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

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

Alison Basden

Centre for Ecostratigraphy and Palaeobiology, School of Earth Sciences, Macquarie University, NSW 2109;email: abasden@laurel.ocs.mq.edu.au

Abstract - Microvertebrates obtained from measured stratigraphic sectionsnear Buchan in northeastern Victoria and Taemas in southern New SouthWales have been dated as dehiscens and perbonus conodont zones (earlyEmsian). The two microvertebrate faunas are similar; the areas are separatedby only about 300 km and were both shallow marine environments duringthe Early Devonian. Thelodonts are a rare component in the otherwiseabundant microvertebrate fauna recovered from four sections containingalmost 450 samples; 18 turiniid scales have been found. It is consideredlikely that only one species of turiniid is represented. Trunk and transitionalscales share similarites with scales of at least five species of turiniids fromeastern Gondwana, recovered from the Aztec Siltstone in Antarctica, theCravens Peak Beds in western Queensland, the Officer Basin in SouthAustralia and the Silverband Formation in western Victoria. Some head scalesshow a previously undescribed crown morphology of sharply incised,parallel-sided notches separating wide radiating ribs. The scales are referredto Turinia sp. cf. T. allstraliensis.

GEOLOGICAL SETTING AND PREVIOUS WORK

Buchan GroupThe Buchan Group in northeastern Victoria

comprises three formations: the Buchan CavesLimestone (dehiscens Zone), overlain by theTaravale Formation (dehiscens serotinus zones) andthe Murrindal Limestone (perbonus Zone). Anoutline of the geology of the Buchan area is givenby Mawson (1987). The conodont biostratigraphyis documented by Mawson (1987) and Mawson etal. (1988, 1992). The Buchan Caves Limestonecontains abundant stromatoporoids and corals(Webby et al. 1993), associated with brachiopods,bivalves and ostracods (Talent 1995). The TaravaleFormation contains more diverse faunas ofbrachiopods, corals, and trilobites (Teichert andTalent 1958), the oldest known ammonoids(Mawson 1987), dacryoconarids, ostracods,agglutinated foraminiferans, scolecodonts and 55species of chitinozoans (Winchester-Seeto 1996).

Young (1979) discussed early discoveries of fossilfish from the Buchan area and the similar sequencefrom the Taemas/Wee Jasper region of NSW (seebelow). Placoderms are the most common elementin the macrovertebrate fauna, and brachythoracidarthrodires are the most common placoderms. Thearthrodire skull first described by Chaprnan (1916),prepared by Hills (1936a) and eventually assignedto Buchanosteus confertituberculatus by Stensib (1945)came from the Buchan area, as did a mandible of

the lungfish Dipnorhynchus (Hills 1936b). The onlyrecent description is by Long (1984), who assignedvarious disarticulated plates from the McLartyMember of the Murrindal Limestone toBuc1wnosteus confertituberculatus, and described twonew acanthothoracids: Murrindalaspis wallacei andM. bairdi. Also identified were several genera firstdescribed from the Taemas/Wee Jasper fauna: thebrachythoracids Arenipiscis westolli, Errolosteus sp.cf. E. goodradigbeensis, and Taemasosteusmaclartiensis, and the petalichthyid Widjeaspiswarrooensis. Turner (1991, 1993) discussed elementsof the Buchan microvertebrate fauna; faunallists aregiven in Young (1993) and De Pomeroy (1995).Microvertebrates from Buchan have not previouslybeen illustrated, except for the scales ofDipnorhyncillls figured by Thomson and Campbell(1971, figures 86-88).

Murrumbidgee GroupThe Murrumbidgee Group in southeastern New

South Wales is of similar age and lithology to theBuchan Group (pireneae - serotinus zones), andoccurs in two separate outcrops on the shores ofBurrinjuck Dam. The stratigraphy of the easternoutcrop (Taemas area), as first outlined by Browne(1959), comprises, from oldest to youngest, theCavan, Majurgong and Taernas Formations. Thegeology of the western outcrop, in the valley of theGoodradigbee River at Wee Jasper, was outlined by

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Pedder et al. (1970). The same sequence is evident inthe lower part of the Wee Jasper section, but theupper part contains more extensive and massivelimestone horizons up to a total of approximately450 m thick, and is overlain by the terrigenousHatchery Creek Fonnation. Mawson et al. (1992)identified the Pragian-Emsian boundary within theCavan formation at the base of the Wee Jaspersuccession.

