Records of the Western Australian Must'lll11 Supplement No. 58: 65~80 (2000).

Devonian trilobites from the Broken River region of northeastern Australia

Raimund Feistl and John A. TalenF

lInstitut des Sciences de l'Evolution, UMR 5554 du CNRS, Universite de Montpellier 11,34095 Montpellier Cedex 05, France

2 Macquarie University Centre for Ecostratigraphy and Palaeobiology (MUCEP),Department of Earth and Planetary Sciences, Macquarie University 2109, Australia

Abstract - The Broken River Group in northeastern Queensland hasproduced the first Middle Devonian trilobites from Australia. With theexception of an undetermined phacopid from Emsian shales, all faunas wererecovered from earliest Eifelian through Givetian to ?earliest Frasnian shallowwater bioclastic limestones and marlstones. The specifically determinable partof the material consists entirely of five new taxa: Proetus (Devonoproetlls)sparsinodosus (Eifelian), Proetlls (Devonoproetus) latil11argo (Eifelian), Burgesinal11awsonae (Eifelian), Seutellum tenuistriatum (Givetian) and Pllacops (Phaeops)broeki (Givetian) A new genus of proetine, Burgesina, with type species B.mawsonae n. sp. is proposed. Preliminary palaeogeographical comparisonssuggest only weak linkages with typical North Gondwanan faunas fromBohemia, southwestern Europe and North Africa. By contrast, the trilobitesfrom the Broken River region are closely related to faunas from InnerMongolia.

INTRODUCTION

Middle Devonian trilobites have not previouslyrecorded from Australia. The youngest previouslyrecorded were from the Taemas Group ofsoutheastern New South Wales (Chatterton 1971);these are no younger than mid-Emsian,appoximately perbonus-inversus zones (R. Mawson,pers. comm.). Trilobites are herein recorded for thefirst time from the Middle Devonian of Australia ­from seven localities in the Broken River Group ofnortheastern Queensland - from the late Emsian(serotinus Zone), late Eifelian (kockelianus Zone),various levels in the Givetian, and one possiblyearliest Frasnian (Figure 1) All come from localities(see Appendix) in the Broken River Group on theWandoVale and Burges 1:100,000 topographicsheets south of Greenvale, in the Townsvillehinterland of northeastern Australia.

STRATIGRAPHIC CONTEXT

The late Emsian-early Frasnian Broken RiverGroup (Mawson and Talent 1989; Sloan et al. 1995;Withnall et al. 1992, 1993) cropping out widely inthe Broken River region (Figure 1) can be viewed asconsisting of two platform areas with shallow-watersequences, the Dosey-Craigie and PandanusPlatforn1s, in the south and northwest respectively(Mawson and Talent 1989; Sloan et al. 1995),separated by a northeasterly directed tract of deeperwater sediments, the Burges Submarine Valley

(compare Figures 1 and 2). Sediments in the latterconsist of a stratigraphically cryptic complex ofmudrocks and subordinate arenites with debrisflows, olistoliths and carbonate fans. Thesedimentary complex in the Burges SubmarineValley has been referred to as the Burges Formation(Withnall et al. 1988, 1993: 116-122) but, largelybecause of lack of stratigraphic differentation, hasbeen referred to in subsequent publications, as wellas herein, as. undifferentiated Broken River Group(Sloan et al. 1995). The debris flows and carbonatefans have been built out into the Burges SubmarineValley primarily from the Dosey-Craigie Platform.The best material has come from these carbonatefans.

TRILOBITE FAUNAS

Most of the material is inadequate, either becauseof poor preservation or because of an insufficientnumber of exoskeletal pieces with characteristicsthat would allow specific or even genericattribution. With the exception of a single phacopidfrom a siliceous nodule in late Emsian shales(Bracteata Formation) and a faunule from ?earlyFrasnian but possibly latest Givetian decalcifiedmudstone (Mytton Formation), the Broken Rivertrilobites are from bioclastic crinoidal limestonesand marlstones in the above-mentioned carbonatefans. The lithologies include algae indicating ashallow water context (or shallow water derivation)

66 R. Feist, J.A. Talent

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in which trilobites tend normally to be extremelyrare. Interestingly, the identifiable part of thecollections consists entirely of new taxa. Ourcollections are very poor in individuals; they aremostly present - especially in the case of theEifelian ones - as detached, often fragmented piecesof carapace accumulated in downslope-transporteddebris-flows derived from presumably reefal

limestones. The Givetian brachiopod- and coral-richmarlstones have produced a few entire phacopids,typical of quiet-water, muddy habitats. All thetrilobites have rather large, protruding eyes ­typical for shallow platform environments.

Two associations were encountered in the lateEifelian (kockelianus Zone) where four families arerepresented (phacopids, proetids, tropidocoryphids

Devonian trilobites from northeastern Australia 67

SILURIAN DEVONIAN

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Figure 2 Schematic north-south cross-section of the Broken River area of northeastern Australia, from the Dosey­Craigie Platform in the south to the Pandanus Platform in the north, showing Ludlow-Frasnian units andrelative stratigraphic position of localities from which late Emsian-?early Frasnian trilobites documentedhere have been obtained (modified from Sloan et al. 1995), Note the southern and northern carbonateplatforms and the intervening submarine valley (Burgess Submarine Valley) with undifferentited BrokenRiver clastics with carbonate fans (at localities 3 and 4), debris flows and olistoliths, Note also that theStanley Limestone Member is shown as extending very slightly above the Givetian-Frasnian boundary onthe basis of meagre, though not compelling conodont data, and that locality 6, low in the Mytton Formation,is separated from the Stanley Limestone Member by a break in sedimentation of uncertain duration,

68

and aulacopleurids), two associations in the early tomid-Givetian Papilio Formation (phacopids andscutelluids), and one in the Mytton Formation (withphacopids and aulacopleurids). Due to thestratigraphic position and the depositionalenvironment, the fauna is different in compositionfrom all previously described trilobite faunas fromAustralia. Preliminary palaeobiogeographicalcomparisons reveal only mild affinity with typicalNorth Gondwanan faunas from Bohemia,southwestern Europe or North Africa. The scutelluidseems to be closer related to Avalonian, especiallyRhenish forms, whereas the phacopids and the proetidsare unusual; they appear to have some traits incommon with Mongolian taxa. More material is neededto complete knowledge of the identified taxa and toenable specific assignment of the several taxa for whichspecific and even generic location is problematic.

