An ichnofossil assemblage from the Lower Carboniferous, Snowy Plains Formation, Mansfield Basin,...

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An ichnofossil assemblage from the Lower Carboniferous SnowyPlains Formation, Mansfield Basin, Australia

Jillian M. Garvey a,b,⁎, Stephen T. Hasiotis c,d

a Department of Zoology, School of Life Sciences, La Trobe University, Melbourne, Victoria, 3086 Australiab Archaeology Program, School of Historical and European Studies, La Trobe University, Melbourne, Victoria, 3086 Australia

c Department of Geology, The University of Kansas 1475 Jayhawk Blvd., 120 Lindley Hall, Lawrence, KS 66045-7613, United Statesd Natural History Museum and Biodiversity Research Center, The University of Kansas 1475 Jayhawk Blvd.,

120 Lindley Hall, Lawrence, KS 66045-7613, United States

Received 27 November 2006; received in revised form 9 September 2007; accepted 2 November 2007

Abstract

Continental strata of the Home Station Sandstone Member of the Snowy Plains Formation of the Mansfield Group, AvonSupergroup, Mansfield Basin in Victoria, Australia, contains diverse and abundant trace fossils, including domichnia, fodinichnia,pascichnia, and repichnia. The ichna are found in numerous beds in an approximately 28-m-thick unit referred to as Fish Hill,within the 250- to 500-m-thick Home Station Sandstone Member. The beds consist of alternating laminated mudstone and siltstone,thick-bedded ripple laminated sandstone, and thick-bedded massive sandstone with occasional discontinuous wavy lamination,trough cross-bedding and small-scale planar cross-bedding, interpreted as meandering river deposits.

The trace-fossil assemblage includes Rusophycus, Cruziana, Palaeophycus, and Sagittichnus, a new ichnospecies of Fuer-sichnus and of Margaritichnus, and a new ichnotaxon Platicytes lioparadus. The geological and geographical distributions ofmany of these ichnotaxa are extended. Excellent examples of the compound nature of Cruziana and Rusophycus are illustrated.Ichnotaxa are herein grouped into a Rusophycus, Margaritichnus, and Cruziana ichnocoenoses, each identified according to theirpaleoenvironmental occurrence. This is the first detailed description on ichnofossils reported from Fish Hill, adding importantpalaeoenvironmental and palaeoecological information to what is previously known from these rocks based on vertebrate fossils.© 2007 Elsevier B.V. All rights reserved.

Keywords: Lower Carboniferous; Fish Hill; Snowy Plains Formation; Australia

1. Introduction

This paper is the first detailed description of theichnofossils from the Fish Hill locality in the Home StationSandstone Member of the Lower Carboniferous SnowyPlains Formation of theMansfieldGroup,MansfieldBasin,in southeast Australia (Fig. 1A). The trace-fossil assem-blage contains a diverse and abundant ichnofauna thatincludes domichnia, fodinichnia, pascichnia, and repichnia

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Palaeogeography, Palaeoclimatology, Palaeoecology 258 (2008) 257–276www.elsevier.com/locate/palaeo

⁎ Corresponding author. Archaeology Program, School of Historicaland European Studies, La Trobe University, Melbourne, 3086Australia. Tel.: +61 3 94792806; fax: +61 3 94791881.

E-mail addresses: [email protected] (J.M. Garvey),[email protected] (S.T. Hasiotis).

0031-0182/$ - see front matter © 2007 Elsevier B.V. All rights reserved.doi:10.1016/j.palaeo.2007.11.002


Fig. 1. A, The township of Mansfield (latitude 37°011′ South and longitude 146°224′ East), approximately 200 km northeast from Melbourne,Victoria, Australia. The state abbreviations areWA—Western Australia, NT—Northern Territory, SA—South Australia, QLD—Queensland, NSW—New South Wales, TAS—Tasmania, Victoria is coloured black. B, Generalised stratigraphy of the Mansfield Basin after VandenBerg et al. (2006).Arrow indicates approximate position of the Early Carboniferous (Tournaisian) Fish Hill (37°011′S, 146°224′E) and Broken River fish localities in theHome Station Sandstone Member. C, Detailed map of the Mansfield region indicating the regions of ichnofossil collection (Fish Hill and the BrokenRiver).

258 J.M. Garvey, S.T. Hasiotis / Palaeogeography, Palaeoclimatology, Palaeoecology 258 (2008) 257–276


recovered from the areas of Fish Hill and from along theBroken River.

Ichnofossils in the Snowy Plains Formation were firstnoted by Sweet (1889) while searching for vertebratefossils. He mentioned bilobed pittings and fucoids orworm castings on Fish Hill (formally known as TanneryPaddock, Fig. 1C). Seedsman (1974) recorded anoccurrence of ichnofossils while searching for vertebratetracks and trails fromBridge Creek, whileMarsden (1988)identified trace fossils and bioturbational structures fromthe Snowy Plains Formation,Mansfield Group,MansfieldBasin, Victoria. Ansell and Archbold (2002) identifiedthirteen ichnogenera and three ichnofacies according toBromley's (1996) modified scheme, at a locality alongBridge Creek, a subsidiary of the Broken River, between1.5 and 2 km northwest of Fish Hill (Fig. 1C).

The Fish Hill locality is well known for its EarlyCarboniferous fish fauna (e.g. Sweet, 1889) as well asfrom outcrops along the Broken River, approximately10 km north of Mansfield, Victoria (Fig. 1C). The fishfauna includes: three sarcopterygians, the large rhizo-dontid, Barameda decipiens (Woodward, 1906; see alsoLong, 1989), the small rhizodontid Barameda mitchelli(Holland et al., 2007), and the small dipnoan, Delatitabreviceps (Woodward, 1906; see also Long and Camp-bell, 1985); three actinopterygians, Mansfieldiscussweeti and M. gibbus (Woodward, 1906; see alsoLong, 1988), and Novogonatodus kasantsevae (Long,1988); a relatively large acanthodian, Gyracanthidesmurrayi (Woodward, 1906), and two smaller acantho-dians, Acanthodes australis (Woodward, 1906) and Eu-pleurogmus creswelli (McCoy, 1890). Recently, severalnew chondrichthyan taxa were recorded from the SnowyPlains Formation, including hybodonts, ctenacanths, anda new species of Ageleodus (Garvey and Turner, 2006).

Ichnofossil analyses presented here provide new andimportant information on the palaeoenvironmental andpalaeoecological analyses for this Early Carboniferousfossil fish locality. The ichnofossils represent theactivity of invertebrates that are not represented bybody fossils in the Lower Carboniferous Snowy PlainsFormation, and are herein grouped into Rusophycus,Margaritichnus, and Cruziana ichnocoenoses—eachidentified according to their paleoenvironmental occur-rence. These ichnofossils can be used to infer conditionsin continental settings that cannot be gleaned from thesedimentology or vertebrate palaeontology alone.

