Bollettino della Società Paleontologica Italiana Modena, Novembre 1999
Early Devonian microvertebrates from the Tyers-Boola area of centrai Victoria, Australia
Alison BASDEN Department of Earth and Planetary Sciences Macquarie University
KEY WO RDS - Microvertebrates, Systematics, Early Devonian, Australia.
ABSTRA CT- The vertebrate microfauna from the Coopers Creek Formation, centra! Victoria, comprises platelets and derma! bo ne fragments from brachythoracid and acanthothoracid placoderms, rare chondrichthyan remains, scales of the acanthodians Trundlelepis cervicostulata, Nostolepoides platymarginata, Gomphonchus? bogongensis, Nostolepis sp., Acanthoides sp., Cheiracanthoides sp_. and Poracanthodes sp., acanthodian fin spines, and rare osteichthyan teeth. Accompanying conodonts date the fauna as sulcatus-kindlei zones (Pragian, Early Devonian). Comparisons with coeva! faunas from southeast Australia reveal many common taxa.
RIASSUNTO- [Microvertebrati del Devoniano inferiore nell'area di Tyers-Boola nel Victoria centrale, Australia]- La microfauna a vertebrati rinvenuta nella Formazione di Coopers Creek, nel Victoria centrale comprende placche palata/i e placche derma/i di placodermi brachitoracidi e acantotoracidi, rari resti di condricti, scaglie degli acantodi Trundlelepis cervicostulata, Nostolepoides platymarginata, Gomphonchus? bogongensis, Nostolepis sp., Acanthoides sp., Cheiracanthoides sp. e Poracanthodes sp., spine caudali di acantodi e rari denti di osteicti. l conodonti rinvenuti negli stessi livelli consentono di datare la fauna alle biozone a sulcatus-kindlei (Pragiano, Devoniano inf). Un confronto con faune coeve del! Australia sud-orientale evidenzia come molti taxa siano comuni.
INTRODUCTION determined by conodonts, chitinozoans or corals, are indicated in Text-fìg. 2. Microvertebrates are becoming increasingly useful for both biostratigraphic correlation and palaeo- environmental interpretation. The systematic collection GEOLOGY AND PREVIOUS WORK of vertebrate microremains and conodonts together, in sections spanning severa! conodont zones, is a more The Coopers Creek Limestone at Tyers (Text-fìg. precise technique than isolated spot sampling for 3) comprises limestones with interbedded mudstones, establishing the stratigraphic range of microvertebrate sandstones an d conglomerates. Detailed geology of the species, as previously demonstrated by De Pomeroy area is given by Mawson & Talent ( 1994). Relifìsch & (1994, 1996) and Basden (1999). Webb (1993) interpret the Coopers Creek Limestone The present study documents the fìsh microfossils as a primarily carbonate fan deposited in deep water. in the Coopers Creek Limestone at Tyers and Boola Conodonts recovered from the Coopers Creek quarries in centrai Victoria (Text-fìg. 1), and dates the Limestone atTyers and Boola quarries indicate an age microvertebrate occurrences using the accompanying spanning the sulcatus to dehiscens zones (Mawson & conodonts. The Tyers-Boola vertebrate microfauna is Talent, 1994, fìg. 2); Tyers quarry is the type locality compared with similar faunas from other Early for the zonally signifìcant Eognathodus sulcatus Philip, Devonian limestones in southeastern Australia: while the incoming of lambda and mu morphs of E. Windellama (Mawson, 1986; Basden et al., in press) sulcatus indicates kindlei Zone, an d that of Polygnathus and Burrinjuck (Schultze, 1968; 0rvig, 1969; White, dehiscens dehiscens Philip & Jackson indicates dehiscens 1978; Giffìn, 1980; Long & Young, 1988; Mawson et Zone. Conodonts were recovered consistently al., 1992; Basden, 1999; Basden et al., in press) in throughout the two sections sampled (TQ and BOO, southern New South Wales; around Wellington see Mawson & Talent, 1994, fìg. 4), whereas most of (Mawson et al., 1988; Basden et al., in press) and the vertebrate microremains from the same residues Condobolin (Pickett & McClatchie, 1991; Pickett, described here occur only in the lower part of the 1992; Burrow, 1996, 1997) in centrai New South sections. Wales; Buchan in eastern Victoria (Long, 1984; Other studies in the Tyers-Boola area have revealed Mawson, 1987; Basden, 1999; Basden et al., in press); a diverse invertebrate fauna: tabulate corals and a and Point Hibbs in western Tasmania (Burrow et al., cyanophyte (Chapman, 1907); rugose and tabulate 1998). Stratigraphic ranges of these other limestones, corals, brachiopods and stromatoporoids (Philip, 1960, 528 A. BASDEN
