Toyo nia n) Wirrealp a and Aroo na Creek Limestones, Flinders Ranges, So uth Austra lia. Th e new family Eoobolidae is prop osed and the type spec ies of Eoobo lus, E. tripa rilis (Ma tthew, 19(2) is re-fig ured. The evo lutio n of the botsfordiid s and acro thelids is disc ussed in the light of the new materi al of Karathele napuru, whic h is intermediate in morph ology between Botsf ordia and Eothe le. The superfam ily Acrot heloidea , is referre d to the order Lingulida because of the similarities in muscle sys tem, shell structure and deve lopment of pseud ointerareas. K e y wo r d s : Brachi opoda, Lin gul ata , taxonom y, Ca mbria n, Antarctica, South Shetland Island s. Lar s E. Holme r, Inst itut e ofEarth Scien ces, Departm ent ofHistorical Geo logy and Palaeon tology, No rbyvdgen 22, S -752 36 Uppsa la, Sw eden. Leonid E. POpOI', VSEGEI, Srednij Pr. 74, 199026 St. Petersbu rg, Russia. Rys zard Wro na, Instytut Paleob iologii PAN, A leja Zw irki i Wigury 93, 02-089 Warsw wa, Poland. Rec e ived 27 A pril 1995. ac ce pted 15 O ctob er 1995
38
LARS E. HOLMER , LEONID E. POP OY and RYSZ ARD WRON A
CONTENTS Int roduction
38
Acknowledgements Material
38 .
39
Systematic pale ont ol ogy
40
Class Ungul at a G ROJIINSKY et Po r o v . 1985
41
Order Lingulida W AIIGEN, 1885 .. . . .
41
Superfamil y Lingul oidea M ENKE, 1828
41
Famil y Eoobo lidae fa m . n.
41
Genus Eoobolus MIITrH EW, 190 2
41
Superfam ily Acrothe loidea W IILCOTr et SCHUCIIERT, 190 8 Famil y Bot sfordiidae SCHINDEWOLF, 1955 Genus Karathele KONEV11 , 1986
44 44
.
44
.
46
Superfamil y Ac ro treto idea SCHUCH ERT, 1893
46
Order Acr otretida KUIIN, 194 9
Fa m ily Acrotretidae SCH UCHERT, 189 3
46
Genus Vanda lotreta M ERGL, 198 8
46
.
48
References
INTRODUCTION Cambrian brachi op od s fro m Antarctic a are poorl y known ; o nly five pap er s have been published to dat e (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 m blages. Th e Early Cambrian brachi op od s described belo w were firs t rec or de d by W RONA ( 1989) from erratic boulders of lim estone wi thi n the Earl y Mi oc en e glacio- rnarine Cape Mel vill e Formati on of Kin g George Island , So uth Sh etl and Island s (Tex t-fig. I). Earlier, BRo c K and COOPER ( 1993) describ ed a si mi lar asse mblag e, including Eoo bo lus aff. elatus (PELMAN, 1986 ), Karath ele nap uru (KRUSE, 1990) and vandalotreta djagoran (KRUS E, 1990) from the lat e Ear ly Cambrian (Toyo nian) Wirrealp a and Aroon a Cree k limeston es, Flinders Ran ges, South Au strali a. The poorl y known type spec ies of Eoobo lus, E. tripa rili s (MATTH EW, 190 2) is re-fig ured to d raw co mpa riso n with Eoo bolus aff. elatus (PELMA N, 1986). Previou sly, most Early and Middle Ca mbr ian lingulides of this type have been referred usuall y to " Ling ulella" or "O be lus" , In co ntrast to these ge ne ra, Eoobolus has a pitt ed larval she ll, much like th at of the acrotretoi ds , in addi tio n to a distinctive granulated pos t-larva l she ll. Belo w we prop ose the new fami ly Eoo bo lidae to include th is ty pe of linguIides. Th e evo lutio n of the botsfordiids and ac rothe lids is discu ssed in the ligh t of the new mat eri al of Karathele nap uru ; it is possibl e to tr ace a gradua l tra ns ition from botsfordi ids, suc h as Karath ele, to acrothe lides , suc h as Eothele. Th e mu scl e sy ste m, she ll struc ture , and pseud ointerareas of the bot sfordiids are very similar to those of Earl y Pal aeozoi c Ob olidae (WALCOTT, 191 2; Po POv, 1992 ), and the superfamily Ac ro the loidea, is here referred to the Order Lin gul ida.
Acknowledgements. - Fred erick J. COLLIE R, Jann THOMPSON, Rex DOESCII ER, and the late Richard GRANT (Was hing ton, D.e. ) ass isted by arra nging fac ilities to study mat erial store d in th e U.S . National Mu seum. J anet W ADD INGTON (Toronto) arrange d the loan s from collec tio ns in the Royal Ont ario Mu seum. This work has been supported by gra nts (to L. HOLMER) from the Swedi sh Natural Scien ce Rese arch Council (NFR). Leonid Po POV gratefully ackn owl ed ges receipt of a one year NFR visiting scientis t gra nt and a 10 month visiting sc ientist grant from the Royal Swedi sh Acade my of Scien ces (KVA) th at have enabled him to wo rk ex tens ive ly at the In stitute of Earth Sci en ces, Department of Historic al Geology and Paleontology, Uppsala Uni versit y, as well as two Visitin g Sci entist grants from the Smithsonian In stitution .
39
CAMBRIAN BRACHIOPOD S FROM ANTARCTICA 58' W
59' W
f--------_+_
~~ ~\>Q -------,-=-"""""----
--:;;:=-~.----j
62' S
O'
King George Island
..¥ o
o
W
E
o,
Fig . I Location map of King George Island (arrowed) in Antarctica, and the outcrops of glac io-m arine strata (asterisked) of the Ear ly Miocene Cape Melville Form ation .