SYSTEMATIC PALAEONTOLOGY

Family Styginidae Vogdes, 1890

Subfamily Scutelluinae Richter and Richter, 1955

Genus Scutellum Pusch, 1833

Type speciesScutellum costatum Pusch, 1833

Scutellum tenuistriatum n. sp.Figure 3C

NameReferring to the fine, striated sculpture covering

the surface of the exoskeleton.

Type materialHolotype, AMF 112492, a partly exfoliated

pygidium from locality 8; Figure 3C. Paratype:pygidium, AMFl12493 from locality 2. Age: lateEifelian (kocke1ianus Zone) to possibly earliestGivetian (hemiansatus Zone).

OccurrenceBrachiopod-rich mudstones of the Papilio

Formation; late Eifelian (kockelianus Zone) topossibly earliest Givetian (hemiansatus Zone)

DiagnosisA species of Scutellum Pusch with pygidium

characterised by having a median rib narrowest inits proximal third, expanding towards the axis tobecome wider than median axial lobe; pleural ribshaving proximal ends bent outwards; ribs flatteneddistally, sharply defined by narrow pleural furrows.Prosopon of dense ridges and sparse nodules;border pseudospinose.

R. Feist, l.A. Talent

DescriptionPygidium moderately vaulted, elongate

pentagonal in outline (width to length == 2.9:2.6)with, in lateral view, the proximal third of thepleural field flattened with a slightly depressedmedian rib and a moderately elevated small axisdeclining forward. Pleural field vaultedmedially, with a markedly flexed median rib;outer part gently sloping to broad borderdepression; border flat and horizontal. Axistriangular in dorsal view, sharply separatedfrom the pleural field by a continuous dorsalfurrow, and with narrow median lobe notreaching the dorsal furrow. Doublure two-thirdsthe length of the pleural field (sagitally). Sevenpairs of pleural ribs and a broader unbifurcatedmedian rib, all cylindrical in their adaxial third,flattened to the rear, reaching the posterolateralborders, sharply defined by narrow and deepinterpleural furrows. Median rib, twice as broadas the adjacent ribs at its posterior end, taperscontinuously to the anterior border of thedoublure, at which point it expands again toreach a maximum width at the junction with thedorsal furrow where it is wider than the medianlobe of the axis. In the proximal third of thepleural field all pleurae (except the first one) arestrongly bent forward and outward to meet thedorsal furrow in a rather narrow angle. Theentire surface of the exoskeleton is denselycovered with thin, transversely directed,anastomosing ridges of equal size that, at thejunction with the borders, curve abruptly backin the form of minute hooks that terminateobliquely at the border in a succession of parallelchevrons giving to it a pseudospinose aspect("Pseudoziihnelung" of Erben 1967). In additionto this sculpture sparse nodules of moderate sizeare displayed throughout the surface.

RemarksIn her revision of the genus Scutellum, Archinal

(1994) grouped all taxa with a long pentagonaloutline of the pygidium and an adaxially wideningmedian rib into a new subgenus Scutellum(Calycoscutellum). Because of this feature, the newspecies could be located in S. (Calycoscutellum), butthe diagnostic features and especially thewidening of the adaxial median rib - seem not to besufficiently expressed to allow unequivocalcharacterisation of that subgenus. Moreover, theadaxial portion of the median rib widens during thepostlarval ontogeny; this feature cannot be seen insmaller specimens as for example in S. depressumCooper and Cloud (1938). There is great variabilityof this feature in different morphotypes of S.flabellifer (the type-species of Calycoscutellum) as wellas in other taxa such as S. ("Calycoscutellum") sagittaArchinal1994. As stated by Basse (1996), additional

..._--------------

Devonian trilobites from northeastern Australia

diagnostic features must be identified before thissubgenus can be usefully employed.

Among Givetian species of SClltellunl with aproximally widening median rib and narrowinterpleural furrows the new species ressembles S.bnmopalllusi Archinal from the Eifel Mountains,Germany, but has a much stronger outwardcurvature of the proximal pleurae. It differs from S.delicatunl (Whidbome) from the Lummaton shellbedin Torquay (SW England) by the markedconstriction of the median rib at the anterior borderof the doublure. It resembles S. depressunl Cooperand Cloud from Illinois, but has a much wideranterior termination of the median rib and strongercurved anterior pleurae. Moreover the border of theAmerican species carries very tiny short spines seenonly in large specimens; such true spines are notdeveloped in S. tenuistriatunl. Additionally, thecharacteristic prosopon makes the new speciesdistinct from all other comparable species ofScutellunl.

All previously described Australianrepresentatives of Scutellum come from the EarlyDevonian. S. calvunl Chatterton 1971 and S. hollandiWright and Chatterton 1988 are both from theEmsian of New South Wales. S. droseron Hollowayand Neil 1982 and S. slldorum Holloway 1996 arefrom the Lochkovian and early Emsian respectivelyof Victoria. All four forms may be discriminatedfrom the present species by their prosopon and theconfiguration of the median rib.

Little is known about Givetian forms of SClltcllunlfrom Gondwana-related areas. A single cranidiumof Scutellum sp. 2 gr. costatum has been found inBrittany (Morzadec 1983). The exoskeleton lacksthe characteristic ridges developed in S.tcnuistriatum. A poorly preserved pygidium fromCantabrian Mountains assigned to s. costatum cf.lummatonensis Selwood by Smeenk (1983) hasmuch larger interpleural furrows than the newspecies. An undescribed pygidium of s. costatumcostatum occurs in Stringocephalus limestones atMonumenz in the Central Camic Alps (collectionsof G.B. Vai, number 23925, Bologna). Otherrepresentatives of the costatum group in theMontagne Noire, all with a narrow anteriormedian rib, do not occur earlier than earliestFrasnian (Feist 1974).

Scutelluid indet.

RemarksTwo fragments of a scutelluid cranidium and a

few thoracic segments (AMFI12494) occur inbioclastic, crinoidal and brachiopod-rich spariticlimestones at locality 4 (latest Emsian patulus Zoneor earliest Eifelian, partitlls Zone). They areassociated with Burgesina mawsonac n. gen. n. sp.