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Windellama vertebrate fauna, and the presence or a borehole in Western Australia. lndeed, turiniid absence of thelodont scales. scales occur in localites across Australia from Early No thelodont remains have been recovered from unti! early Late Devonian (Turner, 1993, 1997). Tyers-Boola, Windellama or Pt Hibbs. However, Studies from regions as widely separated as the Baltic severa! centrai NSW localities, Buchan an d Burrinjuck and Australia indicate that nearshore sediments ali have turiniid scales very much like Turinia commonly comprise a high proportion of thelodont australiemis, originally described by Gross ( 1971) from scales relative to acanthodian scales (e.g. Marss & Einasto, 1978; Turner, 1986, 1991, 1995a, 1995b, 1997; Burrow, 1997). The absence of thelodont scales from the faunas a t Tyers-Boola, Windellama and Point Hibbs, and their presence in the otherwise-similar faunas from Buchan, Burrinjuck and the centrai New South Wales localities, possibly indicates that the depositional environment at Tyers-Boola, Windellama and Point Hibbs was in deeper water. Evidence of water depth is also provided by the abundance and composition of both conodont and invertebrate faunas. The greater abundance of conodonts from the Tyers-Boola area, compared with other Early Devonian horizons in southeastern Australia, is considered indicative of deeper water (e. g. Mawson, 1987; Mawson et al., 1992; Mawson & Talent, 1994). In contrast, generally low yields of conodonts from the Buchan Caves Limestone at Buchan and the Cavan Formation at Burrinjuck suggest very shallow water (Mawson et al., 1992). The invertebrate faunas, too, at Burrinjuck - brachiopods, bivalves, and gastropods in the Cavan Formation and spiriferids and rare lingulids in the Majurgong Formation - are typical of nearshore intertidal or brackish environments (Young, 1969). The thelodont scales recovered from section YWJ at Burrinjuck occurred near the Cavan-Majurgong boundary (Basden, 1999). Conodont studies have shown that, in the Devonian, polygnathid-dominated faunas occur in deeper and quieter water than those dominated by icriodontans or ozarkodinans (Mawson et al., 1988). Species of Ozarkodina are abundant at both Windellama (Mawson, 1986) and some sections at Wellington in centrai New South Wales (Mawson et al., 1988). At Buchan, all the thelodont scales recovered were from the SL/SLO section, which spans the Buchan Caves Limestone and Taravale Formation (Basden, 1999). The Buchan Caves Limestone conodont fauna Ed Alluvials N is dominated by ozarkodinans with polygnathids D Tertiary and Cretaceous absent which, together with a low diversity of D Walhalla Group brachiopods and rare corals, is consistent with very Coopers Creek Formation shallow water (Mawson et al., 1992). The overlying Taravale Formation has ozarkodinids dominating the D Beola Formation conodont fauna, but with species of Polygnathus also stratigraphic sections t present (Mawson, 1987). The Taravale Formation, - o 200 400 lorestry roads containing pelagic ammonoids and dacryoconarids me tres (Mawson et al., 1988), represents deepening conditions, a result of a widespread transgression that Texr-fìg. 3- Geology ofTyers-Boola area, Victoria (from Mawson took place in the dehiscens Zone (Talent & Yolkin, & Talenr, 1994, fig. 1). 1987; Talent, 1989). 530 A. BASDEN
CONODONT ZONE sulcatus kindlei PLACODERMS +
...... _.. _,_ ...... -... ---.-...... _...... ! ...... 1 ...... _...... - .... __ L ...... ?...... Q ...... _,_,_,_,...... ,_, ...... "'-""" --"""" """"""' ..... _...... ,_ ...... 1 .. _.. , .., ___ ...... _...... ___ , ....-...... 1 ..... """"""' """"""' """"""' """"'"" ,_, ...... Placodermi oen. et sp. indet. 1 4 2 CHONDRICHTHY ANS :==:::: ::== :::::=:::: :::::::::: :::::::::::· ::::::::· :::==::: ::=:=::: :::::: :=::::: :::::::::: ::::::::· ::::::::: ::::::::::: :::r:::::::::::: :::::::::: ::::::::::::: ::::::::::: :::::::::::: Elasmobranchii gen . et sp. indet. B 1 1
Gnathostomata gen. et sp. indet.
00 a CD N M ,... co M N co co M M Cl! .q- M N .q- co ..... N M c:i ,... C\i M ..,; ..,; u; Ili Ili Ili r-.: c:i ,... C\i M ..,; u; Ili cci cci ,... ,... ,... ,... ,... ,... ,... ,... ,... M M CONODONT ZONE sulcatus kindlei PLACODERMS
...... _...... _...... ______.. ,_. __ ,_, __ ...... ?...... Placodermi g_en . et sp. indet. 1 ACANTHODIANS :::::::::::: ::::::::: ::::::::::: ::::::::: :::::::::: ::::::::::: ::::::::: :::::::::: ::::::::::: :::::::::::. :::::::::::: ::::::::: :::I :: ::::::::: ::::::::: :::::::::::: :::::::::::: ::::::::::: ...... _.. , "'""'"" ...... ""'"""' ... _t ...... """""" ...... """""""" """"...... _, """""" """"""......
..· ...... ···1... ···1·· ...... s -· ....1 ...... '3' ...... } -· ...... 1 _ .....1" ......
Text-fig. 4 - Distribution and abundance of vertebra te microremains from (above) Tyers Q uarry section (TQ) and (below) Boola Q uarry section (BOO) of the Coopers Creek Limeston. EARLY DEVONIAN MICROVERTEBRATES FROM A USTRALIA 531
TYERS Wlndellama Centrai NSW Pt Hlbbs Buchan Burrln]uck
THELODONTS Turinia sp. cf. T. australiansis x x x PLACODERMS ACANTHOTHORACIDS: murrindalaspid x x x x brindabellaspid x x x x weejasperaspid x x x x acanthothoracid indet. x x x x x BRACHYTHORACID: buchanosteid x x x x x ACANTHODIANS CLIMATIIDS: Nostolepis sp. x x x x x x Nostolepoidas platymarginata x x x x ISCHNACANTHIDS: Trundlalapis carvicostulata x x x Gomphonchus? bogongansis x x x Chairacanthoidas sp. x x x x Poracanthodas sp. x x x CHONDRICHTHYANS x x x x OSTEICHTHY ANS x x x x x Text-fig. 5 - Comparison of vertebrate microfaunas from Tyers-Boola and other Early Devonian limestones in southeastern Australia.