The marine limestones in the Burrinjuck Dam areacontain an abundant invertebrate fauna (e.g.Johnston 1993). The diverse fish fauna isdominated by placoderms (White 1952, 1978;White and Toombs 1972; Young 1979, 1980, 1981,1985; Long and Young 1988; Findlay 1996), but alsoincludes lungfishes (e.g. Campbell and Barwick1982-85), other osteichthyans (e.g. 5chultze 1968;0rvig 1969; Giffin 1980), and acanthodians (Long1986). Chondrichthyans are represented only bymicroremains (e.g. Giffin 1980). Turner (1991, 1993)discussed the microvertebrate faunas. The firstthelodont scales from the Burrinjuck area are aresult of the present study; the fonn "Skamolepis",previously assigned to this group by Giffin (1980),is now considered to be a shark scale, and is notthe same as Skamolepis described by Karatajute­Talimaa (1978), and considered to be a thelodont(e.g. Turner 1991, 1993). The vertebrate fauna isapparently similar to, or the same as, that from theBuchan area (see faunallists in Young 1993 and DePomeroy 1995).

RESULTSResidues of the seven stratigraphic sections

sampled for conodonts near Buchan (see Mawson1987) were investigated for microvertebrates.Only three sections - 510combe's (5L/5LO; 42samples), McLarty's (120 samples) and GelantipyRoad (80 samples) contained significantmicrovertebrate faunas, expecially placodermsand acanthodians, while all seven thelodont scalesfrom Buchan were recovered from six samples insection 5L/5LO.

The 11 thelodont scales from Taemas, the first

A. Basden

recorded from the Burrinjuck limestone sequence,are from stratigraphic section YWJ (224 m), whichspans the pireneae and dehiscens zones. It passesthrough the Cavan and Majurgong Fonnations alongthe road on the northern side of Taemas bridge, thetype locality for the index conodont Polygnathusdehiscens (Philip and Jackson 1967). The scales wererecovered from two of the 125 sampled horizons:YWJ1l7.5, uppermost Cavan Formation, andYWJ135.4, basal Majurgong Fonnation, both dehiscensZone. The single scale recovered from the higher ofthese two samples, YWJ135.4, appears abraded andpossibly reworked. Microvertebrate remains occurthroughout the section, but are especially abundantaround the Cavan-Majurgong boundary. Thedepositional environment of the Cavan Fonnation isinterpreted as nearshore intertidal; higher horizons,comprising shales with limestone lenses, contain afauna of brachiopods, bivalves and gastropodsdecreasing in abundance towards the boundary withthe Majurgong Formation (Young 1969). TheMajurgong Fonnation, consisting of siltstones, finesandstones and shales with some thin limestonelenses, is fairly unfossiliferous, although spiriferidsand rare lingulids occur in the lower horizons(Young 1969), suggesting brackish conditions duringdeposition. There is evidence from other areas thatthelodonts preferred very shallow water or non­marine environments (Marss and Einasto 1978;Turner 1997).

Although some of the scale morphologies arepreviously undescribed, it is assumed for thepresent that only one species of turiniid thelodontis represented, since, although possible, it isunlikely that such a small number of specimens (18scales from almost 450 samples) would haveoriginated from several species.

SYSTEMATIC PALAEONTOLOGYFigured material is lodged in the palaeontological

collections of the Australian Museum (prefix AM F).Locality information is given by section codefollowed by metres above base of section of lowestand 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, scaleAM 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, dehiscensZone. M, crown view; N, base view. 0, scale AM F103584 from SL028.5m, Taravale Formation, dehiscensZone. Crown view. P-Q, scale AM F101204 from YW]117.5, Cavan Formation, dehiscens Zone. P, crownview; Q, lateral view. R-S, scale AM F103585 from YW]117.5, Cavan Formation, dehiscens Zone. R, crownview; S, lateral view. T-V, scale AM FlO1177 from SL051m, Taravale Formation, dehiscens Zone. T, crownview; 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.