Lars HOLMER'S fie ld work in So uth A ustralia was fina nced by NF R, and arra nge d by Bru ce R UNNEGAR (Los Ange les) . R. WRONA is deepl y indebte d to Stefan BENGTSON (U ppsala) for the arra ng ing of a o ne month visi ting gra nt from the Royal Swedish Acade my of Sciences, for his immeasu rabl e help during the course of this wo rk which was extende d to three mo nths in Uppsala, and fo r the use of laborat or y and SE M fac ili tes at the Depar tment of Pal aeontol ogy, Uppsala Un iversity. WRONA'S field work in Antarc tica was undertaken as par t of the Fifth (1980-8 1) and Tenth ( 1985-86) Poli sh An ta rctic Ex peditio ns organized and fi nance d by the Po lish Acade my of Sc iences. T he investigatio n of Antarc tic material in the Institute of Paleobiol ogy, Po lish Academy of Sc ience s, was supported by Project M R. I. 29. The manuscript has ben efited from comments on both the lan g uage and the sc ientific content by Michael G . BASSETT (Cardiff), Gertruda BIERNAT (Warszawa), Peter D. KRUSE (Darwi n), John S. PEEL (U ppsala), and by Ant ho ny D. WRIGIIT (Be lfast) . A ll SEM micrograph s were taken at the Electron Mi croscop y La boratory of the Dep artment of Zoo logy, Uppsala Uni versity. Thi s work is a co ntribution to the IGCP Proj ect 366.
MATERIAL Th e Antarctic brachiopod ma ter ia l described in this paper co me s fro m the res idue of lim eston e erratic boul ders etc he d with 10% acetic aci d . T he bo ulders are scattered as iceb erg-rafted drop ston es within the Early Mioce ne glacio-marine sedi me nts of the Ca pe Me lv ille Formation, ex posed in the eastern most part of King George Island, So uth Shetla nd Islands (Tex t-fig. I). Th e for ma tio n is re presented by a 200 m seque nce of highl y fossili ferou s sha les with siltstone, marl and sa nds to ne int ercalat ion s (B IRKENMAJER et at. 1983). Th e indigen ou s fossi l assemblage of the Ca pe Mel ville Fo rma tion co mprises of di atoms, chrysomona d cysts, si licoflagellates, ca lcareous an d are naceous fo ra mi nifera , so lita ry corals, pol ych aetes, bryozo an s, gastro pods, bi val ves, decapods, echinoi ds, asteroids and fis h re ma ins (for references see
40
LARS E. HOLMER, LEONID E. POPOV and RYSZARD WRONA
GAZDZI CKI ed. 1987 ). Th e glac ial charac te r of the sediments is ev ide nce d by the presence of numerou s, poorl y ro unded or sharp-edged boulder s of rocks exotic for King Geor ge Island, with glacially -g enerated features (BIRKENMAJER et al. 1983 ; BIRKENM AJER and B UTKI EWI CZ 1988; W RONA 1989 ). Th ese strata also yield allochtonous Cre taceo us calc ar eou s nann opl ankton (D UDZIAK 1984 ) and belemnites (B IRKENM AJ ER et al. 1987) dropped int o the se dime nt fro m melting iceb er gs. Th e Early Mi ocen e age o f the formation , and co nseq uently of the Mel vill e Gl aciati on , was determined by biostr ati gr aphi c dat a (BIERNAT et al. 1985; GAZDZI CKI ed. 198 7) as we ll as by rad iometri c dating (c f. BIRKENMAJER 1992 ). Th e lim estone erra tics were co llec ted fro m the erosio na l surface of the Mel vill e Penin sul a and are hou sed at the In stitute of Pal eobiology, Poli sh Ac ademy of Sc ien ces in Warszawa (abbrev iated as AElMe). This coll ecti on co ntains severa l hundred boulder s groupe d int o di stinct lith ol ogical typ es (W RONA 1989 ; WRONA and ZHURAVLEV 1996 thi s vo lume) . Ver y characte ristic boulder s of the light co lo ure d calcimic ro bial-a rc haeocyatha n frameston e (ty pe 11 ; W RONA 19 89) co ntain rare she lls of Eoobo lus aff. elatus and an abunda nt blu e-green al gae Gordonophyton-Renalcis and arc haeocyathan assemblage of the sa me age as the uppermost Bot om an units with Syringocnema favu s Bed s di stingui sh ed in South Au st rali a (Z HURAVLEV and G RAVESTOCK 1994 ; WRONA and Z HURAVLEV 1996 thi s vo lume) ; they ca n be also co rre lated w ith faci all y si milar beds of the Sh ackleton Lim eston e ex posed in the Tran sant ar cti c Mountains (DEBRANNE and KRUSE 1986, 1987 ; REES et al. 1989; ZHURAVL EV and G RAVESTOCK 1994 ). Lin gul ate bra chi op od s identified as Eoobolus aff. elatus are abunda nt and we ll pr eserved in black , so me ti mes slig htly bituminous pack ston e and wack eston e (ty pes 12-14; W RONA 1989). Th ese kind s of lim eston e erra tics also co ntain arc haeoc ya ths, spo nge spic ules , gas tro po ds and hyoliths as we ll as a di ver se assem blage of probl em ati ca (s ma ll shelly fossil s), amo ng them numerous sclerites of the tommot iid Dail yatia, which are being describ ed separately (B ENGTSON and WRONA, in p rep aration). Th is sma ll she lly fossi l assemblage ca n be co mpared with a simi lar fauna and sed ime nts of the Parara Limeston e (Bo toman) in South Austra lia (BENGTSON et al. 1990 ; ZHURAVLEV and GRAV ESTOCK 1994 ), but on th e othe r hand it can also be corre lated with an autoch tono us A ntarc tic ken nardi id asse mblage recorde d fro m the Shac kleto n Lim eston e in the Tran sant 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 