69

Family Proetidae Salter, 1864

Subfamily Proetinae Salter, 1864

Genus Proetlls Steininger, 1831

Subgenus Proetlls (Devonoproetlls) Liitke, 1990

Type speciesProetus (Proetus) talenti Chatterton, 1971

RemarksLutke (1990) placed Ocvonoproetus in Gerastos

Goldfuss 1843 despite the close resemblance of itstype species P. talenti to Proetus s.s. (Adrain 1997).Accordingly and because of the direct ancestor­descendant relationship between both taxa weconsider Oevonoproetus as a subgenus of Proetus.

Proetlls (Devonoproetlls) sparsinodoslls n. sp.Figure 3A, E, I, M, N

NameRecalling the sporadic occurrence of small nodes

on the smooth surface of exoskeleton.

Type materialHolotype, cranidium, AMF 112495, from locality

3,216 m above the base of section Br BB, Figure 3/A, E, 1. Paratypes from same locality and stratum:juvenile cranidium, AMF 112496 ; fragmentarypygidium, AMF 112497, Figure 3/M, N;fragmentary pygidium, AMF 112498. Age: lateEifelian (zone uncertain).

OccurrenceBlack crinoidal wackestone (Eifelian; kockelianus

Zone) from a carbonate fan in undifferentiatedBroken River Group in the Burges SubmarineValley.

DiagnosisSpecies of the subgenus Oevonoproetus

characterised by: cranidium with vaulted, longsubrectangular glabella reaching poorly definedanterior border; lateral occipital lobes unswollen,incompletely separated from occipital lobe; sparsetubercles on posterior part of glabella. Pygidiumwith long and high axis reaching to poorly defined,swollen margin; ring and pleural furrows veryweakly impressed.

DescriptionThe elongate subrectangular, parallel sided

glabella is defined by straight dorsal furrows and asemicircular frontal outline. It is continuously butnot very highly vaulted both in lateral and frontalviews with a hemispherical shaped frontal lobe. SIand S2 furrows, weakly impressed, separate

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70 R. Feist, J.A. Talent

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Devonian trilobites from northeastern Australia

unswollen lateral lobes. Basal angles of glabellabehind L1 are slightly truncated by distally forwarddirected occipital furrow. Dorsal furrows, deepbehind and in front of glabella, are shallowanterolaterally. Anterior border broad, slightlyswollen, poorly defined by large border depressionthat merges with prefrontal furrow of glabella,protruding in dorsal view; broadly downcurvingmargin provided with continuous terrace ridges.Occipital ring narrow, strongly vaulted (sag.), ofequal width as the base of glabella, exhibitsincompletely separated, low, uninflated laterallobes of moderate size that are bounded anteriorlyby the deeply incised distal ends of the occipitalfurrow. Fixigenae adjacent to glabella extremelynarrow and equal in width anteriorly andposteriorly of the palpebral lobe. Facial suturebetween b-g and e--z runs parallel to sagittal line.Palpebral lobe of sigmoidal outline, vaultedtranversely, two and a half times longer than wide,situated inmidst between the anterior borderdepression and the posterior border furrow.Prosopon smooth with the exception ofcharacteristic, relatively big-sized nodules that aredistantly dispersed on the posterior two thirds ofthe glabella. Pygidium, largely semielliptical inoutline, has a long, relatively high axis defined bystraight, little converging dorsal furrows and abroadly rounded, elevated end that reaches to thepoorly defined border depression. There are 7 + 1flat axial rings of which the anteriormost isnarrower and more elevated than the followingones; ring furrows sigmoidal with a distallyforward curving swing; only the two anterior ringfurrows are deeply marked, the others are veryfaint. The pleural field is moderately vaulted with apoorly defined border depression and a largeslightly swollen border. There are 4 or 5 flat pleuralribs that are separated by faint pleural furrows.Pleural ribs and furrows terminate at borderdepression far inside from posterolateral margin.Posterior border slightly up-tilted with narrow,swollen margin. Prosopon smooth; only the anteriorring of axis with a few nodules.

71

RemarksThe type species of Proetlls (Oevonoproetlls) is P.

(D.) talenti Chatterton from the late Emsian TaemasFormation, New South Wales. This species isdifferent in exhibiting dense tuberculation, morediverging anterior sutures, a larger pygidial axiswith throughout deeply impressed ring furrows.The new species is closest to Proetlls vesclIs Zhou,Siveter and Owens from the Eifelian YikewusuFormation of Inner Mongolia. It is distinct by thebroader, somewhat flatter anterior margin and thesparse nodular sculpture on the cephalon. TheEmsian and early Eifelian species P. (D.) margoLutke and P. (0.) clIltrijllgati Richter and Richterfrom Germany are clearly distinct from P. (D.)

sparsinodoslIs by the outline of their more taperingglabella and the curved dorsal furrows. Incomparison to these species P. (D.) sparsinodosus hasa longer rachis with a broader end-piece (sag.).

According to Lutke (1990) the subgenusOevolloproetus is represented in Avalonia-Baltica andNorth America whereas, besides Australia andMongolia, its occurrence in Northern Gondwana(Bohemia and Armorica) is questionable. By contrast,the contemporaneous proetine Gerastos is wellrepresented in western North Gondwana terranessuch as Armorica, North Africa and Bohemia.

Proetus (Devonoproetus) latimargo n. sp.Figures 3B, G, K 0, V, W, X.

NameReferring to the large, characteristic border of the

pygidium.

Type materialHolotype, cranidium, AMFl12499, from locality

3, 214.5 m along a metric tape from the base of thesection (Figures 3B, G, K). Paratypes from the samebed: fragment of librigena, AMFl12500;fragmentary pygidium, AMF 112501, Figures 30, X;pygidium with fragmentary axis, AMFl12502,Figures 3V, W.

