https://doi.org/10.24199/j.mmv.1981.42.02 30 September 1981

A NEW SPECIES OF TORNQUISTIA (BRACHIOPODA: ) FROM THE ARTINSKIAN () OF WESTERN AUSTRALIA By N. W. Archbold

Department of Geology. University of Melbourne, Parkville, Victoria, 3052. Abstract The new species Tornquista gregoryi, from the late Baigendzinian (late Artinskian) Wandagee For- mation of the Carnarvon Basin, Western Australia, is described. Internal structures of the species are elucidated by use of the peel technique. The content and evolution of the family Anopliidae is briefly reviewed and revised.

Introduction Furnish (1961), and revised by Dickins (1976) The classification and distribution of the and Waterhouse (1976). chonetacean family Anopliidae and represen- Localities for Tornquistia gregoryi sp. nov. tatives of the family from the Permian sequence are given with the systematic description. of the Carnarvon Basin, Western Australia, Terminology: The terminology applied herein have previously been investigated by the author to the Anopliidae is that used previously by the (Archbold 1980). After that study had gone to author (Archbold 1980). press, Dr. P. Jell of the National Museum of Techniques: Small chonetacean specimens are Victoria passed on to me a collection of frequently preserved with their valves con- chonetid , from the Permian se- joined and hence elucidation of the dorsal inter- quences of Western Australia, that is housed in nal structure is impossible unless thin sections the National Museum collections. The speci- or peels are prepared. The peel technique has mens described below were collected by Dr C. been seldom applied to the Chonetidina. Camp- Teichert in the years 1939 and 1941. The new bell (1953) and Muir-Wood (1962, pp. 112-113) species Tornquistia gregoryi was investigated used peels to assist in the determination of in- with the aid of peels, a technique seldom ap- ternal structures and the present author (op. cit, phed to the Chonetidina as a group, and the 1980) provided serial sections, in the form of value of this technique for the elucidation of peels, to confirm the internal structures of hidden interior structures is shown herein. species of Demonedys and Tornquistia. The Additions and amendments to the inferred peels of Tornquistia gregoryi si^, nov. described phylogeny of the Anopliidae previously herein were prepared by grinding and etching in presented by the author (Archbold 1980, p. the normal manner. Specimens were embedded 182), made necessary by the works of Jing and in clear polyester resin and the block was Hu (1978) and HavHcek and Racheboeuf (1979) ground at set intervals; a distance of 0.1 mm are given below. apart appears most suitable for the determina- Collections: All specimens are housed in the tion of cardinal structures. Etching of the National Museum of Victoria (NMV), Mel- ground surfaces was for three seconds using bourne, Victoria. 10% HCl. Peels were made of the etched sur- Stratigraphy and Age: Tornquistia gregoryi sp. faces on sheet "celluloid" moistened with a

nov. is known only from the Wandagee Forma- solution of "celluloid" dissolved in acetone. The tion of the Carnarvon Basin. The Permian peels were projected on to photographic paper stratigraphy of the Carnarvon Basin has been and outHnes traced accurately. The orientation documented by Condon (1967) and reviewed by of calcite fibres is semi-diagrammatic. Playford et al. (1975); it is not proposed to review the sequence further here. The age of the Wandagee Formation is SYSTEMATIC PALAEONTOLOGY considered to be Late Baigendzinian (Late Ar- Suborder Chonetidina Muir-Wood 1955

tinskian Stage) as indicated by Glenister and Family Anopliidae Muir-Wood 1 962

Memoirs of the National Museum Victoria, *] No. 42. 1981. B 8 N. W. ARCHBOLD

Subfamily Anopliinae Muir-Wood 1962 an area on Wandagee Station on the Minilya Genus Tornquistia Paeckelmann 1930 River in the Carnarvon Basin, was selected by Dr C. Teichert and others, then of the Univer- Type Species: Leptaena (Chonetes) polita Mc- sity of Western Australia, for detailed map- Coy 1852, ping. The mapping was begun in 1939 and con- Diagnosis: Small, smooth anopliids with lateral tinued in 1940 and 1941. The specimens strongly developed accessory septa in septa and described herein were collected during those the dorsal valve. Alveolus distinct. Cardinal years and came from the stratigraphic interval process small, bilobed or weakly quadrilobate named by Teichert the Wandagee Series, internally. Ventral valve strongly convex. Calceolispongia stage. That stratigraphic inter-