2. Geological setting

Fish Hill and the Broken River localities are in theHome Station Sandstone Member of the Snowy Plains

Formation, comprising the majority of the MansfieldBasin (VandenBerg et al., 2006) (Fig. 1B). TheMansfieldBasin is one of four remnant structural basins that overlieolder Palaeozoic rocks in the Howitt Province, whosedevelopment is poorly understood (O'Halloran and Cas,1995). The Mansfield Basin is filled with a redbedsuccession of conglomerates, sandstones, and mudstones(Marsden, 1988). It contains the Avon Supergroup, whichis subdivided into the Mansfield, Wellington Volcanic,and the Delatite Groups (VandenBerg et al., 2006). TheMansfield Group consists of the Snowy Plains Formationand the older Mt. Kent Conglomerate. The Snowy PlainsFormation is subdivided into seven members, which in-cludes the Home Station Sandstone Member from whichthe ichnofossils were found (Fig. 1B).

The ichnofossil assemblages are in numerous beds inan approximately 28-m-thick unit referred to as Fish Hill,within the Home Station SandstoneMember. The beds onFishHill are alternating laminatedmudstone and siltstone,thick-bedded ripple laminated sandstone, and thick-bedded massive sandstone with occasional discontinuouswavy lamination, trough cross-bedding and small-scaleplanar cross-bedding. Three facies each containing alithofacies and ichnocoenoses are identified. The mostcommon facies is the Low Energy Floodplain Facies,which contains the Laminated Mudstone–SiltstoneLithofacies (Garvey, 2005; Table 1). It occurs repeatedlythroughout the sequence and is often divided by relativelythin sandstone beds, especially in the middle of thesequence. The Rusophycus ichnocoenosis is recordedfrom the LowEnergy Floodplain Facies based on the highnumbers of repichnia (crawling or walking traces, e.g.,Cruziana) and cubichnia (resting or nesting traces, e.g.,Rusophycus), and the shallow fluvial environmentdominated by siltstone (Garvey, 2005). Five ichnogeneraare identified from this ichnocoenosis: Fuersichnus, Sa-gittichnus, Platicytes, Cruziana and Rusophycus.

The Moderate Energy Sand-Sheet Facies contains theThin-Bedded Rippled Sandstone Lithofacies. Thislithofacies represents minor sandstone bodies, b0.2 mthick, between strata of the Laminated Mudstone–Siltstone Lithofacies (Garvey, 2005). Desiccation crackson the top of beds of this lithofacies indicate that the topof the lithofacies at least periodically dried out. TheMargaritichnus ichnocoenosis are identified in thisfacies (Garvey, 2005; Table 1), suggesting deposition ina sandy environment resulting from a brief raised-energy episode, with colonisation during a calm periodbefore the river plain area was covered again bysedimentation. Four ichnotaxa are identified from thisichnocoenosis: Rusophycus, Cruziana, Palaeophycusand Margaritichnus.

259J.M. Garvey, S.T. Hasiotis / Palaeogeography, Palaeoclimatology, Palaeoecology 258 (2008) 257–276


The High Energy River Channel Facies contains theThick-Bedded Sandstone Lithofacies. The Thick-Bedded Sandstone Lithofacies comprises approximately11 m of the Fish Hill section (Garvey, 2005). It consistspredominantly of very fine- to fine-grained, red, purple,and brown coloured sandstone beds with small shalelenses, and is the most resistant lithofacies on Fish Hill.Within the High Energy River Channel Facies is theCruziana ichnocoenosis, with only two ichnotaxarecorded throughout the sequence, Cruziana and Ruso-phycus (Garvey, 2005; Table 1).

3. Methods

Ichnofossils were found by one of us (JMG) indiscrete horizons on Fish Hill during excavations forvertebrate fossils between 2000 and 2004 (Garvey et al.,2002; Garvey, 2006). Ichnofossils were also recordedand collected from beds exposed along Broken Riverapproximately 3 km downstream from Fish Hill (markedPlaticytes on Fig. 1C).

The general morphology and orientation of theichnofossils were recorded in the outcrop, and sampleswere collected. The stratigraphic positions of theichnofossils and outcrop description were also recordedwith the degree of bioturbation, determined by means ofthe ichnofabric index (modified fromDroser and Bottjer,1986). The bioturbation recorded on Fish Hill, theidentified ichnocoenoses, the depositional environment,and the taphonomic history is currently being described.

Trace fossils described in this paper are housed inMuseum Victoria, Melbourne, Australia. Slab numbers are

NMV P310999–NMV P311009 and NMV P311630–NMV P311639. Long and slightly curving trace fossilstentatively assigned to Undichnus were previously identi-fied in outcrops along the Broken River, slightly down-stream fromwhere the newMansfield–TolmieRoad bridgecrosses the Broken River and close to where Platicytes,described herein, were recorded (A. Warren pers. comm.2003). Unfortunately, the horizon with Undichnus tracescould not be relocated during the 2000–2004 field seasons,and, therefore, are described only from a photograph.

4. Systematic ichnology

4.1. Ichnogenus Cruziana d'Orbigny, 1842

Cruziana d'Orbigny, 1842 comprises trace fossilspreserved as elongate bilobate furrows covered byherringbone-shaped ridges. They vary in size and surficialmorphology due to varied behaviour of the animal(Häntzschel, 1975). It is possible that trace fossils assignedtoCruziana are associated with deposit feeding on surfacesediments. An animal ploughing into the sandy sedimentand impinging on the underlying mud likely formed mostCruziana (Seilacher, 1955, 1970, 1982; Goldring, 1985).The Lower Palaeozoic Cruziana has been attributed totrilobite activity (Osgood, 1970), in particular Cruzianasemiplicata (Fortey and Owens, 1999).

4.2. Cruziana problematica (Schindewolf, 1928)

Material: Several specimens preserved on red shale onNMVP311630 (Fig. 2) and its counterpart NMVP311631.

Table 1Description of the Lithofacies and recorded trace fossils from the Fish Hill, Home Station Sandstone Member

Lithofacies Description

Laminated mudstone-sandstonelithofacies

Most common lithofaces on Fish Hill, which is characterised by laterally continuous planar laminations. There arealso rare wavy discontinuous laminations and planar cross-bedding towards the top of the sequence. Desiccationcracks are periodically exposed. This bed is interpreted as being characteristic of a low-energy floodplainenvironment, and is assigned to the Rusophycus ichnocoenosis. Fuersichnus commerfordi isp. n., Sagittichnuslincki, Cruziana problematic, Platicytes lioparadus isp. n, and Rusophycus eutendorfensis, R. jenningsi, and R.biloba were recorded.

Thin-bedded rippled sandstonelithofacies

Minor beds of fine calcareous and noncalcareous sandstone, with all bedsb200 m in thickness. Beds are laterallycontinuous with sharp boundaries, and are characterised by uniform fine to very-fine grained sandstone. Small-scale trough cross-bedding often occurs, while small-scale planar cross-bedding and discontinuous wavylamination is also present. Mudcracks and clasts occur in some of the beds. This bed is interpreted as a moderateenergy channel deposit, and assigned to the Margaritichnus ichnocoenosis. Rusophycus isp., Cruzianaproblematica, Palaeophycus tubularis, and Margaritichnus mansfield isp. n. were recorded.