SYSTEMATIC PALAEONTOLOGY postbranchial lamina fragment (Pl. l , fig. 2) bears widely-spaced tubercles with a convex denticulate Prefixes TQ and BOO referto the sections a t Tyers margin. The base is flat overall but is pitted with and Boola quarries, respectively (Text-fig. 3). Sample irregular depressions and occasionallow ridges. numbers following the prefix indicate tlie number of metres above the base of the section (see Mawson & Remarks - AMF l 06387 (P l. l, fig.l) resembles Talent, 1994, fig. 4 for stratigraphic sections). Histo- s_pecimens of Brindabellaspis stensioi described and logical study might improve designation to taxa for fìgured by Young (1980), while the ornament on some specimens. This has not been undertaken at this AMF l 06388 is typical of that found o n the stage because of either low numbers of specimens or postbranchial lamina of Brindabellaspis (e.g. Young, mineralisation. Figured specimens are housed in the 1980, Pl. 2, figs. 4-5), although similar ornament can palaeontological collections of the Australian Museum, be found on other acanthothoracids (C.J.Burrow, pers. Sydney, and bear the prefix AMF. comm., 1999). The specimens described by Young (1980) were recovered from the Receptaculites and Warroo limestones at Burrinjuck. These are the third Phylum CHORDATA and fourth of six units of the Taemas Formation, which Subphylum VERTEBRATA has been dateci by conodonts as late dehiscens to serotinus Superclass GNATHOSTOMATA zones (Emsian) (Pedder eta!., 1970). Microremains of Class PLACO DERMI M 'Coy, 1848 Brindabellaspis from the same horizons have recently Order ACANTHOTHORACI Stensio, 1944 been described (Burrow & T urner, 1998). Scales an d Family BRINDABELLASPIDIDAE Gardiner, 1991 fragments with ornament similar to Brincktbellaspis have been found from the Lochkovian-Pragian, in the Genus BRINDABELLASPIS Young, 1980 Connemarra Formation, Trundle Group (pesavis- BRINDABELLASPIS sp. sulcatus zones), and the Garra and Camelford lime- Pl. l , figs. 1-2 stones (Basden et al., in press).
Description - The ornament on fragment Materia!- AMF106387 from B0010.6 (kindlei AMF l 06387 (P l. l, fig. l) consists of broad, flat, Zone), AMFl 06388 from BOO 1.9 (sulcatus Zone) and smooth-topped tubercles with rounded or crenulated six other specimens from samples TQ6.4 and TQ6.6 margins, separated by deep, narrow troughs. The base (sulcatus Zone), and TQ7.3, B008.2, B0011.4 and is very slightly concave and unornamented. The B0014.1 (kindlei Zone). 532 A. BASDEN
Family WEEJASPERASPIDIDAE White, 1978 ornament comprising elongate, nodulose, ridged tubercles. Similar microremains have been recovered Genus WEEJASPERASPIS White, 1978 from the Emsian Receptaculites Limestone (Burrow & WEEJASPERASPIS sp. Turner, 1998) and Cavan Formation (dehiscens Zone) P l. l, fìgs. 3-4 at Taemas and the Lochkovian-Pragian Garra Limestone in centra! NSW (Basden et al., in press). Remarks- Ornament on AMFl 06389 is similar to the elongate multi-ridged tubercles described by White Materia!- AMF106390 from TQ6.8 (sulcatus (1978, pl. 3) for Weejasperaspis gavini from Burrinjuck. Zone) and 26 other specimens from samples TQ7.3, As noted by Long (1984) and Long &Young (1988), TQ9.1, TQ11.8, B0010.6, B0012.2, B0014.1, the ornament on scales and dermal bone of B0015.3 and B0036.8 (kindlei Zone). Weejasperaspis an d Murrindalaspis is superfìcially similar, the ridges on Murrindalaspis being slightly shorter; the two genera are discriminateci mainly by differences ACANTHOTHORACI gen. et sp. indet. in size and shape of dermal plates. The Burrinjuck Pl. l fìgs. 6-9 materia! was from the Spirifer yassensis and lower Receptaculites limestones of the Emsian Taemas Remarks - Stellate tubercles with acute ridges Formation. separated by deep valleys are typical ornament on acanthothoracids (Goujet, 1984), as are the small Materia!- AMF106389 from B0014.1 (kindlei nodules o n the radiating tubercular ridges of fragment Zone) and another specimen from TQ3.3 (sulcatus AMF106393 (Pl. l, fig. 9), visible under high Zone). magnifìcation. Scales with similar ornamentation, recovered from the Emsian Xiejiawan Formation of Genus MuRRINDALASPIS Long, 1984 Longmenshan, South China, have been assigned to the MuRRINDALASPIS sp. new acanthothoracid genus Xiejiawanaspis by Burrow P l. l, fìg. 5 et al. (in press; pl. 9). However, the ridges on the tubercles of the Chinese specimens are sharper an d fìner Remarks - Murrindalaspis was described by Long than on the Tyers-Boola materia!. ( 1984) from the Murrindal Limestone (perbonus Zone) near Buchan, Victoria. The scales from Tyers-Boola Materia! - AMF106391 from B0012.2, resemble those described by Long (1984), those fìgured AMF l 06392 from Boola Upper Quarry, AMF l 06393 by Long & Young (1988, fig. 9), and severa! from TQ7.3 (all kindlei Zone) and 43 other specimens morphological types held in the CPC collection at the from samples TQ3.5, TQ6, TQ6.4, TQ6.6, BOO 1.9, Australia National University, Canberra, in the B003.3 and B004.2 (sulcatus Zone) and TQ7.3,
EXPLANATION OF PLATE l
Placoderm remains from the Coopers Creek Li m estone. Scale bar = l OOf.im, unless otherwise specified.