Discussion: The geographical and age ranges of val is now known as the Wandagee Formation. the genus have previously been reveiwed by the Specimens collected during those surveys are author (op. cit. 1980) who demonstrated that accurately located stratigraphically although numerous Permian species belonging to smooth geographically they are not so reliably located genera of the Rugosochonetidae had incorrectly for, as Teichert (1949 p. 37) noted, "the type been assigned to Tornquistia (or Paeckelman- section of the Wandagee Series occurs in nia), Tornquistia is readily distinguished from featureless, plain country which is singularly Demonedys Grant (1976) by the absence of a lacking in any kind of geographic features prominent ventral fold. Anoplia Hall and which could be named". Clarke (1892), a genus superficially similar to The localities of the specimens are recorded Tornquistia, possesses a raised, internally as follows. bilobate cardinal process and long, curved, Specimens NMV P60700-NMV P60712 are crenulated socket ridges. Yagonia Roberts (in from Teichert's field locahty WC (16-24)1, Roberts et al. 1976) also possesses an elongate, recorded as "Wandagee Series, zones 16-24, bilobed cardinal process in addition to many northeast side of syncline, north of Minilya pairs of spinose radial ridges between the short River, west of Coolkilya Pool". Specimens lateral septa and the pronounced accessory sep- NMV P60713-NMV P60724 are from Teichert's ta. Yagonia is also distinguished from Torn- field locality WC (21-25)5, recorded as "Coley's quistia by its remarkably large size. locality E9 and station 29 J of the 1939 survey; Wandagee Series, Calceolispongia stage, zones 16-24". Mr H. Coley, of Wandagee Station, Tornquistia gregoryi sp. nov. was an indefatigable fossil collector, who (Plate 3, figs. 1-28; text figs. 1-2.) became familiar with the stratigraphic occur- Holotype: ventral valve, NMV P60709, a from rences of species of Calceolispongia. Teichert's field locality WC(16-24) 1, described Etymology: The species is named in honour of as "Wandagee Series, zones 16-24, northeast Augustus Charles Gregory, pioneer explorer of side of syncline, north of Minilya River, west of the Swan River Colony and one of the first to Coolkilya Pool." Now in Wandagee Forma- investigate the Permian rocks of Western tion. Australia. Material: Four conjoined shells and 21 isolated ventral valves. One specimen was sectioned to Description: Medium size for genus; convexity reveal the internal structures. low in juveniles, higher in adults. Mesial por- Measurements: All specimens were measured tion of ventral valve inflated, at times with a and yielded the following ranges of weak ventral fold posteriorly developed, other measurements in mm. Graphical scatter plots specimens with a higher convex ventral valve. are provided (fig. 1). Greatest width usually at hinge line; occasional Maximum Width Mid-width Thickness Length specimens develop a rounded quadrate outline 11.4-6.7 10.8-6.3 3.8-2.1 8.8-2.1 with mid-width equal to or slightly greater than hinge width. Interareas low. Dorsal valve gently Localities: After a reconnaissance visit in 1938, concave. Exterior of shell smooth with weakly NEW SPECIES OF TORNQUISTIA

10 •• •-• 10 • • • • ••

• • ,

E 5 5

5

_L 5 70 Length Thickness

Fig. 1— Scatter Plots (in mm) of measurements of Torn- quistia gregoryi sp. nov. of r. gregoryi', the first thirty peels being at 0.1 mm distance apart, the subsequent peels at 0.2 developed growth lines. Cardinal spines poorly mm apart. The full set is shown in Fig. 2. The apparently short and well spaced. known, ventral median septum is obvious by peel no. 8 with well developed teeth, Ventral interior and in peels 10 to 12 can be seen to be ar- median septum high delthyrium wide; ventral ticulating with the bifid cardinal process. Peel with slot between posteriorly, articulating 12 indicates the pronounced bifid nature of the process, lower halves of bilobed cardinal interior of the cardinal process. An additional one half valve anteriorly, extending for about weakly developed lateral groove adjacent to the region, length. Valve floor, excluding muscle strong median groove on the internal face of with scattered pustules. the cardinal process in peel 12 may indicate a cardinal Dorsal interior with strongly bilobed tendency for the cardinal process to be weakly Accessory septa process. No median septum. quadrilobate. Weakly developed low lateral anteriorly as gently well developed, extending septa are indicated in peels 17-19 and the ac- for two-thirds valve diverging, distinct ridges cessory septa arise gently in peel 17 and persist short, low. Anterior of length. Lateral septa anteriorly until peel 32, hence being almost 2 except for region be- valve strongly pustulose mm long. The ventral median septum persists septa. tween the accessory until peel 29 and hence is about half the length Observations on the peels of Tornquistia of the valve. The anterior sections of the two gregoryi: Forty-two peels were prepared from valves reveal the thin nature of the shell the parallel serial grinding of a complete shell anteriorly. 10 N. W. ARCHBOLD NEW SPECIES OF TORNQUISTIA 11