Thick-bedded sandstonelithofacies

Consists of very fine to fine grained, red, purple and brown sandstone with small shale lenses. The beds exhibit anover-all coarsening upwards in grain size. Occasional discontinuous wavy lamination, trough cross-bedding andsmall-scale planar cross-bedding occurs. Represents a high-energy river channel environment. This bed isinterpreted as a high energy river channel environment, and assigned to the Cruziana ichnocoenosis. Rusophycusisp. and Cruziana probelmatica were recorded.

Mansfield Group, Mansfield Basin, Victoria (from Garvey, 2005).



Fig. 2. Slab NMV P311630 with ichnofossils from the Snowy Plains Formation, Fish Hill. A, Entire slab NMV P311630. B, Close up of Cruzianaproblematica, Rusophycus biloba, R. eutendorfensis and R. jenningsi on slab NMV P311630. Examples of C. problematica formed by successive,overlapping Rusophycus are located to the left and lower left of the asterisk.



Locality and horizon: Common on Fish Hill, beingidentified from all three ichnocoenoses. It was alsorecorded from the same horizon as specimens ofactinopterygians and Gyracanthides.

Description: Typically narrow, bilobate Cruzianaformed by distinct or indistinct, closely spaced, finestriations typically transverse to the path of the trail, or areoblique at tighter curves (after Keighley and Pickerill,

Fig. 3. Slab NMV P311632 from Fish Hill. A, Entire slab of NMV P311632. B, Close up of slab NMV P311632 with Fuersichnus commerfordi isp.n. and Rusophycus isp. labelled. C, Close up of slab NMV P311632 with Sagittichnus lincki indicated.



1996). Specimens fromFishHill average 10mm long,withthe longest 19 mm and the shortest between 7 and 8 mm.They have an average width between 3 and 4 mm.

Remarks: Cruziana problematica has been found inboth marine and continental deposits, and ranges in agefrom the Cambrian to the Jurassic, and possibly even theCretaceous (Pickerill, 1992; Keighley and Pickerill,1996, 1997; Buatois et al., 2000). Specimens collectedfrom Fish Hill in particular resemble C. problematicafrom ephemeral fluvial channel deposits in the lower-most Port Hood Formation, Cumberland Group (West-phalian A), western Cape Breton Island of easternCanada (Keighley and Pickerill, 1996, Fig. 1B).

Cruziana is often in association with Rusophycus.On NMV P311630 C. problematica is associated withRusophycus biloba, R. eutendorfensis, and R. jenningsi.

Specimen NMV P311630 and its counterpart NMVP311631 provide good examples of compound tracefossils, where several forms of behaviour have combinedto form one type of trace fossil with distinct morpholo-gical parts. The repichnia (crawling or walking) Cruzi-ana was produced by overlapping cubichnia (resting ornestling) Rusophycus structures, where individual bilo-bate furrows merged into a single trace fossil as theanimal moved through the sediment (Fig. 2B). Therepetitive Rusophycus, thus, has merged to form Cruzi-ana. On this bedding plane the overlap of Rusophycusleaves the impression of short, incremental movementstaken by the tracemaker until it stops in a final and deeperRusophycus biloba trace.

We interpret the tracemaker of Cruziana problema-tica to have been an insect or noninsect arthropod (e.g.,Pollard, 1985). The invertebrate most likely had a short,bilaterally symmetrical body with fully functional ap-pendages rather than an elongate body with short, fleshyprolegs, based on the well-defined transverse scratcheswithin the trace fossil.

4.3. Fuersichnus Bromley and Asgaard 1979

Bromley and Asgaard (1979) erected the ichnogenusFuersichnus to describe horizontal to oblique, retrusiveburrow complexes composed of clusters of J-shapedfills, showing varying degrees of organisation. Thesetrace fossils were thought to have been produced by adeposit-feeding animal (Bromley and Asgaard, 1979).Hasiotis (2004) suggested that Fuersichnus isp. mayhave been constructed by insect nymphs similar toextant mayfly larvae based on comparison with modernanalogues (e.g., Hunt, 1953; Ward, 1992). Mayflies andtheir relatives first appear during the Carboniferous(Grimaldi and Engel, 2005).

4.4. Fuersichnus commerfordi isp. n.

Material: One slab, NMV P311632 (Fig. 3A and B).Holotype: Seven specimens on slab NMV P311632.Etymology: Named for the Commerford families on

whose property most of the excavations conducted byJMG from 2000–2004.

Locality and horizon: The specimens on slab NMVP311632 were collected from the Home Station Sand-stone Member of the Snowy Plains Formation, MansfieldBasin, Victoria (37°011′S, 146°224′E). Slab NMVP311632 came from the Low Energy Floodplain Facieson Fish Hill (Garvey, 2005) in association with Sagit-tichnus and Rusophycus. Actinopterygians, gyracanthid,and dipnoan fossil material were also recovered from thisbed.

Diagnosis: Fuersichnus occurs within small clusters,have a smooth surface, and is gently crescent shaped.

Description: Fuersichnus commerfordi displays acrescent shape, is parallel to the bedding plane, and ispreserved in convex hyporelief. They are relativelyelongate, between 26 and 38 mm long, and between 5and 6 mm wide. Two specimens (Fig. 3A and B) depictslight curving downwards and sideways movement intheir construction. The longer and straighter trace fossilsappear to be in desiccation cracks where either theanimal has constructed burrows within the desiccationcrack fill, or the desiccation cracks formed as a result ofthe burrows weakening the surface sediment. One of us(STH) has observed similar interactions betweenmoderninsect traces in and along drying pools of water, whichhas also been observed in ancient deposits (e.g., Bromleyand Asgaard, 1979). Overall, the traces have a smoothsurface and are gently curving in a crescent shape. Giventheir simple morphology they represent the lowest levelof organisation for this ichnogenus.

Remarks: The Fish Hill specimens differ from Fuer-sichnus communis from the Triassic of East Greenland(Bromley and Asgaard, 1979), as the East Greenlandspecies are more J- and deeply U-shaped spreites, aremuch more densely packed per specimen. F. commerfordidiffers from Fuersichnus isp. in Jurassic strata of NewEngland (Gierlowski-Kordesch, 1991), as F. commerfordidisplays crescent shape and smooth, rather than notched orknobbly on the surface. F. commerfordi differs fromFuersichnus isp. from New Brunswick (MacNaughtonand Pickerill, 1995), as F. commerfordi is not foundparallel to one another and gently curve into almost acrescent shape rather than the J-shape seen in theNew Brunswick specimens. Finally, the Fish Hill speci-mens differ from F. striatus from the Cretaceous ofAntarctica (Buatois, 1995), as F. commerfordi is not



strongly U-shaped, and do not have parallel striationsrunning longitudinally along the axis of the burrow.

Fuersichnusmay be well organised where they appearas either ear- or tongue-shaped, spreite-like structures,may resemble bunches of bananas with uneven spreite(seen as intermediate by Bromley and Asgaard, 1979), ormay look unorganised where they disaggregate into aloose cluster or isolated curved burrows (seen as leastorganised by Bromley and Asgaard, 1979). Bromley andAsgaard (1979) stated that it was difficult to apply aspecies name that could incorporate the gradational rangein morphologies, but introduced Fuersichnus communisto represent members at the well-organised end of theseries, thus, allowing for further species to be described forthe lesser organised members if necessary. F. communiswas interpreted as a record of the probing behaviour of a

deposit-feeding organism that moved along a curved path,backfilling as it moved and periodically depositingsediment immediately adjacent to the previous probe(Bromley and Asgaard, 1979).