~ Figure 3 Trilobites from the Broken River Group, N Queensland. The material is housed in the collections of theAustralian Museum, Sydney (AM). All figured specimens were coated with MgO. A, E, I, M, N, Proetzls(Oevonoproetlls) sparsinodoslIs n. sp. A, E, I, holotype, cranidium, AMF112495, dorsal view (A) x 4.4; frontalview (E) x 4.9; lateral view (I) x 4.4; M, N, pygidium, AMFl12497, fragmentary, lateral view (M) x 5.4; dorsalview (N) x 5.4. B, F, G, H, J, K, O,T, V,W, X, Proetlls (Oevonoproetlls) /atimargo n. sp. B, G, K, holotype,cranidium, AMFl12499, dorsal view (8) x 5.4; frontal view (G) x 5.7; lateral view (K) x 5.2; F, J, T, cranidium,AMF112503, frontal view (F) x 6.5; lateral view (J) x 5.7; dorsal view (T) x 6; H, librigena, AMF112504, dorsalview x 7.4. 0, X, pygidium, AMF 112501, fragmentary, dorsal view (0) x 7.1; lateral view (X) x 7.1. V, W,pygidium, AMF112502, axis broken, dorsal view (V) x 6.4; lateral view (W) x 6.4. C, Selltellllm temlistriatllm n.sp, holotype, pygidium, AMF 112492, partly exfoliated, dorsal view x 1.4. D, Otarion sp. cranidium withbroken glabella, AMFl12534, dorsal view x 8.7. L, P, Astyeoryphe sp. L, librigena, AMFl12529, dorsal view x8.3; P, fragment of cranidium, AMFl12528, dorsal view x 8.8. Q, S, Proetinae gen. et sp. indet., fragmentarypygidium, Q, AMFl12526, lateral view x 6; S, dorsal view x 5.7. R, Cornuproetinae gen. et sp. indet.,fragmentary anterior portion of cranidium, AMF112527, dorsal view x 5. U, Y, AlI/aeop/ellm sp., U, librigena,AMF112533, internal mould, dorsal view x 6.8; Y, cranidium, AMF112532, internal mould, dorsal view, x 7.7.

72

OccurrenceBlack crinoidal wackestone from undifferentiated

Broken River Formation, late Eifelian ?australisZone. Supplementary material assigned to samespecies from locality 4, crinoidal brachiopod spariticlimestone also from undifferentiated Broken RiverFormation: cranidium, AMF 112503, Figure 3/T, F,J; librigena, AMF 112504, Figure 3/H.

DiagnosisSpecies of the subgenus Devonoproetus with

following characteristic features: cranidium withlong tongue-shaped glabella, not overhanginganterior border; border thick, protruding, not up­tilted; genal spines short. Pygidium with large axis,markedly tapering to the rear, with deeply markedaxial ring furrows; margin very large, defined byshallow border furrow.

DescriptionThe cranidium exhibits a tongue shaped glabella,

bounded by slowly but continuously converging,straight dorsal furrows of equal depth. The glabellais strongly vaulted (tr. and sag.), highly arched infrontal view, and is longer than wide (5.5 : 4.5),broadest at base. There is no praeglabella field.Protruding anteriorly, the large, subcylindrical(sag.) border is defined by a marked border furrowthat deepens in front of glabella when merging withthe prefontal furrow. Front lobe of glabella notoverhanging but slightly impinging on border.Glabella furrows inconspicuous; 51 and 52 slightlyimpressed adjacent to dorsal furrow. Occipitalfurrow straight, deepening behind L1 and meetingdorsal furrows at right angles. Occipital ring narrow(sag.), as broad as glabella (tr.), anteriorlysubdivided by deep, discontinuous incisions ofoccipital furrow that isolate elongate, anteriorlyswollen lateral lobes; median lobe protrudingforward. Anterior fixigenae large, vaulted, downsloping. Anterior branches of suture first parallel tothe dorsal furrow then flexing markedly outward. gand e are situated very near to dorsal furrow.Narrow sigmoidal shaped, strongly inclined, smallpalpebral lobes remain behind 52. Posteriorbranches of suture parallel to dorsal furrow. Thelibrigena is provided with a rather robust,cylindrical border, a large eye with a weak eye-socleand a narrow anterior librigenal field. The genalangle protrudes into a small spine. The pygidium islargely parabolic in outline with slightly truncatedposterior margin. Axis robust, high, larger than onepleural region, defined by straight, markedlyconverging dorsal furrows and a truncatedposterior outline. It is provided with 7 + 1prominent, cylindrical rings separated by ratherdeep, slightly sigmoidal ring furrows. Pleural fieldlarge, moderately vaulted, subdivided into a morevaulted inner part with well defined pleural ribs

R. Feist, I.A. Talent

and a very large gently sloping outer marginseparated by a shallow but distinct border furrow.There are 5 - 6 bicomposite ribs defined by deeplyincised pleural furrows. Interpleural furrows areless marked distally. Only the pleural bands of thetwo anterior ribs continue onto the outer margin;the posterior ones terminate at border furrow. Theouter slope of the margin carries two or threeterrace ridges. Prosopon tuberculate. On glabellatubercles increase in size from front to rear.

RemarksThe new species is distinct from P. (DJ talenti, P.

(DJ sparsinodosus and P. (DJ vescus by its tongue­shaped glabella; it shares this feature with Proetuszhusilengensis Zhou, Siveter and Owens from theEmsian Zhusileng Formation of Inner Mongolia;however it is distinct by its much larger pygidialborder and denser nodular sculpture.

Genus Burgesina n. gen.

Type speciesBurgesina mawsonae n. sp.

NameAfter Burges 1:100,000 topographic sheet,

Townsville hinterland, of northeastern Australia,where the trilobites were found.

DiagnosisA genus of the subfamily Proetinae with the

following characteristic features: Cranidium largewith up-tilted border ridge; glabellasubquadrangular, flattened with backwardprotruding basal lobes; lateral occipital lobes high;librigena with prominent eye, lacking genal spines.Pygidium semicircular with thick border; axis long,prominent, larger than pleural region, with 9 + 1rings. Sculpture of coarse discontinuous ridges andtubercles.

SpeciesB. formosa (Zhou, Siveter and Owens 2000), B. sp.

indel. 1 (Zhou, Siveter and Owens 2000), B. sp.indet 2 (Zhou, Siveter and Owens 2000), B.mawsonae n.sp.