Discussion: This distinctive species, charac- Formation to Demonedys. Re-examination of terized by the incipient development of a ven- these specimens indicates that they could just as tral fold posteriorly in many specimens and the readily be immature specimens of T gregoryi high, arched convexity of mature individuals is hence it is now assumed that Demonedys granti readily distinguished from other species of the is restricted to the Cundlego Formation of the genus. Tornquistia magna Archbold (1980 p. Carnarvon Basin until proven otherwise. 186 pi. 25, figs. 5-13) also is characterised by the possession of a highly convex ventral valve with a strongly inflated mesial portion of the valve; PHYLOGENY OF THE ANOPLIIDAE however, no trace of an incipient ventral fold A detailed account of the inferred phylogeny occurs in T. magna unlike T. gregoryi. T. of the Anopliidae has been given by the author gregoryi is prdbably a direct descendant of T. (Archbold 1980) but the works of Jing and Hu magna. Other Western Australian species of (1978) and Havlicek and Racheboeuf (1979) Tornquistia are normally not as strongly con- necessitate amendments to that account. vex as T. gregoryi. True Tornquistia is rela- Caplinoplia Havlicek and Racheboeuf from tively rare in Permian strata (cf. discussions in the Siegenian and Emsian of Czechoslovakia is Archbold 1980 p. 184) and of foreign species a caenanopliinid with strong, long, curved only gibbera T Afanas'yeva (1977) from the lateral and accessory septa and a short, Late or Early Permian (Assel- anteriorly placed median septum, in the dorsal ian) Paren Horizon of the Kolyma-Omolon valve. It is probably a descendent of Eopli- region, need USSR, be compared with T, canoplia Boucot and Harper (1968), a Ludlo- gregoryi. The Siberian species is distinctly con- vian genus with lateral septa and a variably vex; however, no trace of an incipient median developed dorsal median septum, rather than fold is present and the dorsal internal septa are being an off shoot from Plicanoplia Boucot and weakly developed in comparison with the Harper (1968), a Siegenian genus with lateral strongly developed accessory septa of T, and accessory septa but no median septum, in gregoryi. the dorsal valve. The presence of an incipient, posteriorly Pygmochonetes Jing and Hu (1978) a developed, ventral fold in T, gregoryi recalls caenanopliinid from the Artinskian-Kungurian the genus Demonedys Grant (1976). However, of China possesses similar internal structures to that genus is characterised by species small in Costachonetes Waterhouse (1975) but Costa- size and with a pronounced ventral fold and chonetes invariably possesses a variably hence should not be confused with Tornquistia developed ventral sulcus, at least posteriorly

(cf. Grant 1976, Archbold 1980). The ventral (Archbold 1980 pp. 183, 189; ProkoPev 1975 p. median septum of Demonedys is short and high 17, pi. 1, figs 8-14). Pygmochonetes, charac- unlike the relatively long septum of Torn- terised by a strongly inflated ventral valve with quistia, Nevertheless the discovery of a species a highly arched median section, apparently in- of Tornquistia with an incipient ventral fold cludes Chonetella dubia Loczy (1897, p. 67, supports the view of Archbold (1980. p. 183) text fig. 16) from the Late Carboniferous of that Demonedys developed from a Tornquistia China, a species indicated by Archbold (1980, stock. p. 181) to possibly belong to a new genus. Archbold (1980, p. 188) referred several Pygmochonetes and Costachonetes may have poorly preserved specimens from the Wandagee evolved independently from Globosochonetes as indicated by Archbold (1980, p. 183) or Pygnochonetes may have evolved from early Costachonetes. Fig. 2— Tornquistia gregorvi sp. nov. Transverse serial sec- I now include Yagonia Roberts (in et lions of specimen NMV P60706, x5. The larger Roberts numbers refer to the position of the section, in mm, al. 1976) within the Anopliinae. Yagonia was of the shell. Orienta- from the posterior extremity possibly derived from Anoplia, as indicated tion of calcite fibres is shown semi-diagrammatic- on text fig. 3, but this aily. Pseudo-punctae (taleolae) not shown. remains one of the 12 N. W. ARCHBOLD