Given that the Fuersichnus from Fish Hill occurswithin small clusters (Fig. 3A and B), have a smoothmorphology, and are gently crescent shaped from end toend, they exhibit a simpler morphology than othermembers of this ichnogenus. They represent the lowestlevel of organisation of Fuersichnus and, therefore,warrant their own ichnospecies.

Mayflies have been reported to produce shallow J- toU-shaped, retrusive, and backfilled burrows in modernenvironments (e.g., Hunt, 1953; Ward, 1992). Sincemayflies and their relatives first appear during theCarboniferous (Grimaldi and Engel, 2005) Fuersichnus

Fig. 4. Slab NMV P311635 from Fish Hill. A, Slab with Margaritichnus mansfieldi isp. n., Rusophycus isp., and tool marks indicated. B, Siliconmold of Margaritichnus mansfieldi isp. n. with the groove indicated.



commerfordi may have been constructed by an insectnymph similar in form and behavior to extant mayflylarvae.

4.5. Margaritichnus Bandel, 1973

The ichnogenus Margaritichnus was introduced byBandel (1973) to describe a trail from marine deposits ofthe Upper Pennsylvanian Rock Lake Shale Member,Stanton Limestone, Lansing Group, Kansas.While Bandel(1973) stated that it was constructed of ball-like structuresthat were either connected or unconnected, Hakes (1976)did not find any connection between the balls.

Margaritichnus sp. has been interpreted as represent-ing the fodinichnia fecal strings belonging to a sediment-feeding, worm-like animal (Bandel, 1967, 1973; Sudanand Shama, 2000).Hakes (1976) reinterpreted these tracesas domichnia of a soft-bodied animal, possibly anemone-like, while Seilacher (1990) believed them to have beenproduced by probing behaviour. They could also beinterpreted as a subspherical invertebrate, for example aclam, moving through the sediment. Margaritichnus sp.is, therefore, an ichnogenus that has been interpreted inseveral ways; however, neither the Mansfield specimennor the type material shows evidence of probing.

4.6. Margaritichnus mansfieldi isp. n.

Material: Single specimen on slab NMV P311635.Holotype: Single specimen on slab NMV P311635

(Fig. 4).Etymology: Named after the township of Mansfield,

Victoria, Australia.Locality and horizon: The Home Station Sandstone

Member of the SnowyPlains Formation,Mansfield Basin,Victoria (37°011′S, 146°224′E). Slab NMV P311635came from a thin, fine-grained sandstone outcrop towardsthe bottom of the Fish Hill sequence assigned to theMargaritichnus ichnocoenosis. While only one specimenwas collected, several were noted in the field. Also presenton slab NMVP311635 isRusophycus isp. and tool marks.It comes from a horizon fromwhich no vertebrate materialwas collected (Garvey, 2005).

It should be noted that the slab NMV P311635 is thecounterpart as the actual specimen was unable to becollected. The silicon mold (Fig. 4B) shows what thespecimen looked like in the field.

Diagnosis: Margaritichnus mansfieldi isp. n. aresmall, flattened, subspherical aligned structures, circularto slightly elliptical in cross section, unornamented walls,structureless filling, and with a longitudinal groove downthe centre of aligned structures.

Description: Margaritichnus mansfieldi is preservedin convex epirelief, parallel to the bedding plane andcircular to slightly elliptical in cross section. The wallsare unornamented and the filling is structureless. Thereare approximately 10 small, flattened subspherical,aligned structures each with a longitudinal groove run-ning down its centre. In total, the specimen is approxi-mately 30 mm long and 2 mm wide, with the individualstructures measuring about 2 to 3 mm in diameter. Thetrace is a gently curving continuous trail, which may bedue to a tool mark that the animal encountered, alteringits direction.

Remarks: Margaritichnus has been identified fromnumerous deposits including the Precambrian of Russia(Fedonkin, 1976, 1977), Precambrian of Ediacara ofSouth Australia (Glaessner, 1969), the Cambrian Gar-byang Formation of Kumaun in India (Sudan and Shama,2000), the Upper Silurian of Artic Canada (Narbonne,1984), the Late Silurian Holmestrand Formation ofNorway (Davies et al., 2006), the Pennsylvanian(Desmoinesian) of Colorado (Lockley et al., 1987), theUpper Pennsylvanian of Kansas (Bandel, 1967; Hakes,1976), the Permian of the Carnarvon Basin in WesternAustralia (Moore et al., 1980a,b; Skwarko and Seilacher,1993), and possibly the Triassic lacustrine deposits of EastGreenland (Bromley and Asgaard, 1979). Bandel (1967)established the ichnospecies Margaritichnus reptilis,from the Upper Pennsylvanian of Kansas, to describeball-like structures of 15 to 30 mm in diameter that werealigned commonly like a string of pearls.

Margaritichnus mansfieldi isp. n. from Fish Hilldiffers from all other Margaritichnus ichnospecies inthe presence of a continuous longitudinal groove downthe centre of the trace. The identification of thisichnogenus from Fish Hill represents the third occur-rence of the ichnogenus in Australia, and the first fromthe Carboniferous. It is also the third reported occur-rence in freshwater environments.

Margaritichnus reptilis was interpreted as being thetrail of a large, wormlike, sediment-feeding animal thatpackaged its fecal pellets inmucus (Bandel, 1967). It wasthought likely that the pellets were produced just belowthe sediment surface. Bromley and Asgaard (1979)described possible Margaritichnus as clearly having abiogenic origin. They appeared to have been made bydeposit-feeding, aquatic organisms associated primarilywith Rusophycus eutendorfensis. While R. eutendorfen-sis has been identified from Fish Hill, they were notfound in the same beds as Margaritichnus mansfieldi.Instead, at Fish Hill M. mansfieldi was found inassociation with other Rusophycus isp. on slab NMVP311630.



Fig. 5. Palaeophycus tubularis from Fish Hill. A, Slab NMV P311639. B, Small section of NMV P311639 with the P. tubularis specimen that dividesinto three (arrowed). C, Slab NMV P311636.



A bivalve is likely to have constructed Margaritich-nus mansfieldi. Though Margaritichnus has beenattributed to the activity of an anemone (Hakes, 1976),the freshwater fluvial environments of the Snowy PlainsFormation probably did not support anemones. Nofreshwater anemones are known from lower or middlePalaeozoic continental rocks. No freshwater wormstoday are known to deposit feed in such a way as toproduce traces similar toMargaritichnus, and, therefore,it is highly unlikely that they were the constructors of thistrace fossil.

4.7. Palaeophycus Hall, 1847

The ichnogenusPalaeophycus displays a wide range ofmorphologies, and there are still nomenclatural problemswith the ichnogenus Planolites (Pemberton and Frey,1982; Bromley, 1996). They are often cylindrical orsubcylindrical, usually lie parallel to the bedding plane, arefrequently unbranched, but may display occasionalbranching with branches intersecting commonly. Thewalls are mostly smooth but may exhibit faint longitudinalstriae (Häntzschel, 1975), and several types of Palaeo-phycus have longitudinal striae and perpendicular con-

strictions. Palaeophycus can occur as isolated burrows orin groups (Sarkar and Chaudhuri, 1992).