lotreta djagoran and Eoobolus aff. elatus orginate exc lusive ly from one erratic boulder: AE/Me52 . T his com pletel y etched sma ll boul der, size: 4 cm x 5 cm and I cm thic k, co m pr ise d a brachi op od co quino id lim eston e; it was the most fossi liferous sa mp le st ud ied and also co nta ine d cha ncello ri id and spo nge spic ules, gas tro pods, hyoliths and phos phatize d trilobite carapaces. Th e age of th is erratic ca n be es ta blish ed as late Ea rly Cambrian (Toyonian) by the brachiopo d asse mb lage described below, which is closely si m ilar to the brachi op od fauna fro m the late Ea rly Ca mbrian Wirrealpa and Aroona Creek lim eston es, Flinders Ran ges, South Aus tra lia. Equiva lent stra ta co ntai ning these brach iopod co quino id lim eston es has not ye t been discovered o n the A nta rctic landmass. Ho wever, the ge neral lith o log ical co mpos ition of the who le spec tr um of erratics fro m the Ca pe Me lvi lle Forma tion clea rly suggests th at o utcro ps of Low er Ca mbrian roc ks aro und the Wedde ll Sea (in particul ar the Arge ntina Range) and north ern Tran sant arcti c Mountain s ac ted as so urce areas for the g lac ia l boulders (WRONA 1987; BIRKENMAJER and B UTKIEWICZ 1988 ; W RONA 1989). New materi al of Eoobo lus aff. elatus and Karathele nap uru fro m the Wirrealpa Limeston e, Flinders Ran ges (Te n Mile or Mount Bill y Creek sec tion within the Wilkawillina Go rge; see KRUSE (19 91 a) fo r localit y da ta) is illustrate d belo w for co mpa riso n with the A ntarc tic material. Karathel e napu ru and Vandalotreta djagoran we re descr ibed origina lly by KRUSE ( 1990) fro m the ea rly Middle Cambria n Tindall Limeston 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, len gth, height of va lve ; Iw, 11 = wid th, len gth of dorsal pseudointer area ; Pw = wid th of medi an gro ove ; Cw, C l = w idth, len gth of ca rdina l mu scl e field ; VI = len gth of visceral area; SI = len gth of dorsal median rid ge. S = sta nda rd deviation, N = number of measurements, MA X = maximum value, MIN = minimum value .
CAMBRIAN BRACHIOPODS FROM ANTARCTICA
41
Th e terminology of the lin gul ate brachi opods used here mainly fo llo ws th at of Ro wELL (1965); KONEVA (1986); a nd HOLM ER (1989). Th e term med ian tong ue was introduce d by PoPOV and HOLMER (1994) to den ote the na rrow ant eri or ex te nsion of th e dorsal viscera l field of man y lin gul ates (Te xt- fig s 2-3). The illu str ated and/o r di scu ssed material is dep osit ed in th e Insti tut e of Pal eobi ology, Pol ish Academ y of Sciences, War szawa (Z PAL), So uth Aus tralia n Mu seum, Ad el aid e (SA M), Northern Territory Mu seum of Arts a nd Sciences, Dar win (P) . Un ited States Nati on al Mu seum, Was hi ngton D .e. (USNM), Swedi sh Mu seum of Natural Hi story (RM), Stockho lm, and th e Royal Ontar io Mu seum , Toronto (RO M) .
Class Lingulata GROJANSKY et Porov, 1985 Ord er Lingulida W AAGEN, 1885 Superfamily Linguloidea M ENKE, 1828 Fa mi ly Eoobolidae fam . novo
Diagnosis. - Sh ell dors ibicon vex, so mewha t inequ iva lve d, e longate oval to subtr iang u lar; lar val she ll we ll defined, with pitted micro- ornam en tation; postl arval she ll fine ly pu stul ose ; ve ntra l pseudointerar ea elevated a bove va lve floo r, w ith deep pedi cl e groove and we ll-develope d flex ure lin es; dorsal pseudointer area d ivide d, raised above va lve floor ; mu scle sys te m wit h paired umbon al mu scl e scars bisected by v-sha ped impression of pedicl e ne rve ; dorsal visceral fie ld wi th we ll-developed medi an ton gue ex te nd ing to mid-len gth ; mantle canals bac ulate with we ll- develo ped vas cu la medi a. Gen era included. - Eoobolus M ATTH EW, 1902 (= Clivosilingu la USH ATINSKAYA, 1993); Vassilkovia PoPOV et KHAZANOV ITCH (in Po POV et al. 1989 ). Discussion. - The Eoobo lidae show a co m bination of c harac ters , that are un iqu e wi thi n the Lin gu loid ea. T here are no other lingul oids with a fine ly pi tte d larval she ll and pus tulose pos t-larval she ll. O ther types of pitte d micro- orn amentation are distri buted w ide ly with in the superfami ly, but usu all y invo lve both the larval and post-larval she ll as in th e Z hanate llidae, Elka n iidae, Dysorist idae, and th e Paterul idae. It is po ssibl e th at the eoobo lids are re lated so mehow to th e Botsfordiidae, the ea rliest m embe rs of whic h are also characterise d by pi tted lar val and pustul ose post-l arval she lls . Th e det ailed morphol ogy of the larval she ll of vassi lkovia PoPOV et KHAZANOV ITCH (in Poro v et al . 1989), is no t known, bu t the pu stulose post-larval she ll is closel y simi lar to th at of the Eoobolidae. T he fa mily is firs t recorded from the Botoman (U SIIATINSKAYA 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 AT INSKAYA. p. 125 [Type species: Lingulella cli vosa PELM AN. 1983 ; Midd le Cambrian (Amg ian). Siberi a].
Type specie s: Obo lus triparilis MATrIlEW. 1902 (se lected by ROW ELL 1965. p. H263).