Emsian microvertebrates from Buchan and Taemas17

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Superclass Agnatha Haeckel, 1895

Subclass Thelodonti Kiaer, 1924

Order Thelodontida Stensio, 1958

Family Turiniidae Obruchev, 1964

Genus Turinia Traquair, 1896

Turinia sp. cf. T. australiensis Gross, 1971

Material ExaminedEighteen scales: lO head scales, four transitional

(cephalopectoral) scales, four trunk (postpectoral)scales.

LocalitiesSlocombe's section at Buchan, samples SL 46.5m,

SL 48.7m (Buchan Caves Limestone), SLO 28.5m,SLO 5lm, SLO 5l-68m, SLO 213m (TaravaleFormation); Section YWJ at Taemas, samples117.5m (Cavan Formation) and 135.4m (MajurgongFormation).

Stratigraphic RangeEarly Devonian, Emsian: dehiscens - perbonus

conodont zones at Buchan; dehiscens Zone atTaemas.

Description

Head ScalesThe head scales (Figure lA-L) are circular or

subcircular. The central part of the crown isunornamented and, in the elongated scales (FigurelD-L), flat. Approximately 14 main ribs around therim of the crown are separated by deep troughs. Inscale AM FlO1174 (Figure lA-e), the troughs aredeeply incised with parallel sides, and on parts ofthe margin long and short troughs alternateproducing a bifurcation of some of the ribs. Thetroughs in scales AM FlO1175 (Figure lG-I) and AMFI03583 (Figure IJ-L) become narrower towards thecentre of the crown and terminate in shallowelongated depressions. The ribs are subdivided bynarrow vertical grooves; in scale AM FlO1174 (FigureIA-e) these occur only on the sides of the scale andgive the outline a serrated appearance, in scales AMFlO1175 (Figure lG-I) and AM F103583 (Figure lJ­L) the grooves extend a short distance onto thecrown, while in scale AM FlOl203 (Figure ID-F) thevertical grooves on the distal ends of the ribs are notwell developed (see Figure IF).

In scales AM FlO1174 (Figure lA-e), AM FlOI203(Figure ID-F) and AM FlO1175 (Figure lG-I) thecrown and base are separated by a constricted neck.The base is shallow and comprises a low roundedrim surrounding a central large, deep pulp cavity.In scale AM FlO3583 (Figure IJ-L) the neck is notconstricted and is not clearly defined (Figure lL).

A. Basden

The base is deeply convex and contains one smallcentral pulp cavity and a smaller cavity towardsone side (Figure lK).

Transitional ScalesCrown is elliptical (Figure 10) or diamond­

shaped (Figure lM). Deep troughs aligned from theouter anterior margin posteriorly towards the centreof the crown produce coarse ribs covering virtuallythe entire crown surface. In scale AM FI03584(Figure 10) these ribs are separated by a narrowcentral ridge running from anterior to posterior;the ribs alternating in their anterolateral directionfrom this central ridge to the lateral margin of thecrown. A central ridge is less pronounced in scaleAM FlO1l76 (Figure IM), where the ridges andtroughs are all aligned from the anterior and lateralmargins towards the posterior of the crown.Troughs on the crown of scales AM FlOI204 (FigureIP) and AM F103585 (Figure lR) terminate closer tothe scale margin, leaving a smooth, flat central areaon the crown.

The base is the same shape as the crown, butprojects forward from the crown, while the crownoverhangs the base posteriorly. The base contains acentral pulp cavity surrounded by a rounded rim.The crown joins directly to the base without aconstricted neck, although in scales AM FlOl204(Figure IP) and AM FI03585 (Figure IR) there is ashallow trough around the base of the crown,similar to that found on some body scales, e.g.Figure IT.