Palaeophycus represents the passive infilling of anopen dwelling burrow belonging to a predaceous orsuspension-feeding animal (Pemberton and Frey,1982). It has been recorded from a wide variety ofenvironmental settings, from the Precambrian to theHolocene.

4.8. Palaeophycus tubularis Hall, 1847

Material: Numerous specimens preserved on threeslabs of red fine-grained sandstone: NMV P311633,NMV P311636 and NMV P311639 (Fig. 5).

Locality and horizon: The Home Station SandstoneMember of the Snowy Plains Formation, MansfieldBasin, Victoria. They are from a fine-grained sandstoneinterpreted as a channel deposit, and assigned to theMargaritichnus ichnocoenosis. Palaeophycus tubulariswas identified from the same horizon as a semi-articulatedspecimen of rhizodontid (Garvey, 2005; Holland et al.,2007).

Description: Unornamented, horizontal, slightly sin-uous, smooth burrows of varying diameter, preserved in

Fig. 6. Examples of chambers of Platicytes lioparadus isp. n. from Fish Hill. A, NMV P311002; B, NMV P311011; C, P311003; D, NMV P311004the twisted chamber; E, NMV P3110999 (Holotype), the most complete P. lioparadus from Fish Hill, with the longitudinal central groove arrowed.Scale bars are 1 cm.



convex hyporelief. Some of the burrows change diameteralong their length. The majority of the burrows curveslightly, and almost all intersect with at least one othertrace fossil. All of the burrows are unbranched except for asingle example on NMV P311639, which appears todivide into three (Fig. 5B). The burrows are so crowdedthat the entire surface of the sandstone is bioturbated.Burrow diameter ranges from 2 to 7 mm, while the lengthvaries from 4 to 20 mm, depending on how much of theburrow is exposed and whether another subsequentburrow has intersected it. These burrows were likely tohave been much longer than observed, as they appear tocontinue into the sediment. Avariety of burrow shapes arefound on NMV P311633, NMV P311636 and NMVP311639, with some burrows being rather straight, someslightly sinuous, and one S-shaped.

At Fish Hill two different size classes of Palaeophy-cus tubularis were collected. On slabs NMV P311633and NMV P311639 the width of the largest burrows arebetween 4 and 7 mm (Fig. 5A and B), while on slabNMV P311636 the burrows are smaller with the width ofthe largest between 2 and 3 mm (Fig. 5C). It is probable,therefore, that two types of tracemakers were responsiblefor P. tubularis.

Remarks: This ichnogenus is often confused withPlanolites, but in Palaeophycus there is no distinctionbetween the lithology of the filling and the trace fossils,whereas in Planolites there is a difference (Osgood,1970). For detailed discussions on the distinction betweenPalaeophycus and Planolites refer to Pemberton and Frey(1982), Stanley and Pickerill (1994), and Keighley andPickerill (1995).

Palaeophycus has been identified from several fresh-water environments, including Carboniferous fluvial toestuarine facies of Indiana (Archer and Maples, 1984),Carboniferous lacustrine deposits of Argentina (Buatoisand Mángano, 1993) and Canada (Pickerill, 1992),Permian fluviolacustrine deposits of Argentina (Aceño-laza and Buatois, 1993; Buatois et al., 1997), Triassicfluvial deposits of Canada (MacNaughton and Pickerill,1995), Jurassic estuarine deposits of the United States(Hasiotis, 2004), Cretaceous lacustrine deposits of Spain(Fregenal-Martinez et al., 1995), Cretaceous continentaldeposits of southeast Korea (Kim and Pickerill, 2002),and the Palaeogene fluvial rocks of Utah (D'Alessandroet al., 1987). In particular the specimens identified fromthe Snowy Plains Formation, Fish Hill, resembleP. tubularis from Lower Carboniferous continentaldeposits of east Canada (Pickerill, 1992, Fig. 4A). Thisindicates that animals with similar size and behaviourwere present during the Early Carboniferous thatproduced very similar trace fossils in continental

environments in south-eastern Australia and easternCanada. The identification of Palaeophycus tubularis isthe second record of this ichnotaxon from the MansfieldBasin, having been identified previously from an outcropalong Bridge Creek (Ansell and Archbold, 2002).

4.9. Platicytes n. igen.

4.9.1. Type ichnospecies; Platicytes lioparadus isp. n.Diagnosis: Ichnogenus is established to describe hori-

zontal burrows that are oval in cross section, unbranched,perpendicular to the bedding plane, and terminate in ahemispherical chamber. This morphology likely repre-sents a passively filled dwelling burrow (domichnia).

4.10. Platicytes lioparadus isp. n.

Material: Five specimens, NMV P310999, NMVP311001–NMV P311004 (Fig. 6).

Holotype: Single specimenNMVP3110999 (Fig. 6e).Etymology: Greek Plati flat or horizontal, cyte hol-

low vessel, lio smooth and paradus, passage.Locality and horizon: Five natural molds filled with

fine-grained sand were collected, while many more wereidentified in the field. These were all restricted to a smallarea of outcrop along the Broken River, in the HomeStation Sandstone Member of the Snowy Plains Forma-tion, Mansfield Basin, Victoria (37°011′S, 146°224′E).This outcrop was approximately 300–400 m down-stream from where the new Mansfield–Tolmie RoadBridge crosses the Broken River, approximately 1 kmwest of Fish Hill (marked Platicytes on Fig. 1C). Thisbed is equivalent stratigraphically to the outcropsexposed on Fish Hill.

Diagnosis: Platicytes lioparadus isp. n. are dorsoven-trally flattened horizontal burrows with smooth walls thatterminate in a single asymmetrically flattened, bulbouschamber.

Description: All of the burrows are parallel to thebedding plane and dorsoventrally flattened. Theyrepresent simple burrows terminating in an asymme-trically flattened, bulbous, single chamber. Thesechambers vary in size between 50 and 85 mm (Fig. 6).One of the burrows was found to have a twisted chamber(NMV P311004, Fig. 6D). The most complete specimencollected (NMV P310999, Fig. 6E) is approximately400 mm long, and 20 to 30 mm wide, with a chamber60 mm long, 45 mm wide, and 20 mm high. This doesnot represent the entire length of the specimen, as theburrow continued into the outcrop. This burrow isunique as it has a shallow groove or furrow present onthe shaft of the burrow, running longitudinally (Fig. 6E,



arrowed). In cross section the trace fossils are ellipsoid.All of the P. lioparadus were unbranched and they showlittle individual variation.

Remarks: The specimen NMV P310999 from FishHill (Fig. 6E) has a groove along its length. This grooveis interpreted as indicating that it was once an openburrow (similar to Palaeophycus) that later was filledpartially with sediment and compacted to produce thegroove in the ceiling of the trace fossil. This is importantas it suggests that the sediments were relatively firmwhen the trace was created, as it does not have thickwalls for support. This open burrow morphology iscommon in continental environments where terrestrialand aquatic sediments are often gravitationally drainedand compacted (e.g., Hasiotis, 2002, 2004). The otherfour specimens NMV P311001–NMV P311004 do notshow signs of this grove.