Diagno sis. - She ll do rsibiconvex, e longate suboval to elongate subtriang ular; ventral va lve slig ht ly ac um inate with high , ort hoc line pseudo interarea ; ped icl e groove deep, narro w, wi th subparalle l lat eral margins ; do rsa l pseudo inte rarea wit h broad, shallow med ian groove; propareas of both va lves with flexure lines; ve ntra l viscera l fie ld not exte nd ing to mid-valve ; paired ve ntra l um bon al muscle scars b isect ed by v-s haped imp ression of pedic le nerve ; ve ntra l vascula lateralia sub marg ina l, straight, slightly d ivergent pro xim all y; dorsal visceral area w ith median ton gu e usu all y bisected by med ian rid ge and pair of subme dia n ridges ; dorsal vascula lat eral ia marginal, arcuate; vascula media lon g , s lig htly divergent. Other species included . - Ling ule lla wanniecki REDLlCH, 1899; Ea rly Ca mbrian, Pak ist an (Sa lt Ran ge). Ling ulella clivosa PELMAN, 1983; Middle Cambrian (A mgian), Si beria. Ling ulella elata PELMAN, 1986 (= Clivosiling ula dilata ta USHATINSKAYA, 1993); Middle Cambria n (Amgian), Siberi a. Eoobolus aff. elat us (PELMAN, 1986); Early Cambrian (Toyonian), So uth A ustra lia, West Antarcti ca (King Geor ge Isl and ). Discussion. - Eoobo lus has been regard ed as a junior sy no ny m of both Lingul ella (WALCOTT, 1912), Obolus, and Ung ula (ROWELL, 1965; the fig ure d va lves prob abl y represent Ung ula) . However, it see ms that und oubted species of Ling ulella a nd Ung ula are not kn own earlier th an fro m th e Lat e Cambri an ; only Lingulella is int ern all y sim ilar to Eoo bolus, but th e former has poorly de vel op ed dorsal flexure lines and pse udo interareas that are not stro ng ly ra ise d above the va lve floor (Porov and HOLMER 1994). Obol us
42
LARS E. HOLMER, LEONID E. POPOV and RYSZARD WRONA
.,
---- - w- - - -- _ ---- I w ---~
umbonal
: r-
posterolateral muscle field I
impression of gastroparietal ?ands
I
L
- --- -- - - ------ --- - --~-------~
A
anterior muscle field
median tongue
B
anterior lateral
Fig. 2 Eoobolu s aff, elatus (PELMAN, 1986 ), reconstru ction o f vent ral (A) and dorsal interi or (B) . Location of measurements indi cat ed : 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 sc andia and is not similar int ernall y to Eoobolus (Porov et al. , 1989 ). Moreover, our bri ef re- study of the type species Eoobolus triparilis (M ATTH EW, 1902) revealed a di stinctive pustu lose ornamentation on the post-larval she ll, of a type which is not known from any other obolid genera with the exception of Vassl lkovia Por ov et KHAZANOVITCH (in Por ov et al., 1989 ) and Cli vosilingu la USH ATINSKAYA, 1993. E. triparilis al so has a characteri st ic lon g median ridge and a pair of lateral ridges in the dorsal valve . A similar typ e of morphol ogy was recorded in the typ e species of Clivo siling ula, C. clivosa by PELM AN ( 1983 , p. 125) and the pseudointerareas as well as mo st other morphologic al characters are closely similar; Clivosi lingula is here regarded as a junior sy no nym of Eoob olu s. According to USIIATINSKAYA ( 1993) Lin gulella viridis COBBOLD, 1nl , Lingulella rotunda PELMAN, 1977 , and Lin gul ella variabilis PELMAN, 1977 , may be also referr ed to Clivo si lingula , but the det ailed morphology of the se taxa is poorl y kn own. Lingulell a wanniecki REDLICH , 1899 , fro m the Ea rly Cambrian of Paki stan ha s a we ll defined median rid ge and a pair of later al rid ge s in the dorsal val ve as well as a di stincti ve pu stulose ornamentatio n (SCHINDEWO LF and S EILACHER 1955); it is referred her e to Eoo bo lus.
Eoobo lus trip arilis (MATTHEW , 1902 ) (PI. 10: 1-9) 190 2. Obo lus trip arilis MATrl IEW, p. 94, pI. I : 2a-c . 191 2. 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 hori zon and locali ty: Middl e Cambrian (Bo uri not Group), Ca nada (Ca pe Breton ).
Diagnosis. - Sh ell dorsibicon vex , slightly elon gate subtriangular, with maximum width so mewha t anterior to mid-length ; ve ntral valve slig htly acum ina te ; ve ntral pseudointerarea high , tri an gular; dorsal interior with lon g medi an rid ge and pair of sub me dian rid ges bounding narrow median ton gue . R ema r ks. - The micro-ornam entation of the larval shell could not be determined on any of the avai lable specimens. The fragmentary pre servation of the material at hand made it diffic ult obtain good measurements . Materia l. - Fi gured. 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) . O ccur r ence . - Type localit y only.
43
CAMBRIAN BRACHIOPODS FROM ANTARCTICA
Eoobolus aff. elatus (P ELMAN, 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 SHAT INSKAYA, p. 133, fig. 1: 10-11. 1993. Lingulella sp.; B ROCK and C OOP ER, p. 780, fig. 14: 14-15.
Material. - Figured from Antarctica. Ventral valves: ZPAL Bp.XXXIII/33S6, 33S9, 61 S6. Dorsal val ves: ZPAL Bp.xXXIII/50S1 , 61S2. Figured from Au stralia. Complete juvenile shell: SAM P35112. Dorsal val ves: 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
L N X S MIN MAX
27
( PELMA N) ,
S MIN MAX
IwlW
1.17
1.29
135%
0.513
0.545
6.1
0.60
0.28
0.52
0.64
3.1 20%
5.9
1.04
7.46 120%
58%
40 %
5.28
4.28
1.40
2.56
2.36
150%
32 %
79%
63 %
20
20
26
VVL
0.62
W
27
IVL
0.264
L
17
LIW
1.79 0.882
( PEI.MAN),
26
VI
1.117
2.39
26
Iw
26 27%
Table 2. Eoobolu s aff. elatus
N X
11
W 27
average dimensions of ventral valves.
68%
17 50 %
average dimen sions of dorsal valves.