Figs. 1-2 - Brindabellaspis sp. l) Platelet AMF106387 from B0010.6, kindlei Zone. Scale bar= lmm; 2) Postbranchiallamina ornament AMF106388 from B001.9, sulcatus Zone. Figs. 3-4 - Weejasperaspis sp. platelet, AMF106389 from B0014.1, kindlei Zone. 3) side view; 4) top view. Fig. 5 - Murrindalaspis sp. platelet, AMF106390 from TQ6.8-6.9, sulcatus Zone. Figs. 6-9 - Acanthothoraci gen. et sp. indet. platelets. 6) AMF l 06391 from BOO 12.2, kindlei Zone; 7, 8) AMF106392 from Boola Upper Quarry, kindlei Zone. 7) top view; 8) side view; 9) AMF106393 from TQ7.3, kindlei Zone. Figs. 10-13 - Buchanosteus sp.[latelets. lO) AMF10639 from TQ6.4-6.5, sulcatus Zone; 11) AMF106395 from Boola Upper Quarry, kindlei Zone; 12) AMF106396 from B003.3, sulcatus Zone; 13; AMF106397 from TQ4.7-4.8, sulcatus Zone. Figs. 14-15 - Wijdeaspis sp. fragments. 14) AMF106398 from TQ2.5-2.7, sulcatus Zone, Scale bar= lmm. 15) AMF106411 from B0011.4, kindlei Zone, top view. Figs. 16-18 - Placodermi gen. et sp. indet. remains. 16) Fragment AMF106399 from TQ3.3, sulcatus Zone; 17) Platelet AMF106400 from TQ11.8-11.9, kindlei Zone; 18) Fragment AMF106401 from TQ3.3, sulcatus Zone. A. BASDEN, EARLY DEVONIAN MICROVERTEBRATES FROM AUSTRALIA P!. l 534 A. BASDEN
TQ9.1, TQ11.8, TQ12.2, B008.2, B008.3, Materia!- AMF106398 from TQ3.3 (sulcatus B0010.6, B0012.2, B0013.3 and B0014.1 Zone), AMF106411 from B0011.4 (kindlei Zone) (kindlei Zone). and four other specimens from samples TQ2.5 (sulcatus Zone) and TQ7.3, B0011.4 and B0014.1 (kindlei Zone). Order ARTHRODIRA Woodward, 1891 Suborder BRACHYTHORACI Gross, 1932 Family BucHANOSTEIDAE White, 1952 Placodermi gen. et sp. indet. Pl. l, figs. 16-18 Genus BucHANOSTEUS Stensio 1945 BucHANOSTEUS sp. Description - AMF106399 (Pl. l, fig. 16). P l. l, figs. l 0-13 Ornament comprises stellate tubercles, either narrow and crescent-shaped or broad and rounded. Parts of Remarks - Similar platelets from Burrinjuck some of the tubercular ridges are indistincdy nodose (unknown locality), Buchan (Murrindal Limestone, under high magnification, suggesting possible perbonus Zone) and Tyers (Coopers Creek Formation) acanthothoracid affinity. The base of the fragment is with ornament of rounded stella te tubercles, sometimes flat an d ornamented by low, irregular indentations an d surrounded by an outer margin of radiating 'mucus ridges. grooves' (see Pl. l, figs. 12, 13), have been ascribed by AMF106400 (Pl. l, fig. 17). The crown is orna- Burrow and Turner (1998) to Buchanosteus orto the mented with inclined spatulate or pointed tubercles, family Buchanosteidae. Buchanosteid scales have also with longitudinal ridges extending either pardy or fully been recovered from the Taravale Formation at Buchan from the tip of the tubercles to the base. The base of (perbonus Zone) and the Upper ReefFormation at Wee the platelet is smooth and moderately concave. Jasper (Basden eta!., in press). Turner (1991) reported AMF106401 (Pl. l, fig. 18). Ornament consists of buchanosteid scales from horiwns as old as Lochkovian. wide, smooth-topped parallel ridges separated by deep troughs of approximately the same width. The edges Materia!- AMF106394 from TQ6.4 (sulcatus of the ridges are indented by small rounded Zone), AMF l 06395 from Boola Upper Quarry (kindlei embayments. The base of the fragment is fairly flat Zone), AMF106306 from B003.3 (sulcatus Zone), with low, irregular ridges. AMF l 06397 from TQ4. 7 (sulcatus Zone) and 19 other specimens from samplesTQ0-2, TQ2, TQ3.3, TQ4.7, Remarks - The ornament on the fragments TQ6 and TQ6.6 (sulcatus Zone) and TQ11.8, illustrateci (P late l, figs. 16 -18) cannot be classified as B0012.2 and B0014.1 (kindlei Zone). any of the placoderm types discussed by Goujet ( 1984) and so this material is placed in open nomenclature at this stage. Order PETALICHTHYIDA}aekel, 1911 Materia!- AMF106399 & AMF106401 from Genus WIJDEASPIS Obruchev, 1964 TQ3.3 (sulcatus Zone), AMF106400 from TQ11.8 WIJDEASPIS sp. (kindlei Zone) and six other specimens from samples Pl. l, figs. 14-15 TQ2, TQ3.3, TQ6.8 and B002.4 (sulcatus Zone) and TQ11.8 (kindlei Zone). Description- The ornament consists of fine laterally- compressed tubercles arranged in dose parallel rows. The underside of the fragments is smooth and very slighdy concave, following the curvature of the Class CHONDRICHTHYES Huxley, 1880 fragment as a whole. Subclass ELASMOBRANCHII Bonaparte, 1838