Platicytes is a passive infilling of an open dwellingburrow belonging to a predaceous or scavengingarthropod similar in anatomy to such crustaceans asfreshwater crayfish, crabs, and burrowing shrimp (e.g.,Hasiotis and Mitchell, 1993; Hasiotis et al., 1993;Marshall et al., 2002). The burrow was likely in ashallow-water, low-energy, nearshore setting wheresediments were firm enough to construct an openburrow. The burrow likely remained submerged as thatthe burrow walls remained plastic enough prior to burialnot to record any substantial surficial morphology.

4.11. Rusophycus Hall, 1852

The ichnogenus Rusophycus Hall, 1852 describesshort, bilobate, coffee-bean shaped structure usually witha median furrow. One of the most diagnostic features ofthe ichnogenus is that the lobes are transversely wrinkledby anteriolaterally orientated fine striae (Häntzschel,1975). Rusophycus are preserved mostly in convexhyporelief in silt or sand-sized strata. It is predominatelya Palaeozoic ichnogenus applied to cubichnia (nestling orresting behavioural trace fossils) (Osgood, 1970).

The diversity of Rusophycus-like traces has beenextended to include continental environments (e.g.,Trewin, 1976; MacNaughton and Pickerill, 1995; Cookand Bann, 2000), so the assumption that marineorganisms solely produced Rusophycus has been re-evaluated. Diversification of continental organisms andichnofossils were at least present in the Ordovician, andcomplex aquatic and terrestrial ecosystems had evolvedby the Devonian (Maples and Archer, 1989). Therefore,the identification of Rusophycus in predominatelyfreshwater depositional environments as preserved inthe Snowy Plains Formation is not surprising.

There have been numerous discussions as to thedistinction between Rusophycus, Isopodichnus, andCruziana (e.g., Bromley and Asgaard, 1979; Pollard,1985; Keighley and Pickerill, 1996). Numerous species ofRusophycus are classified according to small differencesin shape, and there has been debate as to the nomenclaturestatus of Rusophycus and Cruziana, with the former oftenregarded as a synonym of Cruziana (Häntzschel, 1975).Seilacher (1955) and Osgood (1970), however, arguedthat Rusophycus be retained for short bilobate restingtrails.

4.12. Rusophycus biloba Vanuxem, 1842

Material: Several specimens identified on a slab ofred shale NMV P311630 (Fig. 2), and its counterpartNMV P311631.

Locality and horizon: Fish Hill, in the Home StationSandstone Member of the Snowy Plains Formation,Mansfield Basin, Victoria.

Description: Rusophycus biloba are short, bilobate-shaped cubichnia (resting or nestling), parallel to thebedding plane, and preserved in convex hyporelief. Thelobes are covered in anterolaterally directed striae, andmeasure 3 mm long and 2 mm wide.

Remarks: The ichnotaxa Rusophycus (Fucoides) bi-loba was established by Vanuxem (1842) to describebilobate traces from Silurian (Clinton) deposits in NewYork State. The Rusophycus biloba from Fish Hill areassociated with R. eutendorfensis, R. jenningsi andCruziana problematica. Since the Snowy Plains For-mation at Fish Hill is suggestive of a fluvial system, anontrilobite arthropod is likely to have produced theMansfield R. biloba.

This is the second occurrence of Rusophycus in theTournaisian of the Snowy Plains Formation as Anselland Archbold (2002) identified R. carbonarius from anew locality along Bridge Creek, however, this taxonwas not identified from the Fish Hill assemblage.

4.13. Rusophycus eutendorfensis Linck, 1942

Material: NMV P311630 (Fig. 2) and its counterpartNMV P311631.

Locality and horizon: The specimens are preservedon a slab of red shale and were collected fromexcavations by JMG on Fish Hill, in the Home StationSandstone Member in the Snowy Plains Formation,Mansfield Basin, Victoria between 2000 and 2004.

Description: Typically small, subparallel coffee-beanshaped lobes covered with almost radial, commonlybifid, fine scratch marks that extend to the extreme edge



or outside buckle section of the lobed structure (afterLinck, 1942; Fillion and Pickerill, 1990; Keighley andPickerill, 1996).

Remarks: Rusophycus eutendorfensis has beendescribed from numerous localities, including theLower Cambrian marine Mickwitzia Sandstone, south-central Sweden (Jensen, 1997), the lacustrine–fluvialdeltaic Devonian–Carboniferous Albert Formation,south-eastern New Brunswick, eastern Canada (Fillionand Pickerill, 1990; Keighley and Pickerill, 1996), andthe freshwater Triassic Carlsberg Fjord, East Greenland(Bromley and Asgaard, 1979). As the Snowy PlainsFormation at Fish Hill is suggestive of a fluvial system,a nontrilobite arthropod is likely to have produced theMansfield R. eutendorfensis. This is the first identifica-tion of Rusophycus eutendorfensis from the LowerCarboniferous of the Snowy Plains Formation, Victoria.

4.14. Rusophycus jenningsi Fenton and Fenton, 1937

Material: Several specimens identified on a slab ofred shale NMV P311630 (Fig. 2), and its counterpartNMV P311631.

Locality and horizon: Fish Hill, in the Home StationSandstone Member in the Snowy Plains Formation.

Description: These traces are short, typically bilo-bate, with a deep U-shaped profile (arrowed in Fig. 2B),and preserved in convex hyporelief, and measure 3–

6 mm long and 3–5 mm wide. Their lobes are coveredwith almost radial, typically bifid, fine scrape lines thatextend towards the extreme outside edge of the lobe partof the structure.

Remarks: Rusophycus jenningsi was originally iden-tified by Seilacher (1970) to incorporate traces assignedby Fenton and Fenton (1937) to Cruziana irregularis.Bergström (1976) recorded Rusophycus cf. R. jenningsifrom the Cambrian–Ordovician sequence of ConceptionBay in eastern Newfoundland, Canada. He argued thatthese traces were made by trilobites. Fenton and Fenton(1937) stated that it appeared that R. jenningsi of EarlyCambrian age fromAlberta, Canada, had been created bya trilobite belonging to the genus Olenellus. Jensen(1997) recorded R. jenningsi comparable to those fromthe Early Cambrian of Alberta (figured by Fenton andFenton, 1937), from the Lower Cambrian MickwitziaSandstone in south-central Sweden. Jensen (1997) didnot suggest possible tracemakers. Since the SnowyPlains Formation at Fish Hill is suggestive of a fluvialsystem, a nontrilobite arthropod is likely to haveproduced the Fish Hill R. jenningsi. This is the firstoccurrence of this ichnospecies in Australia.

4.15. Rusophycus isp.

Material: Examples of Rusophycus isp. on NMVP311632 and NMV P311635, Figs. 3B and 4A.