Iw 17
LIW 27
IwlW
VIIL
26
26
17
27%
68 %
50 %
2.27
1.81
0.860
0.700
0.464
3. 1
5.9
6.1
1.06
0.84
0.52
120%
20%
58 %
40%
4.32
3.48
2.40
150%
32%
79 %
63%
1.12
135%
IVL
7.46
Description. - Shell slightly dorsibiconvex , elongate suboval, on average 135 % as long as wide, with maximum width somewhat anterior to mid-length . Ventral va lve gently and evenly convex, slightly acuminate; ventral pseudointerarea high triangular, orthocline, on avera ge 27 % as long as wide , occupying 68% 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. Dor sal 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 to circular, around 0 .10-0.15 mm across , ornamented by fin e c ircular pits, around I m acro ss. Post-larval shell co vered by fine , den sely spaced pustules, around 5-6 m acro ss. Ventral viscera l area some what thickened, ext ending to mid-length; ventra l umb onal muscle scar pair ed , bisected by v-s haped impression of pedicle nerv e; vent ral vasc ula lateralia subperipheral, slightly arc uate. Dorsal visce ral area, with narrow median ton gue ex tending somew hat anterior to mid -val ve ; dorsal median ridge weakl y developed; dor sal vasc ula lateralia margin al, arcuate; vasc ula media weakl y impressed. Discussion. - The spec ime ns fro m Antarctica are similar to Eoobolus elatus (PELMAN ) from the Kuonamka Formation of Sib eri a (USHATINSKAYA 1993, figs 1-11 ) in size, general outline, and in havin g a weakl y de veloped dor sal medi an ridge, as well as in the absence of dor sal submedian ridges; the onl y differen 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 ca nnot be compared in detail with the Antarctic materi al. A second species, Eoobolus dilitatus described by USHATINSKAYA (1993, fig. 1: 5-9) from the same horizon, differs from Eoobolu s etatus in bein g transversely oval and wider, but the se features are known to be very variable in lingul ids and moreover, change significantly dur ing ontogeny; thu s, E. dil itatu s may be a junior synony m of E. etatus . The Antarct ic material of E. aff. elatus is also clo sely similar to Lingul ella sp. de scribed by BROCK and COOPER (1993 ) from the Early Cambrian Wirrealpa and Ram say limestones of the Flinders Ranges and Yorke Peninsula, South Au str alia . 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 cat es that they may be con specifi c, as the Au stralian specimens have an identical type of extern al and internal morphology. Occurrence. - Earl y Cambrian (Toyonian), South Au stralia ; King George Island erratics , Ant arctica; Middle Cambrian (Amgian), Sib eria. Sup erfamil y Acrotheloidea WALCOlT et SCHUCH ERT, 1908 Diagnosis. - Shell with short, con vex posterior mar gin ; postl arval orn ament of fine, evenly distributed gra nules or pustul es; larv al shell pitt ed , with apica l spines or tub ercles; ventral valve convex or low conical; ventral pseud ointerarea ves tigial or lacking; pedicl e eme rging through delthyrium or for amen posterior to apex ; dorsal pseudointerarea vestigial or lacking; muscl e sys tem linguloidean-like, with paired umb onal mu scle; mantl e canal sys te m of both valves bacul ate; vascula media we ll-deve loped; she ll struc ture bacul ate. Discussion. - The close relation ship bet ween the Botsfordiidae and Acrothe lidae has lon g been recogn ized (ROWELL 1965 ). As noted by ROWELL (196 5) it is prob abl e that the botsfordiides were ancestral to the acrothelides; moreover, it now see ms possibl e to trace an almost continuous gradual transiti on in morphology from Botsfo rdia (PI. 10: 10-15 ; Text-fig. 3A-B ) through the intermed iate Karath ele, with a deep , but unr estricted delth yrial ped icle opening and rud imentary ventral pseud ointer area (Text-fig. 3C- D) to ear ly acrothe lides, like Eothele (PI. 12: 6-8) with a low, subco nical ventra l valve, a reduced pseud oin terarea that is gradually transform ed into the pedi cle tub e, and with an elonga te, suboval ped icle fora me n that is formed post-l arvall y (PI. 12: 6- 8). The muscle sys tem of botsfordiides is very similar to that of Early Palaeozoic Obol idae (W ALCOTT 191 2; POPOV 1992); the earliest known ge nera, Botsfordia and Edreja have sca rs that appea r to match a full set of obolide mu scles (Text-fig. 3A-B), whilst the numb er of mu scles in mo st ac rothe lides was prob abl y reduc ed . The scars of a lingul oid type of v-shaped pedi cle nerve in Botsfo rdia and Edreja (Text-fig. 3A) also may indicate a close relat ion ship between the two gro ups. The she ll 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 gro up is still poorl y known . Altho ugh the botsfordi ides and acrothe lides have some characte rs in common with the acro tretoids, such as a pitted larval she ll and a pedi cle fora me n (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 rel ated close ly to the lingulid family Eoo bolidae, which is also charac terized by a pitted larval shell and a pustul ose-granul ar post-larval orn ament ati on . WILLlAMS and ROWELL (1965 ) prop osed that the acrothe lide s prob abl y were ances tra l to the discinides, but in view of the co nside rable differences in their ontoge nies, this is unlikely (C HUANG 1971 ; HOLMER 1989). Family Botsfordiidae SCHINDEWOLF, 1955 Diagnosis. - She ll bico nvex; ventra l pseud ointerarea ves tigia l, divided by deep pedicle groove formin g triangul ar delth yrium; dorsal pseud oint erarea ves tig ial, divide d by medi an groove; mu scle sys te m co nsisting of paired umb on al, transm edi an , outs ide lateral, intern al later al, anter ior lateral, and ce ntral muscles; vasc ula lateral ia straight, submedian, di vergent in both valves ; larv al she ll with one to three apica l tub ercl 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: M idd le Cambri an (Peron opsis ? ultimu s and Ptychagnostus atavu s Zo nes ). Kazakh stan (Malyi Karatau ).