Time and occurrenceEmsian to Eifelian; Western Inner Mongolia and

Broken River Region, North Queensland

RemarksAmong the proetines with pronounced lateral

occipital lobes the new taxon shares features of thecranidium such as the up-tilted anterior border andthe shape of the glabella with some Americanspecies of Crassiproetus Stumm such as C. sibleyensis

Devonian trilobites from northeastern Australia

and C. nonooodensis (cf. Lieberrnan 1994, I~igure 20).But the particular trait of backward projected basallobes does not occur in Crassiproetlls. Besides, thenew taxon has a much shorter and less archedpygidial axis with fewer rings. In this regard, thepygidium is nearer to that of Coniproetlls Albertithough it has a much thicker border (Alberti 1966).The Silurian genus Hedstroemia Pribyl and Vanek1978 to which Zhou, Siveter and Owens originallyattributed the Mongolian taxa is characterised byits anteriorly tapering glabella, the presence of apreglabellar field, and the deeply incised lateralglabeIlar furrows. The new genus is distinct fromHedstroemia by characteristic features such asabsence of the praeglabellar field, uptilted ridge­like anterior border, faint lateral glabellar furrows,very broad and inflated pygidial border, fewer axialrings and straight pleural bands of even height andlength. In the outline of the large quadrangularglabella and in the shape of its posterior border, thenew taxon differs considerably from all otherproetines.

Burgesina mawsonae n. sp.

NameFor Prof. Ruth Mawson who has contributed so

importantly to improving knowledge of thechronologic framework for the Broken River andother mid-Palaeozoic sequences in Australia.

DiagnosisType species of Burgesina n.g. characterised by:

quadrangular, highly vaulted glabella with notchedbase and elongated lateral occipital lobes.

Type materialHolotype, cranidium, AMF112505, from locality

4, Figure 4/ A, U. Para types from same bed:cranidium, AMF112506, Figure 4/ F, R ; fragmentof anterior cranidium, AMF112507, Figure 4/ B;librigena, AMFl12508, Figure 4/ M; pygidium,AMF112509, Figure 4/ I, T; pygidium, AMF11251O,Figure 4/ K, 0, S; pygidium, AMFl12511, Figure 4/N. Supplementary material from same bed: 5cranidia (AMFI12512-516), 2 librigenae(AMF1l2517-518), 2 thoracic segments(AMF112519-520), 5 pygidia (AMF1l2521-525).

OccurrenceBioclastic, crinoidal and brachiopod rich sparitic

limestones at the base of Fish Hill, locality 4 (latestEmsian patllllls Zone or earliest Eifelian, partitllsZone).

DescriptionCranidium square-shaped with large,

subquadrangular glabella, slightly broader at base

73

than in sagittal length, nearly parallel-sided,tapering in front, strongly vaulted anteriorly (sag.),less transversely, flattened behind; frontal lobesemicircular, vertically sloping, not overhanging.Glabellar furrows unimpressed, large, sculpturless;S 1 broad, backward directed without reaching tooccipital furrow; lateral lobes unswollen. Base ofglabella notched by inserts of occipital furrow andmarkedly backward shifted median and laterallobes (L 1). Dorsal furrows narrow, deep, nearlyparallel behind, thinning and strongly convergentin anterior third. Rather narrow, deeply incisedanterior border furrow merges with prefrontalfurrow of glabella. Anterior border ridge-likeuptilted, with vertical, slightly vaulted outer slope,steeply inclined posteriorly, forming a narrow crestseparated from front of glabella by deep, gorge-likeborder furrow. Occipital furrow homogenouslydeep and narrow, sigmoidal, bifurcated distally.Occipital ring as high and broad (tr.) as base ofglabella, flattened at top, slightly protrudingmedially, with completely separated, prominent,elongated lateral lobes. Fixigenae moderately large(tr.), arched anteriorly to the same degree asglabella. Palpebral lobe short (exsag.), rearwardpositioned, of narrow sigmoidal outline, slightlyflexed inward and backward. e lies nearer to dorsalfurrow than g. Anterior branches of suture long,straight, little diverging until border furrow,strongly inward curved on border. Short portion ofposterior suture e-z straight and parallel to dorsalfurrow. z lies in posterior border furrow opposite tothe posterior branch of occipital furrow. Posteriorborder cylindrical, short and bent backward.Librigena subtriangular with prominent eye onhigh, unswollen eye-socle that merges continuouslywith slightly arched and deeply downslopinglibrigenal field. Border furrow sharp and deeplyimpressed. Lateral border broad, as large (tr.) asgenal field, twice as large as posterior border,flattened at top with thick outer margin. Genalangle without spine; short residual spine may stillbe present in young holaspids. Prosopon ofcephalon: coarse wrinkles and discontinuous ridgeson glabella and occipital ring, discontinuousparallel terrace ridges on anterior fixigenae,continuous terrace ridges on the outer slopes of theanterolateral borders, minute grooves in borderfurrows and on eye-socle. Pygidium semicircularwith thick border separated by continuouslymarked border furrow that interrupts the vault ofthe pleural field. Axis long, highly archedtransversely, anteriorly broader (tr) and, in lateralview, as high as the pleural region, tapers withcontinuously straight dorsal furrows and terminateswith a high, rounded end shortly before reaching atthe border furrow. There are 9 + 1 slightlybackward inclined axial rings, completely separatedby thin ring furrows that tend to vanish rearwards

74

(especially in larger specimens). Pleural fieldstrongly arched and sharply bounded by borderfurrow. There are 8 conspicious, bicomposite,prominent pleural ribs that all abruptly terminatein border furrow. Pleural bands of equal heighth,the posterior slightly broader than the anterior; theanterior ones clearly separated by faint butcontinuous interpleural furrows. Border swollen, ofequal breadth, slowly downsloping from borderfurrow outwards, in lateral view as high as pleuralfield. Outer slope carries 6 continuous, parallelterrace ridges. Prosopon: tubercles on each pleuralband and on the posterior axial rings.

ComparisonThe Mongolian representatives of the new genus

share many traits with B. mawsonae from the BrokenRiver region. In particular, B. formosa Zhou, Siveterand Owens from the Emsian Zhusileng Formationis very close to the new taxon. Whereas the pygidiaare nearly identical, the cranidium of B. mawsonae isdistinct in its more quadrangular, stronger andmore highly vaulted glabella with its deeplynotched base, the elongated lateral occipital lobes,and the broader fixigenal fields with posteriorsutures that run parallel to the dorsal furrows. Bothtaxa doubtless belong to the same genus; they aredistinct from Hedstroemia by characteristic featuressuch as absence of the praeglabellar field, uptiltedridge-like anterior border, faint lateral glabellarfurrows, very broad and inflated pygidial border,fewer axial rings and straight pleural bands of evenheight and length. The combination of thesefeatures justifies integration of the two species intoa separate genus, i.e. Burgesina n.g.

