Early Devonian) Microvertebrates from the Buchan and Taemas Areas of Southeastern Australia

Records of the Western Australian MliSI'll III Supplement No. 57: 15-21 (1999).

Emsian (Early Devonian) microvertebrates from the Buchan and Taemas areas of southeastern Australia

Alison Basden Centre for Ecostratigraphy and Palaeobiology, School of Earth Sciences, Macquarie University, NSW 2109; email: [email protected]

Abstract - Microvertebrates obtained from measured stratigraphic sections near Buchan in northeastern Victoria and Taemas in southern New South Wales have been dated as dehiscens and perbonus conodont zones (early Emsian). The two microvertebrate faunas are similar; the areas are separated by only about 300 km and were both shallow marine environments during the Early Devonian. Thelodonts are a rare component in the otherwise abundant microvertebrate fauna recovered from four sections containing almost 450 samples; 18 turiniid scales have been found. It is considered likely that only one species of turiniid is represented. Trunk and transitional scales share similarites with scales of at least five species of turiniids from eastern Gondwana, recovered from the Aztec Siltstone in Antarctica, the Cravens Peak Beds in western Queensland, the Officer Basin in South Australia and the Silverband Formation in western Victoria. Some head scales show a previously undescribed crown morphology of sharply incised, parallel-sided notches separating wide radiating ribs. The scales are referred to Turinia sp. cf. T. allstraliensis.

GEOLOGICAL SETTING AND PREVIOUS WORK the lungfish Dipnorhynchus (Hills 1936b). The only recent description is by Long (1984), who assigned Buchan Group various disarticulated plates from the McLarty The Buchan Group in northeastern Victoria Member of the Murrindal Limestone to comprises three formations: the Buchan Caves Buc1wnosteus confertituberculatus, and described two Limestone (dehiscens Zone), overlain by the new acanthothoracids: Murrindalaspis wallacei and Taravale Formation (dehiscens serotinus zones) and M. bairdi. Also identified were several genera first the Murrindal Limestone (perbonus Zone). An described from the Taemas/Wee Jasper fauna: the outline of the geology of the Buchan area is given brachythoracids Arenipiscis westolli, Errolosteus sp. by Mawson (1987). The conodont biostratigraphy cf. E. goodradigbeensis, and Taemasosteus is documented by Mawson (1987) and Mawson et maclartiensis, and the petalichthyid Widjeaspis al. (1988, 1992). The Buchan Caves Limestone warrooensis. Turner (1991, 1993) discussed elements contains abundant stromatoporoids and corals of the Buchan microvertebrate fauna; faunallists are (Webby et al. 1993), associated with brachiopods, given in Young (1993) and De Pomeroy (1995). bivalves and ostracods (Talent 1995). The Taravale Microvertebrates from Buchan have not previously Formation contains more diverse faunas of been illustrated, except for the scales of brachiopods, corals, and trilobites (Teichert and Dipnorhyncillls figured by Thomson and Campbell Talent 1958), the oldest known ammonoids (1971, figures 86-88). (Mawson 1987), dacryoconarids, ostracods, agglutinated foraminiferans, scolecodonts and 55 species of chitinozoans (Winchester-Seeto 1996). Murrumbidgee Group Young (1979) discussed early discoveries of fossil The Murrumbidgee Group in southeastern New fish from the Buchan area and the similar sequence South Wales is of similar age and lithology to the from the Taemas/Wee Jasper region of NSW (see Buchan Group (pireneae - serotinus zones), and below). Placoderms are the most common element occurs in two separate outcrops on the shores of in the macrovertebrate fauna, and brachythoracid Burrinjuck Dam. The stratigraphy of the eastern arthrodires are the most common placoderms. The outcrop (Taemas area), as first outlined by Browne arthrodire skull first described by Chaprnan (1916), (1959), comprises, from oldest to youngest, the prepared by Hills (1936a) and eventually assigned Cavan, Majurgong and Taernas Formations. The to Buchanosteus confertituberculatus by Stensib (1945) geology of the western outcrop, in the valley of the came from the Buchan area, as did a mandible of Goodradigbee River at Wee Jasper, was outlined by 16 A. Basden