Diagnosis. - Shell subequibico nvex, subcircular ; ventra l pseud ointerarea narrow triangul ar, procline to catacline, with vest igial prop areas; pedi cle groove deep , with stee p lateral slopes; dorsal va lve slig htly sulcate ; dor sal pseud oin terarea ves tigial, with poorl y defined med ian groove; ve ntral larval shell with three tubercles and one or two pairs of dorsal tub ercl es; ventra l visceral field sma ll, short, onl y slightly thickened ; dorsal visce ral field small, slightly thickened ; med ian ton gue narr ow, extending to mid- valve, bisected by medi an rid ge ; mantl e ca nal sys tem bacul ate ; both valves with straig ht, submedian, divergent vascula lateralia ; vasc ula media long, di vergent.
45
CAMBRIAN BRACHIOPODS FROM ANTARCTICA pedicle groove
umbonal
proparea
impression of gastropa rietal bands
vascula media
median tongue
A
impression of pedicle nerve pedicle groove
transmedian
B umbonal transmedian anterior lateral outside lateral
proparea
-...r--- - ....L
middle ----''------lateral central vascula teterelie
-t-- - .
median tongue
c
vascula media
D
Fig . 3 Recon stru ction of vent ral (A) and dorsal interior (B) of a ge neralised Botsfordiidae and ventral (C) and dorsal interior (D) of Karath ele napuru (KRUSE, 1990 ).
Other species included . - Eotlzele napuru KR USE, 1990; Early Cambrian (Toyoni an)-early Middl e Cambrian, Australia, Antarctic a. Discussion . - Karath ele was referred originally to the Acrothelidae by K ONEVA (1986 ); howe ver, it has an open delthyrium throu ghout onto geny, with a ventr al pseudointerarea and ped icle groove much like those of Botsfordia and Edreja (Text-fig. 3). Karathel e differs from both Botsfordia (PI. 10: 10-15 ) and Ed reja mainly in having a procline to catacline ventral pseudointerarea with vestigial propareas, small , densely spaced post-larval pustule s, one pair of ventral lar val tubercle s, as well as a short ventral viscer al area that does not extend anteriorly to mid-v alve. Externally Karath ele 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 ele spurri are similar to Karath ele in the morphology of the larval shell , delthyrium and rudimentary pseudointerareas, but unlike Karath ele, 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 and cannot be compared dire ctl y. Karath ele napuru (KR USE, 1990) (PI. 11: 1-8; PI. 12: 1-5 ; Text-fig. 3C-D) 1990. Eoth ele 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; BRo c K and COOPER, p. 78 2, fig . IS : 1-14. Holotype : Ventr al valve SAM P85206. Type horizon and locality : Middle Cambrian , Tind all Limestone (bore -hol e NTGS 83/3), Daly Basin , Northern Territory, Australia.
46
LA RS E. HO LME R. LEON ID E. I'OP O V an d RYSZ AR D WRO NA
Material. - Figu red fro m A ntarctica. co mp lete she lls: ZPAL Bp.XXXIII/29S8, 38S4 ; ventr al va lves : ZPAL Bp.XX XI1I/53S2, 5340U, 54S 3; do rsa l va lves: Z PAL Bp .XXXIII/29 S5 , 34S3 . 50S2, 50S 3, 53S4. Figured from Australi a. Ventr al valves : SAM P351 00. SAM P35 107, SAM P35 113. Me asu re men ts: See Tabl es 3--4 . Tab le 3 . Karathele 11
11
11
X
11
S i\IIN 11
MAX
II
Il a p ll nt
I.
1.12 0.242 0.82 1.40
I
(KRUSE). av erage d ime ns io ns o f 5 vent ral val ve s.
I
W
1.18 1 I 0.331 I I 0.76 I 1.46
Iw
.1
0.33 0.130 0. 16 I 1.34
I
I' w
0.16 1 II 0.038 I 0. 10 I 0.20
1
1
I
VI
Vw
0.38 0.060 0.32
0.4 3 0.103 0.32
I
0.50
I
I OAO
i-
I
1
LlW
Iw/W
979c 8.6 84 'le I089c
279c 5.9 2 1'le 349c
I
1
1
VI/I.
349c I I 3.8 I 299c I 39%
I
1 I
!! 11
Table 4 . Karath ele napuru ( KRlISE). av erage d im e ns ions o f 6 dorsa l va lves.
I
I.
X
IA I
S
0.334 1.02 2. 14
MI N MAX
W
Iw
1.59 00409 1.1 6 2.14
0.38 0.083 0.28 1.22
SI
I
1'~ _~
0.28 0.58 1.22
LlW
Iw/W
89'7e
24%
2.5 85% 92%
3.5 20% 29%
SilL 60% 604 52% 69%
Dia gnosis. - Pedi cl e groove deep , for ming narrow slit in mature specimens; do rsal larval she ll with two tubercle s. Descript ion of materia l fro m A nta rct ica : She ll s lightly ve ntribico nvex, subcircular. about 97 % as lon g as wide. Ventral valve low subconica l with maximum hei ght at umbo ; ven tra l pse udoi nterarea narrow tria ngular. proc line to slightly catac line, occupying on avera ge 27 % of valve widt h: pedic le groo ve very deep . forming high tria ngular to scmiov al delth yr ial ope ning: posteri or part of delth yrium forming narro w slit in mature spec ime ns ; ve ntra l propareas vesti gial. Dor sal valve ge ntly con vex with maximum heig ht at abo ut umbo: dor sal pse udointer area vesti gial. oc cupy ing abo ut 24 % of valve width. Ventral larval she ll with high medi an tubercle above delt hyri um and pair of tub ercl es in postero lateral part. Dor sal larva l shell with two high tube rc les posterolatera lly. Ventral visce ral area small. oc cupy ing about 34% of valve len gth and 38 % of valve width : vascula lateralia of bot h va lves submed ian. straight. di vergen t. Dor sal interi or with slig htly raised viscera l area occ up ying about 60 % of valve lengt h: dorsal median ton gu e bisect ed by lo ng rid ge. Discu ssion . - Kara thele napuru differs from the typ e spec ies , K. co ronata (KONEV,\ , 1986 ). in havin g on ly two tube rcles o n the dorsal larval she ll, and a much deep er pedi cle groove . T here are virtually no di ffe re nces between our material and the material described by KRUSE ( 1990 , 199 I b). However, the maxi mum recor ded size from the A ntarctic (max L = 2 . 14) is much sma ller tha n that recorded by KRUSE (1990: max L = 7 .6) , and the de lthyri um rema ins ope n thro ugho ut ontogeny in the A ntarctic spec ime ns, whilst the larger va lves described by KRlJSE ( 1990) have a fine slit or s uture. A similar type of de lth yrium is also found in the large r sp ecim en s from the Wirrea lpa Lim eston e, Fli nders Ranges (PI. 12: 4-5 ), but in all know n spec ime ns of K. napuru the delth yria l mar gin s rem ain separated (eve n thou gh they mig ht "to uc h" one another) and do not join comp letel y as in mature she lls of Eot hele (P I. 12: 6-8). Occurrence . - Ear ly Cambrian (Toyoni anj-early Midd le Ca mbr ian. Au stral ia (Nothern Territory). King George Island erratics, A ntarcti ca.