Proetinae gen. et sp. indet.Figure 3Q, S

RemarksA undetermined fragmentary pygidium, AMF

112526, from locality 4, occurring in associationwith Proetus (Devonoproetus) latimargo. It may,however, belong to another species because theaxial rings are not completely separated, and thering furrows are displayed only in the centre of theaxis. Effaced axial furrows occur on pygidia ofOrbitoproetus Pillet 1969, but representatives of thatgenus have a broader and less elevated axis.

Subfamily Comuproetinae Richter and Richter,1956

Comuproetinae gen. et sp. indet.Figure 3R

RemarksAn undetermined fragment of the anterior of a

-------

R. Feist, J.A. Talent

cranidium, AMF 112527, from locality 3, 216 malong a metric tape from the base of section Br BB,late Eifelian (?australis Zone). The fragment isassigned to the Cornuproetinae because of theconfiguration of the broad anterior glabella lobe, therather narrow preglabellar field, the deep, flexedanterior border furrow, and the broad cylindricalborder protruding slightly forward adaxially, andits continuous terrace lines.

Family Tropidocoryphidae Pribyl, 1946

Subfamily Tropidocoryphinae Pribyl, 1946

Genus Astycoryphe Richter and Richter, 1919

Type speciesAstycoryphe senckenbergiana Richter and Richter,

1919

?Astycoryphe sp.Figure 3L, P

RemarksFragments of tropidocoryphine trilobites have

been obtained from locality 3, at 214.5 m along ametric tape from the base of the section Br BB, lateEifelian (?australis Zone). Figured are onecranidium, AMF 112528, Figure 3P and onelibrigena, AMF 112529, Figure 3L. Supplementarymaterial: one fragment of the anterior of acranidium, AMF 112530, and one librigena,AMF112531.

The material is not sufficiently complete forspecific or even generic attribution. Because of thelarge frontal lobe of the glabella and thecharacteristic lateral border it is tentatively assignedto Astycoryphe. This genus has been reported fromthe Early Devonian of New South Wales (Wrightand Chatterton 1988).

Family Aulacopleuridae Angelin, 1854

Subfamily Aulacopleurinae Angelin, 1854

Genus Aulacopleura Hawle and Corda, 1847

Type speciesArethusina konincki Barrande 1846

Aulacopleura sp.Figure 3D, Y

RemarksA single cranidium, AMF 112532, Figure 3Y and a

free cheek, AMF 112533, Figure 3D have beenrecovered from a small decalcified marlstonenodule at locality 6 in the Mytton Formation in the

Devonian trilobites from northeastern Australia

Page Creek area. The specifically indeterminablematerial exhibits characteristic features ofAlllacopleura such as the rectangular, anteriorlytruncated, low frontal lobe of the glabella, the large,posteriorly projected praeglabella field and theprominent eye-ridges. However, the librigenaressembles those of the genus Cyplzaspis Burmeister1843. If the presence of Alllacopleura can beconfirmed by the discovery of more complete andbetter preserved material a pre-Taghanic age couldbe assigned to the nodule as Alllacoplellra has beenregarded as disappearing at the Middle/Uppervarcus Zone boundary. The matrix, however, isargued (see earlier) to be younger than this, the bestestimate on available data being earliest Frasnianrather than late Givetian.

Subfamily Otarioninae Richter and Richter, 1926

Genus Otarion Zenker, 1833

Type speciesOtarion diffractum Zenker, 1833

Otarion sp.Figure 3D

RemarksA fragment of a cranidium assigned to Otarion,

AMF112534, occurs in association with Burgesina inbrachiopod-rich, crinoidal sparites at locality 4(latest Emsian patulus Zone or earliest Eifelianpartitus Zone). The specimen exhibits a row of 6spines of equal length on the anterior border. Thisfeature occurs in some otarionines such as O.davidsoni Barrande 1852 and was earlier considereddiagnostic of Otarionella Weyer 1965; it is nowconsidered to be a paedomorphic feature (Adrainand Chatterton 1994), and Otarionella is accordinglyviewed as a younger synonym of Otarion.

Phacopidae Hawle and Corda, 1847

Phacopinae Hawle and Corda, 1847

Phacops Emmrich, 1839

Phacops (Phacops) Emmrich, 1839

Type SpeciesCalymmene latifrons Bronn, 1825

Phacops ( Phacops ) brocki n. sp.Figures 4H, L, P, Q

NameFor Or Glenn Brock of Macquarie University

Centre for Ecostratigraphy and Palaeobiology

75

recognising his work in the Broken Riverarea.

Type materialHolotype, AMF112535, enrolled entire specimen

from locality 1, Figure 4P, Q. Paratype: cephalon,AMF112536, from locality 5, approximately halfway between 146 and 147 m above the base of thesection, Figure 4H, L. Supplementary material:enrolled thoraco-pygidium, AMF112537, fromlocality 1; 1 fragmentary cephalon, AMF112538,rom locality 8 at 223 m from the base of the secton; cephalon, AMF112539, from locality 5, approx­imately half way between 150 and 165 m above thebase of the section; cephalon, AMF112540, fromlocality 4 same horizon as the holotype.

OccurrenceFrom brachiopod-rich mudstones in the Papilio

Formation-late Givetian, Izermanni Zone, about 51­52 m above the entry of Sclmzidtognathus (see Sloanet al. 1995, Table 6). Because of rareness, the firstrecord of Schmidtognathus in this section may beappreciably above the base of the hermanni Zone,and conceivably younger in the late Givetian.

DiagnosisA species of Phacops (Plzacops) characterised by

having: cephalon low, flattened; glabella notoverhanging, with slightly convex dorsal furrows;preoccipital ring elongate, large; occipital ringvery narrow, not protruding medially; eyes verylarge and high, reaching to the posterior borderfurrow. Pygidium with narrow, high axis; axialring and pleural furrows deep.