Pedder et al. (1970). The same sequence is evident in recorded from the Burrinjuck limestone sequence, the lower part of the Wee Jasper section, but the are from stratigraphic section YWJ (224 m), which upper part contains more extensive and massive spans the pireneae and dehiscens zones. It passes limestone horizons up to a total of approximately through the Cavan and Majurgong Fonnations along 450 m thick, and is overlain by the terrigenous the road on the northern side of Taemas bridge, the Hatchery Creek Fonnation. Mawson et al. (1992) type locality for the index conodont Polygnathus identified the Pragian-Emsian boundary within the dehiscens (Philip and Jackson 1967). The scales were Cavan formation at the base of the Wee Jasper recovered from two of the 125 sampled horizons: succession. YWJ1l7.5, uppermost Cavan Formation, and The marine limestones in the Burrinjuck Dam area YWJ135.4, basal Majurgong Fonnation, both dehiscens contain an abundant invertebrate fauna (e.g. Zone. The single scale recovered from the higher of Johnston 1993). The diverse fish fauna is these two samples, YWJ135.4, appears abraded and dominated by placoderms (White 1952, 1978; possibly reworked. Microvertebrate remains occur White and Toombs 1972; Young 1979, 1980, 1981, throughout the section, but are especially abundant 1985; Long and Young 1988; Findlay 1996), but also around the Cavan-Majurgong boundary. The includes lungfishes (e.g. Campbell and Barwick depositional environment of the Cavan Fonnation is 1982-85), other osteichthyans (e.g. 5chultze 1968; interpreted as nearshore intertidal; higher horizons, 0rvig 1969; Giffin 1980), and acanthodians (Long comprising shales with limestone lenses, contain a 1986). Chondrichthyans are represented only by fauna of brachiopods, bivalves and gastropods microremains (e.g. Giffin 1980). Turner (1991, 1993) decreasing in abundance towards the boundary with discussed the microvertebrate faunas. The first the Majurgong Formation (Young 1969). The thelodont scales from the Burrinjuck area are a Majurgong Fonnation, consisting of siltstones, fine result of the present study; the fonn "Skamolepis", sandstones and shales with some thin limestone previously assigned to this group by Giffin (1980), lenses, is fairly unfossiliferous, although spiriferids is now considered to be a shark scale, and is not and rare lingulids occur in the lower horizons the same as Skamolepis described by Karatajute (Young 1969), suggesting brackish conditions during Talimaa (1978), and considered to be a thelodont deposition. There is evidence from other areas that (e.g. Turner 1991, 1993). The vertebrate fauna is thelodonts preferred very shallow water or non apparently similar to, or the same as, that from the marine environments (Marss and Einasto 1978; Buchan area (see faunallists in Young 1993 and De Turner 1997). Pomeroy 1995). Although some of the scale morphologies are previously undescribed, it is assumed for the RESULTS present that only one species of turiniid thelodont Residues of the seven stratigraphic sections is represented, since, although possible, it is sampled for conodonts near Buchan (see Mawson unlikely that such a small number of specimens (18 1987) were investigated for microvertebrates. scales from almost 450 samples) would have Only three sections - 510combe's (5L/5LO; 42 originated from several species. samples), McLarty's (120 samples) and Gelantipy Road (80 samples) contained significant SYSTEMATIC PALAEONTOLOGY microvertebrate faunas, expecially placoderms Figured material is lodged in the palaeontological and acanthodians, while all seven thelodont scales collections of the Australian Museum (prefix AM F). from Buchan were recovered from six samples in Locality information is given by section code section 5L/5LO. followed by metres above base of section of lowest The 11 thelodont scales from Taemas, the first and highest productive sample.