Fig. 7. Long and slightly curving trace fossils tentatively assigned to Undichnus previously identified in outcrops along the Broken River, slightlydownstream from where the new Mansfield–Tolmie Road bridge crosses the Broken River and close to where Platicytes, described herein, wererecorded (A. Warren pers. comm. 2003).



Locality and horizon: Fish Hill, in the Home StationSandstone Member in the Snowy Plains Formation. Thisichnotaxon is abundant in floodplain and the riverchannel deposits, and recorded from the same horizonsas actinopterygians, gyracanthid and dipnoan fish.

Description: Short, with a length-to-width ratio of lessthan 2:1, small, interfacial bilobate burrows resembling acoffee bean, preserved in convex hyporelief. The ovatelobes are parallel to slightly divergent. The mediumfurrow is moderately developed and continues down thelength of the ichnofossil, and the lobes are unornamented(after Keighley and Pickerill, 1996). Rusophycus fromFishHill varies in size from2 to 5mm long, and 2 to 4mmwide.

Remarks: Trace fossils on NMV P311632 and NMVP311635 are assigned to the ichnogenus Rusophycus, asthey are short and bilobate, although poor preservationrestricts assignment to species. Rusophycus isp. is themost abundant ichnotaxon, occurring in both shale andsandstone horizons throughout the Fish Hill sequence,and outcrops along the Broken River. It was found inassociation with Margaritichnus, Fuersichnus, and Sa-gittichnus. Since the Snowy Plains Formation at Fish Hillis suggestive of a fluvial system, a nontrilobite arthropodis likely to have produced the Mansfield Rusophycus isp.

4.16. Sagittichnus Seilacher, 1953

Seilacher (1953) established Sagittichnus linki todescribe a trace fossil suggestive of an arrowhead with amedian keel. They were thought to represent cubichnia(resting trace) with the producer unknown.

4.17. Sagittichnus lincki Seilacher, 1953

Material: Many specimens preserved on a red shaleblock, NMV P311632 (Fig. 3).

Locality and horizon: The specimens were collectedfroma horizon onFishHill in theHomeStation SandstoneMember in the Snowy Plains Formation. This bed isinterpreted as being characteristic of a low-energy flood-plain environment, and is assigned to the Rusophycusichnocoenosis. Actinopterygians, gyracanthid, anddipnoan vertebrates were also collected from this horizon.

Description: Horizontal to the bedding plane,preserved in concave hyporelief, and slightly bilobateto elongate arrow-shaped traces 3 to 5 mm long, and 1 to3 mm wide. They show a range of morphology that islikely controlled by the sediment consistency at the timethey were made, which is a taphonomic control.

Remarks: The material from Fish Hill occurs on thesame slab as Fuersichnus and Rusophycus isp (Fig. 3).In several places it appears that the two Rusophycus andSagittichnus may grade into one another, depending onwhether the animal was deposit-feeding or hiding orresting.

This trace fossil was first recorded from the Carboni-ferous by Głuszek (1995), from the Upper Carboniferouscontinental deposits of the Upper Silesia Coal Basin,Poland. This is the first identification of this ichnogenusfrom Australia.

S. linki could have been produced by several differenttypes of arthropods at Fish Hill. Recent neoichnologicalstudies, however, indicate that ostracodes are a likelyconstructor of this ichnotaxon in freshwater environments

Table 2Summary of the preservation, ethologic types, trophic types, tentative tracemakers, and associated figures of ichnotaxa described from the HomeStation Sandstone Member of the Snowy Plains Formation, Mansfield Basin, Victoria

Ichnospecies Preservation Ethology Trophic Type Tentative tracemaker Figure

Cruziana problematica Convex hyporelief Repichnia Deposit feeder Arthropod 2Fuersichnus commerfordi

isp. n.Convex hyporelief Fodinichnia Deposit-feeder Insect larvae 3

Margaritichnusmansfieldi isp. n.

Convex epirelief Fodinichnia/Domichnia orRepichnia?

Deposit feeder Worm or anemone/clam?

4

Rusophycusi isp. Convex hyporelief &epirelief

Cubichnia Deposit feeder Arthropod 3–4

Rusophycus biloba Convex hyporelief Cubichnia Deposit feeder Arthropod 2R. eutendorfensis Convex hyporelief Cubichnia Deposit feeder Arthropod 2R. jenningsi Convex hyporelief Cubichnia Deposit feeder Arthropod 2Sagittichnus lincki Convex hyporelief Cubichnia Unknown Unknown 3Palaeophycus tubularis Convex hyporelief Domichnia Suspension feeder or predaceous

organismArthropod 5

Platicytes lioparadus isp. n. Convex epirelief Fodinichnia/Domichnia Suspension feeder or predaceousorganism

Arthropod 6

Cubichnia (resting or nestling), repichnia (crawling or walking), domichnia (dwelling), fodinichnia (feeding traces).



(Retrum et al., 2005). In laboratory experiments,ostracodes force themselves into the sediment so thatthey are in a position of variable depth beneath thesediment surface. To extract themselves, they extend theirappendages outward to the surface and appear to pullthemselves out of the sediment. Such behaviour hasproduced slightly bilobate to arrow-shaped traces. Similartrace fossils have been described from Carboniferousdeposits of England interpreted as an ephemeral lake(Pollard and Hardy, 1991).

4.18. Undichna Anderson 1976

Anderson (1976) established Undichna for single orhorizontal sets of wave traces interpreted as fish trails.

Material: The material was previously located in thesame area as the current study, but was unable to berelocated. It is described from a photograph taken at thatoutcrop (Fig. 7).

Locality and horizon: Long and slightly curving tracefossils that appear to be preserved in convex hyporelieftentatively assigned to Undichnus were previouslyidentified in outcrops along the Broken River, slightlydownstream from where the new Mansfield–TolmieRoad bridge crosses the Broken River and close to wherePlaticytes, described herein, were recorded (A. Warrenpers. comm. 2003). The block also contains what appearto be large sets of desiccation cracks that crosscutthese traces and support our interpretation that they arepreserved in convex hyporelief. They were collectedfrom a horizon in the Home Station Sandstone Memberin the Snowy Plains Formation.

Description: Horizontal trials preserved as convexhyporelief consisting of asymmetrically arcuate, dis-continuous curving traces where the crests of the trail areirregular. Random overlap of traces is common on theblock. Each trace is approximately 200 mm long and∼10 mm wide at its widest point. The gentle slopingportion of the arcuate shape has an angle of ∼10–15°,whereas the steep side has an angle of∼65–75° (Fig. 7).In some cases, only the nearly straight portion of thetrace is preserved. In other examples these tracescontinue to recurve on themselves to form parallelridges. Amplitude and wavelength, however, cannot bedetermined because of the lack of a sinusoidal pattern.

Remarks: The Mansfield traces fossils are verysimilar to the lateral levees of Undichna uniscaladescribed by de Gilbert et al. (1999, fig. 4c and e) inLower Cretaceous lacustrine deposits in Spain. Furtherichnotaxonomic assessment of the Mansfield Undichnais not possible because the outcrop specimen could notbe relocated. This trace fossil was likely made by the

dorsal or pectoral fins of a fish swimming close to thechannel bottom prior to subaerial exposure thatproduced the desiccation cracks, which crosscut thearcuate trace fossils. While it is not possible to ascertainwhich fish taxon made the traces, it is possible that suchan acanthodian as Gyracanthides murrayi with pectoraland pelvic fin spines produced the traces with sweepingmotion.