Order Acrotretida
K UHN ,
1949
Superfami ly Acro treto idca SCII UCII ERT. 1893 Fami ly Acr ot r eti d a e SCH UCII ERT. 1893 Genus 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 spec ies : vanda lot reta vafra M EJ{(;L. 1988. Typ e hori zon and localit y: Mi ddl e C am brian (A mg ia n). Hig h A tlas ( Yag o ur inli e r). M or occo.
47
CAMBRIAN BRACHI OPODS FROM ANTARCTICA
Diagnosis. - Sh ell tran sversel y o va l to subcircular with narrow, convex po sterior margin; ventral val ve wide conical ; ve ntra l pseudointerarea procline to ca tacl ine, poorly defined laterall y with int ertrou gh : for am en not e nc losed w ithin larval she ll; dor sal va lve weakly convex; dorsal pseudointer ar ea shor t with bro ad ly triang ula r medi an groove ; ap ica l process for mi ng bo ss-like thi ck ening ante rior to int ernal for ame n; dor sal viscera l fie ld with narro w me d ia n ton gu e , usu all y bi sect ed by ves tig ia l med ian ridge ; dorsal medi an buttress de vel op ed . Other species included. - Acrotre ta pompeckji M ERGL et SLEIIOFERovA, 1990; Middle Cambri an , Jin ce Formati on , Ce ntra l Bohemi a (Czec h Republi c ). Hadrot reta dja goran KRUSE, 1990; Ea rly C ambri an (Toyonia n)-early M iddle Ca mbr ian. A ustralia, West A ntarctica. Acrotreta lim oensi s WI MAN, 1903; earl y Middle Ca mbr iant") , Sou th Bothni an Sea (Limo n) , Sw ed en. Discussion. - Luh ot reta M ERGL et SLI~ HO FERovA , 1990, from the ea rly M iddl e Ca m brian of Bohemia di ffer s o nly s lig htly from vandalotreta in det ail s o f o rna me nta tio n a nd the co m plete a bse nce of a dorsal medi an rid ge ; the ge nus is re gard ed her e as ajuni or sy no ny m. Vandulot reta is mo st simila r to Hadrotreta ROWELL, 1966, but differ s ma inl y in havin g a boss-shaped apical proc ess th at do es not fill the e ntire apex as in Hadrotreta ; mo reo ver, the do rsal medi an ridge is vesti gial to co m plete ly ab sent in vandalotreta a nd the dorsal va lve lack s a medi an sulcus. vanda lotreta is a lso so me wha t s im ilar to the ceratre tide Bozshakolia HOLMER et USHATINSKAYA, 1994 , in the poor de velopment of a dor sal median ridge and having a fo ra me n that is not e nc losed w ith in the larval she ll, but di ffer s in the lack of a rid ge-like apical process enc los ing the ped icl e tube. Th e poorl y known Acrotre ta lim oen si s WIMAN, 1903, from early Middl e Ca m brian(?) g lac ia l e rrat ics on the Isl and o f Lim on in the South Bothn ian Sea can also be referred to vandalotreta (HOLMER and M ERGL in pr ep ar ati on ). vanda la treta d jagoran (KRUSE, 1990) (PI. 13 : 1-9 ; Te xt-fi g. 4) 1990 . Had ro treta djagoran KRUSE. p. 29 . fig . 5. pl. II A-N .
1991 b. Had ro treta dja goran KRUSE. p. I n. jig. 6H-L. 1993 . Had rotreta primaevu (W ALCOn) : BROCK and COOPER . p.
n2. fig . 14 : 1-1 3.
Holotyp e: Ventral va lve SAM 1'851 38. Type hor izon and local ity: M idd le Camb rian, Tinda ll Li mestone (bo re-ho le NTGS 83/3) , Daly Bas in, Northern Territ ory. Austra lia .
Material. - Fig ured fro m A ntarctica ; complet e she lls: ZPA L Bp .X X XIII/29 S I, 39S8, 54 S6 , 54S 7; ve ntra l va lves: Z PA L Bp .XX XIII/5 4 S5, 6 1S80, 5337U ; dor sal va lves : Z PAL Bp .XX XIII/5 333U. Measu remen ts: See Tab les 5-6. Tab le 5. vando lotreta dja goran (KRlISE). average di mensions of ventra l valves. L
W
N
8
X
0.55
S
T
LIW
8
8
8
8
0.65
0 .26
85 %
48 %
0 .14 8
0.239
0 . 118
1.82
1.61
MI N
0 .30
0 .34
0 . 10
83 %
33 %
MA X
0.80
0 .94
0.46
88%
80 %
-
T/L
Tab le 6. vanda iotreta djagoran (KR USE). average dimensio ns of dorsa l va lves . L
11
W
PI
Iw
Cl
Cw
SI
N
5
5
4
4
4
4
4
4
X
0 .85
0.99
0 .055
0.54
0 .21
0 .23
0 .64
0 .66
S
0.439
0.503
0 .025
0.230
0 .099
0 .079
0 .189 0 .388
MIN
0.30
0 .36
0.02
0 .24
0 .08
0 . 14
M AX
1.30
1.56
0 .08
0 .76
0.32
0 .32
Diagnosis. - See KRUSE ( 1990, p. 29).