DescriptionCephalon with a broad subtrapezoidal, low,

flattened glabella with a vertically dipping anteriorlobe that does not overhang but covers the narrow,medially withdrawn anterior border (in dorsalview). Glabella defined laterally by slightlyoutcurved dorsal furrows. Anterior and posteriorcorners of glabella blunt. SI continuously marked,curved forward and shallowing medially.Preoccipital ring well developed, exhibiting anelevated, elongate median lobe protrudingforwards medially, and low, small, moderatelyswollen lateral lobes. Occipital furrow straight, ofeven depth. Occipital ring rather narrow, straght, prominent, as broad (sagitally) as the preoccipial ring, not protruding medially, as high as the g­labella. Eyes very large, extending to posterior bor­der furrow, as high as the posterior glabella.Palpebral lobe narrow, horizontal, archedtraversely, with shallow, curved palpebral furrow.Visual surface steeply inclined, kidney-shaped with17 dorsoventral files of eye-lenses, with a maximumnumber of 10 lenses in a row in large specimens (for

76 R. Feist, J.A. Talent

)

Iu

M

E

c

H

L

J

p

T

G

o

B

• ',~ '1l-~-:"_._~"'.

.~~.~~,-... ~ -':~';'''', ~'-",. ~ . ~,( "-

":;;...~,...~,~. 1"",1~, ,....,,-~" '''---' ~ .~. ' .. ..-' ....~'.: ,~ . ~' ';

o

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r,....·.... ,..I' •

,~

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Devonian trilobites from northeastern Australia

example, from front to rear, 7899 10 9 10 9 9 9 9 87 6 5 4 2), yielding a total of 130 lenses. Lens-filesstart shortly above base of eye which is surroundedby a large, shallow depression. Posterior genal fieldswollen between palpebral furrow and dorsalfurrow. Outer genal field narrow, uninflated,moderately down-sloping; genal corners blunt.Vincular furrow very shallow and large medially.Prosopon consisting of low tubercles of mediumsize on glabella and occipital lobe. Pygidiumsemielliptical in posterior outline, with axis ratherhigh and narrow, a little broader than half of onepleural region. Dorsal furrows straight, convergingvery gradually; end of axis elevated, higher thanpleural regions. Axial rings 8 + 1, narrow, elevated,separated by straight, deep ring furrows. Pleuralregion homogenously arched, with 5-6 pleural ribsclearly defined by deep pleural furrows. Prosopontuberculate.

RemarksThe combination of features such as low, non

protruding glabella, outwardly convex dorsalfurrows, very large, backwardly extending eyes,and narrow, medially non- protruding occipital lobeare characteristic of the new, first mid-Devonianphacopid reported from Australia. Europeanspecies from the Givetian with comparably largeeyes, such as Phacops (Phacops) sobolewi Kielan 1954or Pedinopariops simulator Basse 1998, haverectilinear diverging dorsal furrows and mediallyenlarged occipital rings. A similar configuration offlexed dorsal furrows, medially broadeningintercalating ring and narrow occipital ring isdeveloped in Toxophacops (Zhusilengops) ejinensisZhou and Campbell from the Emsian of westernInner Mongolia. However, this echinophacopidspecies differs from P. (P.) brocki in having a highlyarched anterior border and triangular pygidium.

Phacops sp.Figure 4C, E

RemarksAn exfoliated, fragmental cephalon of an

77

unidentified species of Plzacops, AMF1l2541, hasbeen recovered from a decalcified marlstonenodule, associated with AlIlacoplellra sp., at locality5 in the Mytton Formation in the Page Creek area(see earlier for discussion of age of matrix at thislocality).

Remarkable features of the specimen are the veryflat, evenly forwardly expanding glabella with arather low, dorso-ventrally flattened frontal lobe andbackwardly displaced, large eyes that are higher thanthe glabella. Conceivably it is from a new species,but is poorly preserved; additional material IS

necessary for adequate delineation of the species.

Phacopidae indet.Figure 4G, J

RemarksAn unidentified phacopid cephalon, was

recovered from a small siliceous nodule in lateEmsian age (serotinus Zone) shales outcropping inthe gully at locality 7. The specimen is preserved asan internal mould. Glabella very low and flat,narrow, subpentagonal, with a medially pointed,dorso-ventrally compressed frontal lobe thatstrongly overhangs the anterior border. Borderfurrow faint, retreated on ventral side behind thetip of the glabella. Vincular furrow shallow butcontinuously marked, strongly arched adaxially.Medially very high occipital lobe that is higher thanthe glabella. Preoccipital intercalating ring broadwith suppressed median lobe (on internal mouldthere is a large, gorge-like depression) and small,low lateral lobes. Very high eyes, backward shifted,reaching to the posterior border furrow, anteriorlyintruding into the dorsal furrows, slightlyimpinging on the anterior sides of glabella (oninternal mould only?). Visual surface steeplysloping, with 17 lens files with a maximum of 6lenses per file. Palpebral lobes reniform, horizontal,as high as the glabella. Genal angles pointed,probably with short spines.

RemarksThe general outline of the glabella, the ventrally

.... Figure 4 Trilobites from the Broken River Group, N Queensland. The material is housed in the collections of theAustralian Museum, Sydney (AM). All figured specimens were coated with Mgo. A, B, D, F, I, K, M, N, R-V mawsonae n.gen. et n. sp A, V, holotype, cranidium. AMF1l2505, dorsal view (A) x 5.9; lateralview x 5.9; B, cranidium, AMF1l2507, fragment of anterior part, dorsal view x 7; D, K, S, pygidium,AMF1l251O, posterior view (D) x 8; dorsal view (K):7.3; lateral view (S) x 8; F, R, cranidium, AMF1l2506,dorsal view (F): x 7.6; lateral view (R) x 7.6; I, T, pygidium, AMF1l2509, dorsal view (I) x 7.6; lateral view(T) x 7.4; M, librigena, AMF112508, partially exfoliated, dorsal view: x 6.3; N, pygidium, AMF1l2511, dorsalviex: x 78. C, E, Phacops sp., cranidium, AMF1l2541, internal mould, dorsal view (C) x 7.7; frontal view (E): x7.9. G,], Phacopidae gen. et sp. indet., cranidium, AMF1l2542, internal mould, dorsal view (G) x 5.1; frontalvIew (J) x 4.9. H, L, P, Q, Phacops (Phacops) brocki n. sp. H, L, cephalon, AMFl12536, partly exfoliated, lateralview (H) x 28; dorsal view (L) x 2.8; P, Q, holotype, entire enrolled specimen, AMFl12535, partly exfoliated;cer, 11:' Inn. dorsal view (Q) x 3; pygidium and posterior thoracic region, exfoliated, dorsal view (P) x 2.2.