Trunk ScalesThere are two distinct morphological types of

trunk scales. The more commonly occurring type isillustrated in Figure IT-W. Scale AM FlO1l77(Figure IT-V) is elongated horizontally with thebase extending as a long anterior processapproximately two-thirds the total length of thescale. The complex crown structure comprisesseveral overlapping, deeply subdivided layerssurrounding a central lanceolate platform. Thiscentral platform bears ridges and troughssUbparallel to the margin. The central platform issuccessively underlain towards the posterior of thecrown by three similar, larger, ridged layers. Theposterior part of the crown has been broken;overall crown shape is difficult to determine. Theposterior part of the crown substantially overhangsthe base (Figure 1U), with 10 parallel ribs extendingfrom the posterior crown-base interface to theposterior edge of the crown (Figure 1U). Theanterior neck region is a wide, shallow trough. Thebase is shallow and flat with a small central pUlpcavity, and the tapering anterior process extendssmoothly from the base in the same horizontalplane. Other scales of this type, e.g. Figure 1W, havecrown ornament of similar overlapping layers with

Emsian microvertebrates from Buchan and Taemas

less deeply incised troughs, and a shorter anteriorprocess on the base.

The second scale type is represented by only onespecimen (broken during electron micrography;Figure IX). The flat diamond-shaped crown has acentral, diamond-shaped, slightly raised portionwith three wide, shallow anterior ridges separatedby wide, shallow troughs. This central area issurrounded laterally and posteriorly by a larger,flat, diamond-shaped area with the lateral cornersslightly stepped down, and the posterior cornerextending slightly into a rounded point. On eitherside of this posterior point there is a lower, narrow,elliptical platform. There is no dearly-defined neckarea. The base is extended into a short, stout,tapering anterior process approximately one-thirdthe total length of the scale.

DISCUSSION

Some Buchan/Taemas thelodont scales sharefeatures with those of Turinia australiensis fromwestern Queensland (Turner et £11. 1981) and fromthe Jerula Formation of central NSW (pesavis ­sulcatus zones) (Turner 1997, figures 3C, 3K, 3N, 4G,41, 4M); T. antarctica from southern Victoria Land,Antarctica (Turner and Young 1992); T. gondwanafrom Bolivia (Gagnier et £11. 1988); T. ftlscina fromwestern Victoria (Turner 1986), and scales fromwestern Queensland described as TuriniagavinyOlmgi by Turner (1995a) and from SouthAustralia described as Turinia d. T. pagoda by Longet £11. (1988). Scales from the Cravens Peak Beds ofwestern Queensland (Eifelian), and the AztecSilts tone in Antarctica (Givetian), held in thecollections of the Queensland Museum and theAustralian National University, also showsimilarities with the Buchan/Taemas specimens.

Crown ornamentation of distinct, parallel-sidedtroughs between sometimes bifurcating ribs onhead scale AM F101203 (Figure 10) is seen in headscales of T. australiensis (Turner et £11. 1981, figurellA). The Buchan specimen lacks the deep roundedbase of typical head scales of T. australiensis (Turneret al. 1981), although the depth of the base inthelodont scales is related to ontogenetic stage (S.Turner pers. comm. 1995). Similar crown ornamentoccurs in specimens from samples C654 from theJerula Formation at Condobolin (see also Turner1997, figure 3F) and C657 from the Trundle Groupat Kadungle, held in the collections of theQueensland Museum.

The second type of head scale morphology, withtapering troughs terminating in shallowdepressions (Figure IG-L), can be found inspecimens from the Officer Basin, South Australia(Turinia sp. aff. T. australiensis - Long et £11. 1988,figure 5A), Cravens Peak Beds (T. australiensis ­Turner et a1. 1981, figure 100), and in collections at

19

the Australian National University from shot pointand outcrop samples from the Cravens Peak Beds(T. australiensis, sample SP813, and T. gavinyollllgi,sample GB77, respectively), and from the AztecSiltstone (T. antarctica, sample V2616).