Remarks - Plates of the petalichthyid Wijdeaspis Elasmobranchii gen. et sp. indet. A warrooensis, described and figured by Young (1978) Pl. 2, fig. 14 from the Crinoidal Limestone of the Emsian Taemas Formation at Burrinjuck, bear similar ornament to the Description - AMF l 06412 is a tricuspid tooth material from Tyers. A fragment with similar ornament attached to a base. The cusps are conical and tapering, t o AMF l 06411, al so from the Coopers Creek the middle cusp is larger than the two lateral ones, and Formation, was described and figured by Philip (1965, the broken distai end shows a narrow longitudinal pl. 8, fig. 42) as a possible acanthodian fin spine. The centrai canal. The lateral cusps originate from the labial six specimens described here constitute the earliest side of the base and diverge distally, while the middle report of this genus. cusp is placed further towards the lingual side of the EARLY DEVONIAN MICROVERTEBRATES FROM AUSTRALIA 535 base and curves lingually. The base, of cancellous bony TQ0-2, TQ2, TQ4.7, TQ6.6, TQ6.8, B003.3, tissue, is slightly concave. B004.2 (sulcatus Zone) and TQ9.1, TQ11.8, B008.2, B008.3, BOO 14.1 an d BOO 16.2 (kindlei Materia!- AMF106412 from TQ7.3 (kindlei Zone). Zone).
Genus NosTOLEPOIDES Burrow, 1997 Elasmobranchii gen. et sp. indet. B NosTOLEPOIDES PLATYMARGINATA Burrow, 1997 Pl. 2, figs. 15-16 Pl. 2, figs. 2-3
Remarks- Both specimens are the broken, abraded, Description - The crown consists of a centrai broadly striated distai ends of conica! teeth or denti cles. triangular section with short subparallel ribs extending They are placed in open nomenclature because of a from the rounded anterior margin a short distance lack of distinguishing features. Irregular vascular canals towards the pointed posterior (Pl. 2, fig. 2). The lower penetrating the base of AMF l 06414 (P!. 2, fig. 16) section of the crown is a lateral platform of varying suggest even a possible placoderm affinity (S. Turner, width that extends beond the upper posterior point. pers. comm., 1999). The neck is clearly defined, and shallower anteriorly than posteriorly (Pl. 2, fig. 3). The subrhombic base is Materia!- AMF106413 from TQ7.7 (kindlei gently convex, centrally vaulted an d separated from the Zone), AMF106414 from TQ6.4 (sulcatus Zone). neck by a distinct rim.
Remarks - These scales resemble Morphotype l scales of Nostolepoides platymarginata, described by Class ACANTHODII Berg , 1922 Burrow (1997) from the Gleninga (pireneae-dehiscens Order CLIMATIIFORMES Berg, 1940 zones) and Troffs formations (kindlei-dehiscens zones) Family CuMATIIDAE Berg, 1940 in centrai NSW. The scales are morphologically similar to the Late Silurian-Early Devonian Nostolepis striata Genus NosTOLEPIS Pander, 1856 Pander but can be distinguished from that species NosTOLEPIS sp. histologically and by the posterolateral ledge of the P!. 2, fig. l crown being wide and/or irregular and continuing around the posterior point, as discussed by Burrow Description-The crown is subtriangular, and slopes (1997). Probable Nostolepoides scales have also been up from a rounded anterior margin to a pointed recovered from the early Emsian Taravale Formation posterior that extends beyond the base. Short, low, and Murrindal Limestone at Buchan (Basden et al., in parallel ridges ornament the anterior p art of the crown. press, Text-figs 9.1 - 9.3). One of the scales figured by The neck is shallow anteriorly an d dee per an d concave Philip (1965, Pl. 8, fig. 38) from the Tyers area has posteriorly, with no or few visible p ore canal openings. been assigned by Burrow (1997) to Nostolepoides The base and neck are separated by a distinct rim. The platymarginata. centrally convex base is subrhombic, with anterior and posterior corners slightly rounded and latera! corners Materia!- AMF106404 from TQ11.8 (kindlei more pointed. Faint concentric lines are visible on the Zone) and 12 other specimens from samples TQ2, base. TQ2.5, TQ6.4, TQ6.6, TQ6.8 and B004.4 (sulcatus Zone) and TQ11.8, TQ12.2 and B0012.2 (kindlei Remarks - See Denison (1979) for diagnosis and Zone). De Pomeroy ( 1996) for comments o n the distribution of this cosmopolitan genus. Recorded occurrences of nostolepid scales in the early Devonian of eastern Order lSCHNACANTHIFORMES Berg , 1940 Australia include the Broken River Group and Martins Family lSCHNACANTHIDAE Woodward, 1891 Well Limestone in Queensland (Turner, 1991, 1993; De Pomeroy, 1996; Turner eta!., in press) and, in NSW, Genus TRUNDLELEPIS Burrow, 1997 the ]erula, Connemarra and Condobolin formations, TRUNDLELEPIS CERVICOSTULATA Burrow, 1997 Garra and Windellama limestones, Trundle Beds and P!. 2, figs. 4-5 Yarra Yarra Creek Group (Turner, 1991, 1993; Burrow, 1997). Description - The diamond-shaped crown is relatively Bat and bears short ribs extending from the Materia!- AMF106403 from B008.3 (kindlei anterior margin towards the centre of crown. The Zone) and 50 other specimens from samples TQO, posterior part of the crown is thin, overhangs the base 536 A. BASDEN and is ornamented by overlapping, V-shaped growth Genus GOMPHONCHUS Gross, 1971? lines (Pl. 2, fìg. 4). The neck is deep, slighdy more so GOMPHONCHUS? BOGONGENSIS Burrow, 1997 at the posterior, and concave (Pl. 2, fìg. 5). lt is separated Pl. 2, fìgs. 6-9 from both crown and base by a distinct rim, which may extend slighdy into flanges at the latera! points. Description- In both Morphotype l (P l. 2, fìgs. 6- The posteri or neck bears vertical slits an d pillars, giving 7) and Morphotype 2 (Pl. 2, fìgs. 8-9), the crown is a "buttressed" appearance (Burrow, 1997, p.59). The fan-shaped with a scalloped anterior margin, and a base is convex, more so anteriorly, and has concentric posterior point that is more elongated in Morphotype markings. 