Figure 1 Thelodont scales TlIrinia sp. cf. T. allstraliensis from Buchan and Taemas. A-C, scale AM F101174 from .. SL051-68m, Taravale Formation, perbonlls Zone. A, crown view; B, base view; C, lateral view. D-F, scale AM F101203 from YW]117.5, Cavan Formation, dehiscens Zone. D, crown view; E, base view; F, lateral view. G-I, scale AM FlO1175 from SL0213m, Taravale Formation, perbonlls Zone. G, crown view; H, base view; I, lateral view. J-L, scale AM F103583 from SL46.5m, Buchan Caves Limestone, dehiscens Zone. J, crown view; K, base view; L, lateral view. M-N, scale AM FlO1176 from SL48.7m, Buchan Caves Limestone, dehiscens Zone. M, crown view; N, base view. 0, scale AM F103584 from SL028.5m, Taravale Formation, dehiscens Zone. Crown view. P-Q, scale AM F101204 from YW]117.5, Cavan Formation, dehiscens Zone. P, crown view; Q, lateral view. R-S, scale AM F103585 from YW]117.5, Cavan Formation, dehiscens Zone. R, crown view; S, lateral view. T-V, scale AM FlO1177 from SL051m, Taravale Formation, dehiscens Zone. T, crown view; U, base view; V, detail of crown ornament. W, scale AM F101205 from YW]117.5, Cavan Formation, dehiscens Zone. Crown view. X, scale AM F103586 from SL46.5m, Buchan Caves Limestone, dehiscens Zone. Crown view (specimen broken and crown misoriented by 180°). Scale bars = 100 Jlm. 17 Emsian microvertebrates from Buchan and Taemas 18 A. Basden

Superclass Agnatha Haeckel, 1895 The base is deeply convex and contains one small central pulp cavity and a smaller cavity towards Subclass Thelodonti Kiaer, 1924 one side (Figure lK). Order Thelodontida Stensio, 1958 Transitional Scales Family Turiniidae Obruchev, 1964 Crown is elliptical (Figure 10) or diamond shaped (Figure lM). Deep troughs aligned from the Genus Turinia Traquair, 1896 outer anterior margin posteriorly towards the centre of the crown produce coarse ribs covering virtually Turinia sp. cf. T. australiensis Gross, 1971 the entire crown surface. In scale AM FI03584 Material Examined (Figure 10) these ribs are separated by a narrow Eighteen scales: lO head scales, four transitional central ridge running from anterior to posterior; (cephalopectoral) scales, four trunk (postpectoral) the ribs alternating in their anterolateral direction scales. from this central ridge to the lateral margin of the crown. A central ridge is less pronounced in scale Localities AM FlO1l76 (Figure IM), where the ridges and Slocombe's section at Buchan, samples SL 46.5m, troughs are all aligned from the anterior and lateral SL 48.7m (Buchan Caves Limestone), SLO 28.5m, margins towards the posterior of the crown. SLO 5lm, SLO 5l-68m, SLO 213m (Taravale Troughs on the crown of scales AM FlOI204 (Figure Formation); Section YWJ at Taemas, samples IP) and AM F103585 (Figure lR) terminate closer to 117.5m (Cavan Formation) and 135.4m (Majurgong the scale margin, leaving a smooth, flat central area Formation). on the crown. The base is the same shape as the crown, but Stratigraphic Range projects forward from the crown, while the crown Early Devonian, Emsian: dehiscens - perbonus overhangs the base posteriorly. The base contains a conodont zones at Buchan; dehiscens Zone at central pulp cavity surrounded by a rounded rim. Taemas. The crown joins directly to the base without a constricted neck, although in scales AM FlOl204 Description (Figure IP) and AM FI03585 (Figure IR) there is a shallow trough around the base of the crown, Head Scales similar to that found on some body scales, e.g. The head scales (Figure lA-L) are circular or Figure IT. subcircular. The central part of the crown is unornamented and, in the elongated scales (Figure Trunk Scales lD-L), flat. Approximately 14 main ribs around the There are two distinct morphological types of rim of the crown are separated by deep troughs. In trunk scales. The more commonly occurring type is scale AM FlO1174 (Figure lA-e), the troughs are illustrated in Figure IT-W. Scale AM FlO1l77 deeply incised with parallel sides, and on parts of (Figure IT-V) is elongated