78

situated anterior border furrow, and highposteriorly shifted eyes bring to mind species ofToxophacops (Atopophacops ) Zhou and Campbellfrom Inner Mongolia and especially T.(Atopophacops ) sp. indel. (Zhou and Campbe1l1990:PI. 5, Figure 12 ) which is approximately the sameage (late Emsian). However, representatives ofAtopophacops all have an elevated median lobe inthe preoccipital intercalating ring and have strongtuberculation. Both features are lacking in ourspecimen. Confident location in Atopophacopsrequires more and better material as well as apygidium.

ACKNOWLEDGEMENTS

Part of the trilobite material forming the basis ofthe present report was accumulated as a by-productof many years of field work in the Broken Riverregion focused on conodont- and brachiopod­biostratigraphy and palaeoecology funded byAustralian Research Council and MacquarieUniversity research grants to JAT and RuthMawson. Postgraduate students participated insome these field ventures and, with an eye fortrilobites, helped augment these collections. RF isgrateful to Macquarie University for a researchgrant enabling him to undertake field work withJAT and Ruth Mawson and other MUCEPcolleagues, particularly Brett Pyemont, Ken Bell andRoss Orury, focusing on a quest for trilobites. Thisconsiderably augmented the collections of thissparse component in the diverse faunas of theBroken River Group. We are indebted to RuthMawson for the conodont-based agedeterminations, Glenn Brock for sorting out thetrilobites in the voluminous Broken Rivercollections, Dean Oliver who made the drawings,Michel Pons (Montpellier) for printing thephotographs of the trilobites, and to our anonymousreviewers for helpful suggestions. This is acontribution to ISEM, UMR 5554 CNRS (pubI. 99­069) and to IGCP 421 North Gondwana mid-Palaeozoicbioventlbiogeography patterns in relation to crustaldynamics.

APPENDIX: LOCALITY REGISTER

Locality numbers refer to locations on Figure 1.LOCALITY 1. Uppermost Papilio Formation on

stratigraphic section S0210 at spot-samplelocality originally designated SO 43E. Thislocality (not shown on Mawson and Talent1989, Figure 5) is in Storm Dam Creek where itmeets the base of the Mytton Formation in theupper part of the widespread, approximately 1m calcareous interval referred to by Mawsonand Talent (1989) informally as the "Cinderellalimestone" .

R. Feist, J.A. Talent

Age: No older than late Givetian (hermanniZone, conceivably disparilis or norrisi Zones).Trilobites: Phacops (Phacops) brocki n. sp.

LOCALITY 2. Papilio Formation on stratigraphicsection SO 128 of Mawson and Talent (1989).Trilobites obtained during initial sampling ofthe section, labelled interval B-C, from 50-70m along a metric tape from the top of the OoseyLimestone.Age: Latest Eifelian (kockelianus Zone) topossibly earliest Givetian (hemiansatus Zone).Trilobites: Scutellum tenuistriatum n. sp.

LOCALITY 3. Undifferentiated Broken River Group onstratigraphic section Br BB of Sloan et al. (1995:Table 8, Text-fig. 8). Section commenced at thetop of the Shield Creek Formation and wasoriented to cross a prominent 13 m limestonelens (Sloan et al. (1995) interpreted as acarbonate fan extending from the Oosey­Craigee Platform into the Burges SubmarineValley.Age: Late Eifelian (?australis Zone).Trilobites:At 216 m: Proetus (Devonoproetus) sparsinodosusn. sp. Comuproetinae gen. et sp. indel., This isapproximately 2.3 m stratigraphically abovethe base of the fan.At 214.5 m: Astycoryphe sp., Phacops sp., Proetus(Devonoproetus) latimargo n. sp. This isapproximately 1 m above the base of the fan.

LOCALITY 4. Prominent carbonate interval in theundifferentiated Broken River Groupinterpreted as a carbonate fan extendingnorthwards into the Burgess Submarine Valley(referred to informally as the "Fish Hilllimestone") on stratigraphic section Br 15 ofSloan et al. (1995: Text-fig. 8, Tables lOA, lOB)at 348 m along a metric tape from the top of theShield Creek Formation - approximately 8 mabove the base of the "Fish Hill limestone".Age: Latest Emsian (patulus Zone) or earliestEifelian (partitus Zone).Trilobites: Burgesina mawsonae n. sp., Proetus(Devonoproetus) latimargo n. sp., Proetinae gen.et sp. indel., Otarion sp.

LOCALITY 5. Spanner Limestone Member of PapilioFormation on stratigraphic section SO 216 ofMawson and Talent (1989), cf. Sloan et al. (1995:Table 6). Trilobites obtained from 145-146 malong a metric tape (equivalent toapproximately 80 m thickness) from the base ofthe Spanner Member.Age: Late Givetian (hermanni Zone or possiblyslightly younger).Trilobites: Phacops (Phacops) brocki n. sp.

LOCALITY 6. Low Mytton Formation, 24.7 to 30 mabove the top of the Stanley Limestone Member

Devonian trilobites from northeastern Australia

of the Papilio Formation, same horizon asproducing charophytes (M. Feist et al. thisvolume).Age: Postdating entry of IcriodllS symmetricllsand thus more likely early Frasnian rather thanlatest Givetian, but caution has been suggested(Mawson and Talent 1997).Trilobites: Au/acop/eum sp., Phacops sp.

LOCALITY 7. Undifferentiated Broken River Groupmudrock in gully approximately 15 m east ofthe base of the "Fish Hill limestone" onsampled section Br 15 of Sloan et al. (1995).Age: Late Emsian (serotinlls Zone) - laterallyequivalent to part of the Bracteata Formationfarther south.Trilobites: Phacopidae indet.

LOCALITY 8. Papilio Formation on stratigraphicsection SO 130 of Mawson and Talent (1989).223 m and 243 m along a metric tape above thetop of the Oosey Limestone.Age: Not well constrained by conodont databut stratigraphic position accords with lateGivetian.Trilobites:At 223 m: Phacops (Phacops) brocki n. sp.At 243 m: Scutellum tenuistriatllm n. sp.

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Manuscript received March 1999; accepted December 1999.

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