2. Morphotype 2 also has lower latera! crown ledges. Sharp ridges extend from che anterior margin of the Remarks - Scales of che acanthodians Trundlelepis crown towards che centre of the scale. The neck is cervicostulata were originally described from the pesavis- concave, deeper posteriorly, and separated from the base sulcatus zones of centrai NSW (Trundle Beds and ]erula by a distinct rim (Pl. 2, fìg. 7, 9). The base is centrally Formation; Burrow, 1997), but dating of these occur- convex and may show concentric striations (Pl. 2, fìg. rences has subsequendy been revised to pesavis-?sulcatus 7). and pireneae-dehiscens zones (Sherwin, 1996). The species is also reported from the sulcatus Zone at Point Remarks-These scales resemble those described by Hibbs (Burrow et al., 1998), but a recent study Burrow (1997) from the Trundle Beds and Gleninga (Winchester-Seeto & Carey, in press) has found the Formation in centrai NSW Revised dating (Sherwin, kindlei Zone indicator, the chitinozoan Bulbochitina 1996) now places the horizons with these scales imo bulbosa Paris, in sample PH 85-35, which contained pireneae-dehiscens zones. G.? bogongensis scales are also one T. cervicostulata scale (Burrow et al., 1998, p. 14). reported from the sulcatus Zone at Point Hibbs (Burrow The Tyers-Boola scales are indistinguishable from those et al., 1998). ofMorphotype l ofBurrow (1997). Morphotypes l and 2 can both be distinguished from Nostolepoides platymaginata scales by the scalloped Materia!- AMF106405 from TQ6.6 (sulcatus anterior margin of the crown, and the increased depth Zone) and 120 other specimens from samples TQO, of che neck towards che posterior, and Morphotype l TQ2, TQ2.5, TQ3.3, TQ4.7, TQ5, TQ6, TQ6.4, also by the lack of lowered latera! crown ledges. TQ6.6, TQ6.8, B000.3, B001.9, B003.3, B004.2 and B004.4 (sulcatus Zone) and TQ7.3, Materia!- AMF106406 from TQ6.8 (sulcatus TQ7.7, TQ9.1, TQ11.8, TQ12.2, B008.3, Zone), AMF106407 from TQ9.1 (kindlei Zone) and B0014.1, B0016.2, B0016.8 and B0037.3 56 other specimens from samples TQ6.4, TQ6.8 and (kindlei Zone). B005.8 (sulcatus Zone) and TQ7.3, TQ7.7, TQ9.1,
EXPLANATION OF PLATE 2
Acanthodians (1-13), Chondrichthyans (14-16) and ?Osteichthyans (17-19) from Coopers Creek Limestone. Scale bar= lOOf!m
Fig. l - Nostolepis sp. scale AMF106403 from B008.3, kindlei Zone, crown view. Figs. 2-3 - Nostolepoides p!arymarginata scale AMF106404 from TQ11.8-11.9, kindlei Zone. 2) crown view; 3) side view. Figs. 4-5 - Trundlelepis cervicostulata scale AMF106405 from TQ6.6-6.7, sulcatus Zone 4) crown view; 5) side view. Figs. 6-9 - Gomphonchus? bogongensis. 6, 7) Morphorype l scale AMF106406 from TQ6.8-6.9, sulcatus Zone. 6) crown view; 7) side view. 8, 9) Morphotype 2 scale AMF106407 from TQ9.1, kindlei Zone. 8) crown view; 9) si de view. Figs. l 0-11 - Poracanthodes sp. lO) Scale AMF106408 from TQ7.3, kindlei Zone, crown view. 11) Scale AMF106409 from TQ2-2.1, sulcatus Zone, crown view. Figs. 12-13 - Acanthodii gen. et sp. inder. fin spines. 12) AMF106410 from TQ6.8-6.9, sulcatus Zone, side view; 13) Fragment AMF106402 from TQ3.3, sulcatus Zone, top view. Fig. 14 - Elasmobranchii gen. et sp. inder. A tooth AMF106412 from TQ7.3, kindlei Zone. Figs. 15-16 - Elasmobranchii gen. et sp. inder. B. 15) Tooth fragment AMF106413 from TQ7.7-7.9, kindlei Zone; 16) Tooth fragment AMF106414 from TQ6.4-6.5, sulcatus Zone. Figs. 17-18 - Gnathostomata gen. et sp. inder. fragment AMF106415 from TQ7.3, kindlei Zone; 17) front view, 18) side view. Fig. 19 - Teleostomi gen. et sp. inder. fragment AMF106416 from TQ4.7-4.8, sulcatus Zone. A. BASDEN, EARLY DEVONIAN MICROVERTEBRATES FROM AUSTRALIA Pl. 2 538 A. BASDEN
TQ11.8, B008.2, B008.3, B0011.4, B0012.2, (Lochkovian-Pragian) of centrai NSW (Burrow, 1997, B0014.8, B0015.3 and B0016.8 (kindlei Zone). pl. 3, figs. 12-13); although some of the younger of these scales have subsequendy been reassigned to a new species of Cheiracanthoides (see discussion in Burrow Genus PoRACANTHODES Brotzen, 1934 et al., in press), the older specimens are stili considered PoRACANTHODES sp. to beMachaeracanthus (C. Burrow, pers. comm., 1998). P l. 2, figs. l 0-11 However, under high magnification, pore openings are visible o n the posterior crown surface connecting with Description - Generally large scales with a kite- slits underneath, indicating that the scales are, in fact, shaped crown. The posterior part of the crown is thin poracanthodid. Lack of well-preserved scales from and commonly broken but a complete specimen (Pl. Tyers-Boola precludes histological examination. 