horizontally with the the margin long and short troughs alternate base extending as a long anterior process producing a bifurcation of some of the ribs. The approximately two-thirds the total length of the troughs in scales AM FlO1175 (Figure lG-I) and AM scale. The complex crown structure comprises FI03583 (Figure IJ-L) become narrower towards the several overlapping, deeply subdivided layers centre of the crown and terminate in shallow surrounding a central lanceolate platform. This elongated depressions. The ribs are subdivided by central platform bears ridges and troughs narrow vertical grooves; in scale AM FlO1174 (Figure sUbparallel to the margin. The central platform is IA-e) these occur only on the sides of the scale and successively underlain towards the posterior of the give the outline a serrated appearance, in scales AM crown by three similar, larger, ridged layers. The FlO1175 (Figure lG-I) and AM F103583 (Figure lJ posterior part of the crown has been broken; L) the grooves extend a short distance onto the overall crown shape is difficult to determine. The crown, while in scale AM FlOl203 (Figure ID-F) the posterior part of the crown substantially overhangs vertical grooves on the distal ends of the ribs are not the base (Figure 1U), with 10 parallel ribs extending well developed (see Figure IF). from the posterior crown-base interface to the In scales AM FlO1174 (Figure lA-e), AM FlOI203 posterior edge of the crown (Figure 1U). The (Figure ID-F) and AM FlO1175 (Figure lG-I) the anterior neck region is a wide, shallow trough. The crown and base are separated by a constricted neck. base is shallow and flat with a small central pUlp The base is shallow and comprises a low rounded cavity, and the tapering anterior process extends rim surrounding a central large, deep pulp cavity. smoothly from the base in the same horizontal In scale AM FlO3583 (Figure IJ-L) the neck is not plane. Other scales of this type, e.g. Figure 1W, have constricted and is not clearly defined (Figure lL). crown ornament of similar overlapping layers with Emsian microvertebrates from Buchan and Taemas 19 less deeply incised troughs, and a shorter anterior the Australian National University from shot point process on the base. and outcrop samples from the Cravens Peak Beds The second scale type is represented by only one (T. australiensis, sample SP813, and T. gavinyollllgi, specimen (broken during electron micrography; sample GB77, respectively), and from the Aztec Figure IX). The flat diamond-shaped crown has a Siltstone (T. antarctica, sample V2616). central, diamond-shaped, slightly raised portion The crown of transitional scale AM F101176 with three wide, shallow anterior ridges separated (Figure IM-N) also resembles that of T. allstraliensis by wide, shallow troughs. This central area is (e.g. Turner et a1. 1981, figure 111), with crenulations surrounded laterally and posteriorly by a larger, deeply incising the crown almost to the centre, flat, diamond-shaped area with the lateral corners leaving only a narrow central smooth area (Figure slightly stepped down, and the posterior corner IM). Scales of T. gavinyOlmgi figured by Turner et £11. extending slightly into a rounded point. On either (1981, figure 7A) have a similarly crenulated crown, side of this posterior point there is a lower, narrow, but the crenulations here are more widely spaced elliptical platform. There is no dearly-defined neck and the central smooth ridge is wider than in the area. The base is extended into a short, stout, Buchan specimen. Subparallel troughs extending tapering anterior process approximately one-third from the margin almost to the central antero the total length of the scale. posterior axis of the crown are also found in scales of TlIrinia d. T. pagoda from the eastern Officer Basin, South Australia, figured by Long et £11. (1988, DISCUSSION figure 2A,B). Although the number and orientation Some Buchan/Taemas thelodont scales share of the troughs is similar to those on the crown of features with those of Turinia australiensis from Buchan specimen AM FI01176 (Figure 1M-N), the western Queensland (Turner et £11. 