20% 32 %
LIW
IwlW
CI/L
C wlW
SUL
5
4
4
4
4
86 %
45%
26 %
56 %
60 %
4.5
6.9
3. 1
6. 1
6.4
0. 12
8 1%
39 %
23 %
49 %
5 2%
0 .98
90 %
65 %
30%
63 %
69 %
48
LARS E. HOLMER, LEONID E. POPOV and RYSZARD WRONA
. umbonal (apical pits)
:~ I w--: cardinal muscle field
11
' --~- t :::: j:--: :~: --
: :
:
i
': ~ -----j------- --
-
~lrQJ):
T--
9 1
-- SI
L
vascuta tateralia ___________________
__ _ _
~_o__,~
A
apical process (muscle platform)
B
-+-_median ridge
c
Fig . 4
Vandalotreta djagoran (KRUSE, 1990), lateral view of the ventral valve exterior (A), reconstruction of ventral (B) and dorsal interior (C) . Location of measurements indicated : W - width of valve, Cw - width of cardinal muscle field , Iw - width of dors al 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 near umbo ; pedicle foramen subcircular, situated immediately posterior to apex , not enclosed within larval shell; 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 narrow propareas. Ventral interior with apical process forming low boss-shaped projection with median depression anterior 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 to low, triangular median buttress; dorsal cardinal muscle fields relatively small, somewhat thickened, extending anteriorly for about 26 % of valve length; dor sal 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 a more transversely suboval outline, and a more well defined dorsal median ridge, as well as less well defined 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.
REFERENCES BENGTSON, S., CONWAY MORRI S, S., COOPER, B.J. , JELL, P.A. and RUNNEG AR, B.N . 1990. Early Cambrian fossils from South Australia. - Association of Australasian Palaeontologists, Memoir 9, 1-364. BIERNAT, G., BIRKENM AJER, K. and POPIEL-B ARCZYK, E. 1985. Tertiary brachiopods from the Moby Dick Group of King George Island (South Shetland Islands , Antarctica). - Studia Geologica Polonica 81, 109-141 . BIRKENMAJER, K. 1992. Cenozoic glacial history of the South Shetland Islands and Northern Antarctic Peninsula. In: J . L6pez-Martinez (ed.) Geologia de la Antartida Occidental. Simposios T 3 , 251-260. III Congreso Geol6gico de Espafia y VIII Congreso Latinoamericano de Geologia. Salamanca, Espafi a, BIRKENMAJER, K. and B UTKIEWICZ, T . 1988. Petrography and provenance of magmatic and metamorphic erratic blocks from Lower Miocene glacio-marine deposits of King George Island (South Shetland Islands, Antarctica). - Studia Geologica
Polonica 95, 23-51 .
CAMBRIAN BRACHIOPODS FROM ANTARCTICA
49
BIRKENMAJ ER, K. GAZDZICKI , A. and WRONA, R. 1983. Cretaceo us and Tertiar y foss ils in glacio-mari ne strata at Ca pe Melville, Antarctica. - Nature 303, 56-59. BIR KENMAJ ER, K. GAZDZICKI , A., PUGACZEWSKA, H. and WRONA, R. 1987. Recycled Cre taceo us belemnit es in Lower Miocene glacio-marine sediments (Ca pe Melville Formation) of King George Island . Antarctica. In : A. Gaidzic ki (cd.) Palaeont ological Results of the Polish Antarc tic Expeditions. Part I. - Palaeon tologia Polon ica 49, 49- 62. BROCK, G.A. and COOPER, B.l . 1993. Shelly fossils from the Early Ca mbrian (Toyo nian) Wirrealpa, Aroo na Cree k, and Ramsay limestones of South Australia. - Journ al of Paleo nto logy 67, 758- 778. CHUANG, S.H. 197 1. Th e morph ology and paleob iology of Trema tis elliptopora Coo per (Inarticulata, Brach iopod a).ln: 1.T. Dutro (cd.) Paleozoic perspectives: a paleo ntolog ical tribute to G. Arthur Cooper. - Sm ithsonian Contribution s to Paleobiology 3, 93- 100. COIlIlOLD, E.S. 1921. The Cambria n hori zons of Co mley (Shropshire) and their Brachiopoda, Pteropoda, Gas tropoda. Geo logical Society of London , Quarterly Jou rnal 76, 325-386. DEIlRENNE, F. and KRUSE, PD . 1986. Shac kleton Limestone archaeo cya ths. - A lcheringa 10, 235- 278. DEBREr-.'NE, F. and KRUSE, P.D. 1987. Shackleton Limestone archaeocyaths. Corrigendum. - A lcheringa 11, 138. DUDzIAK,l. 1984. Cre taceo us calca reous nannoplankto n from glac iomarine dep osits of the Cape Mel ville area , King Geo rge Island (South Shetland Island s, Antarctica). - Studia Geo log ica Polon ica 79, 37-5 1. EVANS , K.R. and ROWEL L, A.1. 1990. Small shelly fossils from Antarctica: an Early Cam brian faun al co nnectio n with Australia. - Jou rnal of Pala eontology 64, 692-700. EVANs, K.R. 1992. Maroce l/a : Antarctic specimens of an enigmatic Ca mbrian animal. - Journ al of Pal aeon tology 66 , 558-562. GAZDZICK I, A. (cd.) 1987. Palaeont ological Result s of the Polish Antarctic Expeditions. Part I. - Palaeon tologia Polon ica 49, 1-1 68. GORJ Ar-.'SKY, V.1. and Po ro v , L.E. (I'ops ucxmi , B.IO. , OOIlOB, JI.E.) 1985. MO P