2, fig.10) measures approximately 2.2 mm in length. Anteriorly, the crown is rounded and ornamented with Materia!- AMF106408 from TQ7.3 (kindlei short radiating ribs. The posterior margin is pointed Zone), AMF106409 from TQ2 (sulcatus Zone) and and broadly denticulate. Overlapping growth zones on 15 other specimens from samples TQ2, TQ3.5 and the posterior part of the crown follow the oudine of TQ6.4 (sulcatus Zone) and TQ7.3, TQ9.1, TQ11.8 the posterolateral margin. These undulating growth and B008.2 (kindlei Zone). zones define low radiating ridges, each ridge running into an extension of the margin. Pores between the low ridges o n the posterior part of the crown open into Genus CHEIRACANTHOID ES Wells, 1944 slits on the underside of the crown. The neck is clearly CHEIRACANTHOID ES sp. defined and deeper at the posterior, where vertical pillars separate slit-like openings. The neck is separated Description-The elliptical or subrhombic crown is from the base by a distinct rim at the lateral and flat and ornamented on the anterior third to half by posterior edges that is less marked at the anterior. The between four and ten radiating or subparallel ridges. base is convex, with greatest depth towards the anterior, These ridges are connected along the anterior margin is rounded anteriorly and posteriorly and drawn into or the crown by a distinct flat rim that clearly separates flanges laterally, and shows faint concentric striations. the crown from the indented neck. The crown extends beyond the base posteriorly. The base is diamond- Remarks-The morphology of the Tyers-Boola scales shaped and centrally convex, with the neck-base margin is consistent with the diagnosis for Poracanthodes marked by a thickened rim. quijingensis by Wang & Dong (1989) from the Late Silurian of China, although the anterior margin of the Remarks- This cosmopolitan genus is frequently Chinese scales slopes smoothly towards the base (e.g. reported from Early and Middle Devonian horizons Wang & Dong, 1989, pl. IIIA) , in contrast to theTyers- (e.g. Wells, 1944; Valiukevicius, 1979, 1985; Veith, Boola scales where the anterior crown is flat and is 1980; Veith-Schreiner, 1983). Scales from the Recepta- separated from the base by a definite rim around the culites Limestone at Burrinjuck have been assigned to crown margin and a deeper, concave neck (Pl. 2, fig. the genus (Giffin, 1980, fig. 5), bur it is possible that 11). The Tyers-Boola specimens also resemble scales Cheiracanthoides has become a "bucket" taxon from the Silurian-Devonian ofEurope (Marss, 1997) comprising several different form genera with a similar assigned t o ?P. menneri. Scales recovered from an morphology; De Pomeroy (1996) discusses this unnamed Late Silurian sequence near Cumnock and problem. the Lochkovian-Pragian Camelford Limestone, both in centrai NSW, ascribed by Parkes to Poracanthodes Materia!- Seven scales from TQ2.5, TQ6, BOO 1.9 (in Basden et al., in press, text-figs. 5.11-12, 6.11-14), and B003.3 (sulcatus Zone), and B0014.1 (kindlei have similar crown ornament to the Tyers-Boola scales. Zone). Dental elements, possibly from P. menneri, have been recovered from the Lochkovian Windellama Limestone in sourhern NSW (Parkes in Basden et al., in press) Order ACANTHODIDA Berg 1940 and the Trundle Beds, Gleninga an d ]erula formations (Burrow, 199 5, 1997). The scales are also grossly similar Genus ACANTHOID ES Brotzen 1934 to Machaeracanthus scales (e.g. Gross, 1973; Goujet, AcANTHOID ES sp. 1976; Mader, 1986) and in particular resemble some of those figured specimens in overall shape an d crown Description-The crown is flat and unornamented, ornament (e.g. Gross, 1973,. pl. 29, figs. 1-8; Goujet, and the pointed posterior margin extends beyond the 1976, pl. 61, figs . 4-6, 8-10, Mader, 1986, pl. 2, figs . base. The neck is deep an d indented. The highly convex 2, 8d, 15). Machaeracanthus scales have been described base is deepest towards the anterior of the scale. from the Trundle Beds and Gleninga Formation EARLY DEVONIAN MICROVERTEBRATES FROM AUSTRALIA 539
Remarks- De Pomeroy (1996) discusses confusi o n specimen, the cusp-base boundary is marked by a between Acanthodes Agassiz 1833 and Acanthoides slight transverse ridge; d. ust a bo ve this ridge o n o ne Brotzen 1934. The scales from Tyers-Boola are assigned si de is a pore opening Pl. 2, fig. 18). The specimen to the latter genus, using Wells's concept of a form genus bears a slight resemblance to both branchia! denticles for smooth-crowned scales considered too old to belong (A. Blieck, pers. comm., 1999) and acanthodian tooth to the Carboniferous and Permian genus Acanthodes. whorls (e.g. Burrow, 1995). These scales do, however, resemble those figured by Gross (1973), Vieth (1980) and Storrs (1987) assigned Materia/- AMF 106415 from TQ 7.3 (kindlei to Acanthodes? dublinensis. Zone).