1981) and from crown of the South Australian scale is more the Jerula Formation of central NSW (pesavis shallowly incised. The base, too, of the Buchan scale sulcatus zones) (Turner 1997, figures 3C, 3K, 3N, 4G, (Figure IN), which is shallow with a large central 41, 4M); T. antarctica from southern Victoria Land, pulp cavity and small anterior process, resembles Antarctica (Turner and Young 1992); T. gondwana that of body scales of Turinia sp. cf. T. pagoda (Long from Bolivia (Gagnier et £11. 1988); T. ftlscina from et £11. 1988, figure 2A-C,E). However, although the western Victoria (Turner 1986), and scales from Buchan specimen shares these features with the western Queensland described as Turinia TlIrinia sp. cf. T. pagoda material from South gavinyOlmgi by Turner (1995a) and from South Australia, it differs from the original description of Australia described as Turinia d. T. pagoda by Long T. pagoda from western Yunnan, China (Wang et £11. et £11. (1988). Scales from the Cravens Peak Beds of 1986) in lacking the lateral neck-spurs and by western Queensland (Eifelian), and the Aztec having more deeply incised grooves on the crown Siltstone in Antarctica (Givetian), held in the (see Wang et £11. 1986, figure 4G-J). A specimen of T. collections of the Queensland Museum and the !lIscina, figured by Turner (1986, figure 2N), Australian National University, also show resembles the transitional scales from Buchan and similarities with the Buchan/Taemas specimens. Taemas in crown ornament. Crown ornamentation of distinct, parallel-sided The crown of trunk scale AM FlO1177 (Figure troughs between sometimes bifurcating ribs on 1T,V) resembles scales of T. australiensis from the head scale AM F101203 (Figure 10) is seen in head Jerula Limestone Member of the Glenmga Formation scales of T. australiensis (Turner et £11. 1981, figure in western New South Wales (Turner 1997, figure llA). The Buchan specimen lacks the deep rounded 4M), T. antarctica, described from the Aztec Siltstone base of typical head scales of T. australiensis (Turner of southern Victoria Land, Antarctica by Turner and et al. 1981), although the depth of the base in Young (1992, figure 5H), and T. gondwana from the thelodont scales is related to ontogenetic stage (S. Catavi Formation, Bolivia (Gagnier et £11. 1988, figure Turner pers. comm. 1995). Similar crown ornament 40,E). Common features are the central raised occurs in specimens from samples C654 from the elliptical section with subparallel ribs, lateral Jerula Formation at Condobolin (see also Turner segments that converge with the posterior tip of the 1997, figure 3F) and C657 from the Trundle Group central platform in a thin ridge extending towards at Kadungle, held in the collections of the the posterior of the crown (e.g. Turner and Young Queensland Museum. 1992, figure 5h; Gagnier et al. 1988, figure 40,E), The second type of head scale morphology, with parallel ribs on the ventral side of the crown (Turner tapering troughs terminating in shallow and Young 1992, figure 6; Gagnier et al. 1988, figure depressions (Figure IG-L), can be found in 4), a smooth wide trough around the anterior edge of specimens from the Officer Basin, South Australia the crown (Turner and Young 1992, figure 5a, c-m), (Turinia sp. aff. T. australiensis - Long et £11. 1988, and a long anterior process on the base (Turner and figure 5A), Cravens Peak Beds (T. australiensis Young, 1992, figure 5d-€, g-m; Gagnier et al. 1988, Turner et a1. 