DORASHAMIAN Glabrichonetina / TATARIAN DJULFIAN

CHHIDRUAN Chonetina KAZANIAN Permochonetes KUNGURIAN

ARTINSKIAN

SAKMARIAN

ASSELIAN

GZHELIAN Costachonetes STEPHANIAN KASIMOVIAN

o WESTPHALIAN MOSCOVIAN cc Anoph'opsis BASHKIRIAN Globosochonetes NAMURIAN

cc< u VISEAN Yagonit Subgfobosochonetes TOURNAISIAN STRUNIAN FAMENNIAN FRASNIAN /^ Corbicularia GIVETIAN EIFELIAN Cyrtoniscus Caplinoplia ^^^ / EMSIAN ^^

SIEGENIAN \Plicanoplia Squamatina

GEOINNIAN

PRIDOLIAN

LUOLOVIAN Eoplicanoplia <

WENLOCKIAN

LLANOOVERIAN NEW SPECIES OF TORNQUISTIA 13

more hypothetical relationships within the BoGER, H. AND Ftebig, H., 1963. Die Arlicuiaten Anopliinae. Brachiopoden des West Deuischen Oberkarbons. Palaeontographwa Abt. A. 122: 111-165, Finally, Squamatina Havlicek and Rache- BoucoT, A. J. AND Harper, C. W., 1968. to boeuf a genus typical (1979, p. 109), of Lower Chonetacea. J. Paleont. 42( 1 ): anopliinid size and shape from the Siegenian of 143-176. Czechoslovakia may be an atypical descendent Brand, P. J., 1970. Scottish Carboniferous chonetids. Bull. geoL Surv. Gt. fir., 31: 89-137. of Shagamella Boucot and Harper (1968). Campbell. K. S. W., 1953. The Fauna of the Permo- Havlicek and Racheboeuf (1979, p. 109) noted Carboniferous Ingelara Beds of Queensland. Pap. the external similarity of Squamatina to Dep. GeoL Univ. Qd.. 4(3): 1-30. Anoplia and that the internal structures of Condon, M. A., 1967. The Geology of the Carnarvon 2. Permian Stratigraphy. Bull. Bur. Squamatina were atypical of the Anopliidae, Basin, W.A. Part Miner. Resour. Geol. Geophys. Ausi., 11: 1-191. with which I agree, nevertheless, Shagamella, DiCKiNS, J. M., 1976. Correlation Chart for the Permian the earliest known member of the Anopliinae, System of AustraUa. Ibid. 156B: 1-26. possesses poorly developed or no accessory sep- Glenister, B. F. and Furnish. W. M., 1961. The Per- mian ammonoids of Austraha. J. Paleont., 35: ta and hence is similar to its chonetid ancestory. 673-736. Squamatina can be interpreted as a member Grant, R. E., 1976. Permian brachiopods from southern interior of the Anopliidae also possessing Thailand. J. Paleont., 50 (supp. Paleont. Soc. Mem. characters of the Chonetidae, the ancestors of 9): 1-269, the Anopliidae. Hall, J. and Clarke. J. M. 1892. An Introduction to the the genera of Palaeozoic Brachiopoda. Other genera and relationships shown in text study of Paleont. New York, 8(1): 1-367. discussed in Archbold with the fig. 3 are (1980) Havlicek, V. and Racheboeuf. P. R., 1979. Chonetacea exception that Chonetes {Chonetina) westphali- (Brachiopodes) du Silurien et du Devonien de Boheme CW5 Boger & Fiebig (1963, pp. 147-148) from the (Tchecoslovaquie). Annal. de Paleont. (Invert.), 65(2): 69-138. Westphalian of Germany is now placed in JiNG. Y. AND Hu. S., 1978. Brachiopods of the Kuhfeng Tornquistia following Winkler Prins (1970, p. Formation in South Anhui and Nanking Hills. Acta 3) and Brand (1970, p. 100). This, in turn, Palaeont. Sinica, 17(2): 101-127. leaves the origin of Chonetina open to ques- LoczY, L., 1897. A fosszilis emlos — es puhatestii allat- palaeontologiai — tion. maradvanyok leirasa es a stratigraphiai eredmenyek. In, Szechenyi, B. (Grof)., Keletazsiai Utjanak, Tudomdnyos Eredmenye Acknowledgements 1877-1880. Kot. 3: I-VII. 1-192. Budapest. Kilian Frigyes Egyetemi. I thank Dr P. Jell, National Museum of Vic- McCoy, F., 1852. Contributions to British Palaeontology. toria, for the loan of specimens in his care. Dr On some new Brachiopoda from the Carboniferous 421-429. G. A. Thomas, University of Melbourne, Limestone. Ann. Mag. nat. Hist. Ser. 2. 10: MuiR-WooD, H. M., 1%2. On the morphology and critically read the manuscript. Mrs G. Water- classification of the suborder Chonetoidea. man typed the manuscript. The assistance of British Museum (Natural History), pp. 1-132. the staff of the Baillieu Library is ack- Paeckelmann, W., 1930. Die Brachiopoden des deutschen nowledged. Unterkarbons. I Teil: Die Orthiden, Strophomeniden und Choneten des Mittleren und Oberen Unterkar- bons. Abh. Preuss. Geol. Landesanst. N.f. 122: 143-326. References Playford, p. E., Cope, R. M., Cockbain, A. E., Low, Afanas'yeva, G. a.. 1977. Suborder Chonetidina. In, G. H., AND LowRY, D. C, 1975. Phanerozoic. In, Sarycheva, T. G., (Ed.), Pozdnepaleozoiskie Pro- Geology of Western Australia. West. Australia GeoL Inst. diktidy Sibiri i Arkliki. Trudy Paleont. Akad. Surv. Mem.. 2: 233-432. NaukSSSR, 161-5-41. Prokof'ev, V. A., 1975. Brakhiopody verkhnego Kar- Archbold, N. W., 1980. Studies on Western Australian bona Samarskoi Luki. Trudy Vses. nauchno — issled. Permian Brachiopods. I. The Family Anopliidae Geol.-razved. neft. Inst. (VNIGNI), 162: 1-144. (Chonetidina). Proc. Roy. Soc. Vict., 91(2): 181-192. Roberts, J., Hunt. J. W., and Thompson, D. M. 1976. Late Carboniferous marine invertebrate zones of eastern Australia. Alcheringa 1: 197-225. Teichert, C. 1949. Permian crinoid Calceolispongia, 3 - Inferred phylogeny of the Family Anopliidae Muir- Fig Mem. geol. Soc. Amer., 34: 1-132, Wood (modified from Archbold 1980, p. 182). 14 N. W.ARCHBOLD