Materia/- Seventeen scales from TQ2-2.1, TQ4.7- 4.8 andTQ6.6-6.7 (sulcatusZone), andTQ7.3, TQ9.1, CONCLUSION TQ11.8-11.9 and B0014.1 (kindlei Zone). Comparison of the abundance and composition of microvertebrate, conodont and invertebrate faunas, as Acanthodii gen. et sp. indet. well as sedimentological studies, can assist with Pl. 2, figs. 12-13 interpretation of depositional environment. Such a comparison suggests that the Coopers Creek Limestone Description-The fin spine fragments are commonly in the Tyers-Boola area was deposited under conditions slightly curved longitudinally (e.g. Pl. 2, fig. 12) and that, while stili shallow marine, were slightly deeper are either single (Pl. 2, fig. 12) or multiple (Pl. 2, fig. than some other Early Devonian localities in sourheast 13) elements. The upper surface bears rows of round, Australia. Further, conodonts recovered from the same pointed tubercles. The lower surface is unornamented residues give a precise age to the accompanying and gently grooved longitudinally; faint lines marking vertebrate microremains, thus contributing to a the join between individuai elements are visible o n the microvertebrate biozonation for eastern Australia so base of the multiple elements. that eventually vertebrate microremains can be used alone as a biostratigraphic tool in marginai environ- Materia/- AMF106402 from TQ3.3 and ments where conodonts are not preserved, and also for AMF106410 from TQ6.8 (sulcatus Zone), and 225 inter-regional and inter-continental correlation. other fin spine fragments from samples TQO, TQ1, TQ1.3, TQ2.5, TQ3.2,TQ3.5,TQ4.7, TQ6, TQ6.8, B001.9, B002.4, B004.2, B004.4 and B006.7 (sulcatusZone) and TQ7.3, TQ9.1, TQ11.8, B008.2 and B0010.6 (kindlei Zone). I thank Professar John Talenr and Associate Professar Ruth Mawson for providing the materia! for srudy and giving ongoing encouragement and assistance, Carole Burrow and Ross Parkes fo:{roviding copies of unpublished manuscripts and translations, Teleostomi gen. et sp. indet. an far useful discussion, Theresa Winchester-Seero far helpful Pl. 2, fig. 19 comments and suggesrions ro the manuscript, and the Macquarie Universiry Postgraduate Fund for providing fìnancial assistance ro Description-AMF l 06416 (P l. 2, fig. 19) is hollow. presenr the resulrs of this srudy at the ECOS VII-IGCP 421 The two cusps are conica! and longitudianlly striated. conference in Bologna & Modena in June 1998. This is a contribution to IGCP421: North Gondwana M id-Palaeozoic The outer layer of the cusps is shiny dentinous tissue. Biodynamics. A bulbous margin separates the cusps from the base. Just below the cusp-base margin are severa! pore openings. The base appears socketed into bone, which implies either acanthodian or osteichthyan affinity. REFERENCES BASDEN, A.M., 1999, Emsian (Early Devonian) microvertebrates Materia/- AMF 106416 from TQ 4.7 (sulcatus from the Buchan and Taemas areas of southeastern Australia: Zone). Records of the Western Australian Museum, Supplement No. 57: 15-21, l fig. BASDEN, A.M., BuRRow, C., H ocKI NG, M., PARKES, R. & YoUNG , G.C., 1999, Siluro-Devonian microvertebrates from Gnathostomata gen. et sp. indet. southeastern Australia. In Blieck, A. & Turner, S. (eds.), P l. 2, figs. 17-18 Courier Forschungs-Institut Senckenberg, 12 fìgs. (in press) . BuRROW, C.J., 1995, Acanrhodian denral elemenrs from the Trundle Beds (Lower Devonian) ofNew South Wales: Records Description- AMF 106415 (Pl. 2, figs. 17-18) of the Western Australian Museum, 17: 331-341. comprises a seri es of varying-sized cusps, socketed onto BuRROW, C.J., 1996, Placoderm scales from the Lower Devonian a base. The cusp materia! is smoother than the spongy- of New Sourh Wales, Australia: Modern Geology, 20: 351- looking materia! of the base. At the back of the 369, l pl. 540 A. BASDEN
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