The crown of transitional scale AM F101176(Figure IM-N) also resembles that of T. allstraliensis(e.g. Turner et a1. 1981, figure 111), with crenulationsdeeply incising the crown almost to the centre,leaving only a narrow central smooth area (FigureIM). Scales of T. gavinyOlmgi figured by Turner et £11.(1981, figure 7A) have a similarly crenulated crown,but the crenulations here are more widely spacedand the central smooth ridge is wider than in theBuchan specimen. Subparallel troughs extendingfrom the margin almost to the central antero­posterior axis of the crown are also found in scalesof TlIrinia d. T. pagoda from the eastern OfficerBasin, South Australia, figured by Long et £11. (1988,figure 2A,B). Although the number and orientationof the troughs is similar to those on the crown ofBuchan specimen AM FI01176 (Figure 1M-N), thecrown of the South Australian scale is moreshallowly incised. The base, too, of the Buchan scale(Figure IN), which is shallow with a large centralpulp cavity and small anterior process, resemblesthat of body scales of Turinia sp. cf. T. pagoda (Longet £11. 1988, figure 2A-C,E). However, although theBuchan specimen shares these features with theTlIrinia sp. cf. T. pagoda material from SouthAustralia, it differs from the original description ofT. pagoda from western Yunnan, China (Wang et £11.1986) in lacking the lateral neck-spurs and byhaving more deeply incised grooves on the crown(see Wang et £11. 1986, figure 4G-J). A specimen of T.!lIscina, figured by Turner (1986, figure 2N),resembles the transitional scales from Buchan andTaemas in crown ornament.

The crown of trunk scale AM FlO1177 (Figure1T,V) resembles scales of T. australiensis from theJerula Limestone Member of the Glenmga Formationin western New South Wales (Turner 1997, figure4M), T. antarctica, described from the Aztec Siltstoneof southern Victoria Land, Antarctica by Turner andYoung (1992, figure 5H), and T. gondwana from theCatavi Formation, Bolivia (Gagnier et £11. 1988, figure40,E). Common features are the central raisedelliptical section with subparallel ribs, lateralsegments that converge with the posterior tip of thecentral platform in a thin ridge extending towardsthe posterior of the crown (e.g. Turner and Young1992, figure 5h; Gagnier et al. 1988, figure 40,E),parallel ribs on the ventral side of the crown (Turnerand Young 1992, figure 6; Gagnier et al. 1988, figure4), a smooth wide trough around the anterior edge ofthe crown (Turner and Young 1992, figure 5a, c-m),and a long anterior process on the base (Turner andYoung, 1992, figure 5d-€, g-m; Gagnier et al. 1988,figure 5B). The same pattern of tiered subdivisions of

20

the crown occurs in body scales from the EarlyLochkovian Read Bay Formation of CornwallisIsland, Canadian Arctic (Turner and Burrow 1997),and in specimens from outcrop locality GB77 of theCravens Peak Beds in western Queensland (Turner1995a, figure 3V,W; Queensland Museum collection).

CONCLUSIONThe turiniid scales described in this study show

most morphological similarity to scales of T.australiensis (see Turner 1995b, 1997 for discussion ofT. australiensis scale morphology, histology andsignificance of the taxon in biostratigraphicsubdivision of the Early Devonian). However,because of slight differences from previously figuredspecimens of T. australiensis, similarities with otherspecies of Turinia, and the unique morphology ofscale AM FlO1l74 (Figure 1A-e), the turiniid scalesfrom the Buchan/Taemas fauna described herein arereferred to Tuninia sp. cf. T. australiensis. The age ofthis fauna (perbonus - dehiscens zones) is the same asthat of the T. sp. cf. T. australiensis Assemblage(Fauna 3) of Turner (1997).

ACKNOWLEDGEMENTSI thank John Talent and Ruth Mawson for

providing the material for this study and givinghelp and encouragement throughout the project;Sue Turner for assistance with identification, accessto collections and financial help to visit Brisbane tostudy material; Gavin Young for continued helpand encouragement, and for providing access toAGSO and AND collections; the Linnean Society ofNSW and the SA and ACT branches of theAustralian Federation of University Women forfinancial assistance; David Mathieson for help withprinting; and Theresa Winchester-Seeto forongoing practical advice and encouragement. Partof this study was carried out during tenure of anAustralian Postgraduate Award.

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Manuscript received 14 January 1998; accepted 6 February1998.