Dorsal and ventral views of Waterhouse, J. B., 1975. New Permian and Figs. 7-8. NMV P60713. ventral valve. brachiopod taxa. Pap. Dep. Geol. Univ. Qd. 7(1): Figs. 9-10. P60718. Ventral and posterior views of 1-23. NMV ventral valve. Waterhouse, J. B., 1976. World correlations for Permian Figs. 11. NMV P60724. Ventral view of ventral valve. marine faunas. Pap. Dep. Geol. Univ. Qd. 1(1): 1-232. Figs. 12. NMV P60707. Ventral view of ventral valve. Winkler-Prins, C. F., 1970. The brachiopod faunas of Figs. 13. NMV P60716. Ventral view of ventral valve. the Aegir Marine Band and of the British, Belgian and Figs. 14-17. NMV P60706. Posterior dorsal, anterior and Dutch marine bands of the same age. Rapport. Geol. ventral views of shell. Holotype. Posterior, dorsal Bureau Heerlen, 1153: 1-8. Figs. 18-20. NMV P60709. and ventral views of ventral valve. views of Explanation of Plate Figs. 21-22. NMV P60717. Ventral and posterior ventral valve. PLATE 3 Figs. 23. NMV P60708. Ventral view of ventral valve. Tornquistia gregoryi sp. nov. All figures x4.5. Figs. 24. NMV P60705. Ventral view of ventral valve. Figs. 1-3. NMV P60715. Dorsal, ventral and posterior Figs. 25. NMV P60702. Ventral view of ventral valve. views of shell. Figs. 26-27. NMV P60710. Ventral and posterior views of Figs. 4-6. NMV P60712. Dorsal, ventral and anterior ventral valve. views of shell. Figs. 28. NMV P60700. Ventral valve in ventral view. MEM. NAT. MUS. VICT. 42 PLATE 3