1981, figure 100), and in collections at figure 5B). The same pattern of tiered subdivisions of 20 A. Basden the crown occurs in body scales from the Early Speonesydrion, an Early Devonian dipnoan with Lochkovian Read Bay Formation of Cornwallis primitive toothplates. Palaeoichthyologica 2: 1-48. Island, Canadian Arctic (Turner and Burrow 1997), Campbell, K.S.W. and Barwick, RE. (1985). An advanced and in specimens from outcrop locality GB77 of the massive dipnorhynchid lungfish from the Early Cravens Peak Beds in western Queensland (Turner Devonian of New South Wales, Australia. Records of 1995a, figure 3V,W; Queensland Museum collection). the Australian Museum 37: 301-316. Chapman, F. (1916). On the generic position of 'Asterolepis ornata var.' McCoy, with description of a new variety. Proceedings CONCLUSION of the Royal Society of Victoria 28: 211-215. The turiniid scales described in this study show De Pomeroy, A. (1995). Australian Devonian fish most morphological similarity to scales of T. biostratigraphy in relation to conodont zonation. australiensis (see Turner 1995b, 1997 for discussion of Courier Forschungsinstitut Senckenberg 182: 475-486. T. australiensis scale morphology, histology and Findlay, CS. (1996). Placoderm (Pisces: Placodermi) significance of the taxon in biostratigraphic remains from Lower Devonian rocks at Taemas, New subdivision of the Early Devonian). However, South Wales. Proceedings of the Linnean Society of New South Wales 116: 161-186. because of slight differences from previously figured Gagnier, P.-Y., Turner, S., Friman, specimens of T. australiensis, similarities with other L., Suarez-Riglos, M. and ]anvier, P. (1988). The Devonian species of Tu rinia, and the unique morphology vertebrate and of mollusc fauna from Seripona (Dept. scale AM FlO1l74 (Figure 1A-e), of Chuquisaca, the turiniid scales Bolivia). Neues Jahrbuch fiir Geologie from the Buchan/Taemas lmd Paliiontologie, fauna described herein are Abhandlungen 176: 269-297. referred to Tuninia sp. cf. T. australiensis. The age of Giffin, E. (1980). Devonian vertebrates from Australia. this fauna (perbonus - dehiscens zones) is the same as Postilla 80: 1-15. that of the T. sp. cf. T. australiensis Assemblage HiIls, E.s. (1936a). On certain endocranial structures in (Fauna 3) of Turner (1997). Coccosteus. Geological Magazine 73: 213-226. Hills, E.s. (1936b). Records and descriptions of some Australian Devonian fishes. Proceedings of the Royal ACKNOWLEDGEMENTS Society ofVictoria 48: 161-171. ]ohnston, P.A (1993). Lower Devonian Pelecypoda I thank John Talent and Ruth Mawson from for southeastern Australia. Memoirs of the Association providing the material for of this study and giving Australasian Palaeontologists 14: 1-134. help and encouragement throughout the project; Karatajute-Talimaa, V.N. (1978): Telodonti Silura i Devona Sue Turner for assistance with identification, access SSSR i Shpitsbergena [Silurian and Devonian thelodonts to collections and financial help to visit Brisbane to of the USSR and Spitsbergen]: 1-344, Mokslas, Vilnius, study material; Gavin Young for continued help Lithuania. and encouragement, and for providing access to Long, ].A (1984). New placoderm fishes from the Early AGSO and AND collections; the Linnean Society of Devonian Buchan Group, eastern Victoria. Proceedings ofthe Royal NSW and the SA and ACT branches of the Society ofVictoria 96: 173-186. Long, ].A (1986). New Australian Federation of University Women for ischnacanthid acanthodians from the Early Devonian of Australia, financial assistance; David Mathieson for help with with comments on acanthodian interrelationships. printing; and Theresa Winchester-Seeto Zoological Journal ofthe for Linnean Society ofLondon 87: 321-339. ongoing practical advice and encouragement. Part Long, ].A., Turner, S. and Young, G.C (1988). A of this study was carried out during tenure of an Devonian fish fauna from subsurface sediments in the Australian Postgraduate Award. eastern Officer Basin, South Australia. Alcheringa 12: 61-78. Long, ].A and Young, G.C (1988). Acanthothoracid REFERENCES remains from the Early Devonian of New South Wales, including a complete sclerotic capsule and Browne, I. (1959). Stratigraphy and structures of the pelvic girdle. Memoirs ofthe Association of Devonian rocks of the Taemas and Cavan Australasian areas, Palaeontologists 7: 65-80. 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