5. Discussion

Trace fossils are important indicators of biologicalactivity in depositional environments (e.g., Häntzschel,1975; Ekdale et al., 1984; Miller, 2007). They areparticularly useful in continental deposits where thepreservation potential for fossil remains is relatively low(e.g., Chamberlain, 1975; Bown, 1982; Hasiotis, 2002,2004). Ichnofossils preserved at the Fish Hill localityindicate that likely a variety of noninsect arthropods,insect larvae, ostracodes, worms, and clams were present(Table 1) even though no invertebrate body fossils wereidentified from the Snowy Plains Formation. Our studyillustrates how ichnofossils can be used as a proxy forbiodiversity when the animal itself is not preserved(Hasiotis, 2002, 2004), particularly if they contain uniquecharacteristics definitive of a specific form of organism.

The depositional environments represented on FishHill are characterised by intervals of relatively low-energy conditions when clay accumulated, punctuatedby periods of higher energy when sand was deposited(Garvey, 2005; Table 1). Trace fossils identified fromthese different environments reflect these energyfluctuations with three ichnocoenoses identified: Ruso-phycus, Margaritichnus, and Cruziana (Garvey, 2005;Tables 1 and 2). These assemblages of ichna areconsidered ichnocoenoses because they represent morelocalized spatial and temporal occurrences of trace fos-sils. The Rusophycus and Cruziana ichnocoenoses aremost common in low-energy environments, indicatingthat the animals responsible for these traces preferredslow moving or still water. They are also present in thetwo higher energy facies, however, indicating that afterthese beds were deposited there must have been arelatively quiet period allowing invertebrates to colonisethe sediments. The high density of Rusophycus andCruziana from Fish Hill suggests that the environmentmust have been suitable to support large amounts ofbenthic activity.

While such trace fossils as Rusophycus and Cruzi-ana have been identified from both marine andcontinental environments, they may have entirely dif-ferent meanings due to the different lifestyle, adaptive



requirements, and trophic requirements of the trace-makers (Tevesz and McCall, 1982; Hasiotis and Bown,1992; Hasiotis, 2002). Continental (freshwater andterrestrial) environments experience extreme fluctua-tions in water level, temperature, clarity, salinity, oxygenrichness, and depositional energy levels with changes inweather conditions, temperature, and seasonality uniqueto these environments (Tevesz and McCall, 1982; Ward,1992; Hasiotis, 2002, 2004). Variations in these factorsaffect the distribution of organisms and the ichnotaxaproduced by their activity.

Ichna described here also have palaeoecologicalsignificance to the continental deposits at Fish Hill. Theterrestrial and freshwater–aquatic ecosystems are likelyto have contained abundant detritus and algae fromprimary production based on the abundance of deposit-and suspension-feeding ichna represented by Cruziana,Margaritichnus, Rusophycus, and Sagittichnus pre-served in the red mudstone and sandstone. Such detritusand primary productivity would have been necessary tosupport the nektonic and benthic invertebrates thatproduced the majority of trace fossils described fromFish Hill. The overall amount of organic matter thataccumulated in the system, however, would have beenless than that found in green or black siltstones andshales and coal deposits. The majority of organic matterat Fish Hill was likely consumed and oxidized by thebiota and environmental conditions. Such simpleburrows as Palaeophycus and Platicytes likely representpredaceous invertebrates that feed on the deposit andsuspension feeders. Predaceous arthropods would havelived with the herbivorous and saprophytic invertebratesinhabiting the sediment. Nearly all of the invertebratesthat constructed the fluvial trace fossils likely served as amajor food source for the diverse fish fauna in theseEarly Carboniferous aquatic systems. The fish fauna,represented by body fossils, would have been sustainedon aquatic and terrestrial arthropods as they are inmodern systems (Ward, 1992). The relationship betweenthe fish fauna and bottom sediments is also representedby the ichnotaxon Undichna; however, since it couldnot be relocated in the outcrop it is described from aphotograph.

6. Conclusions

Nine ichnotaxa are identified from the LowerCarboniferous (Tournaisian) Snowy Plains Formationat Fish Hill. Trace-fossil assemblages are dominated bycubichnia (Rusophycus, Sagittichnus), repichnia (Cruzi-ana as well as pasichnia), fodinichnia (Margaritichnus,Fuersichnus), and domichnia (Palaeophycus and Plati-

cytes). A summary of the ichnotaxa, their ethologictypes, trophic types, and the possible animals respon-sible is presented in Table 2. Based on these ichnotaxaand the corresponding deposits, three ichnocoenoses areidentified: the Rusophycus, Margaritichnus, and Cruzi-ana (Garvey, 2005, 2006).

This study of the ichnofauna from the Home StationSandstone Member of the Snowy Plains Formationextends the known distribution of several ichnotaxa andidentified of three new ichnospecies. Fuersichnus haspreviously been confined to the Triassic of EastGreenland and eastern Canada (Bromley and Asgaard,1979; MacNaughton and Pickerill, 1995), the Jurassic ofNew England and the Rocky Mountain region (Gier-lowski-Kordesch, 1991; Hasiotis, 2004), and the LateCretaceous of Antarctica (Buatois, 1995). The identifi-cation of F. commerfordi from the Early Carboniferousextends the geological distribution of this ichnogenus,the first occurrence of the ichnogenus in Australia, andrepresents a new ichnospecies. The identification ofM. mansfieldi extends the known number of ichno-species and represents the first occurrence of theichnogenus in the Carboniferous, and the second recordof it in Australia. Besides broadening the distribution ofFuersichnus and Margartichnus, Rusophycus biloba,R. eutenodorfensis, R. jenningsi, and Sagittichnus linckiare all recorded for the first time from Australia. Plati-cytes lioparadus is established for the first time.

Identifying trace fossils from Fish Hill may beimportant in establishing biogeographical relationshipsin the Late Palaeozoic. Aceñolaza and Buatois (1993)examined the ichnotaxa recorded from Late PalaeozoicGondwanan continental deposits, which reflected a highdiversity of Gondwana freshwater biota. The Fish Hillichnotaxon Palaeophycus tubularis shares some simila-rities with the Carboniferous deep lacustrine and thePermian fluviolacustrine trace-fossil assemblages fromArgentina, suggesting that a similar type of arthropodcould have had a wide Gondwanan distribution in theLate Palaeozoic.

Trace fossils in Carboniferous continental depositsare also important because the body-fossil record ofinvertebrates representing components of terrestrial andfreshwater ecosystems is depauperate at this time (e.g.,Grimaldi and Engel, 2005; Stanley, 2005). Invertebrateichnofossils indicate behaviour in response to stimulifrom the local environment, including depositionalenergy levels, sediment texture, sediment stability andconsistency, food availability, and oxygen levels (Carbo,1979; Hasiotis, 2002, 2004). Such palaeoenvironmentaland palaeoecological information is difficult to extractfrom continental deposits without the use of trace fossils.



Acknowledgements

We thank Anne Warren, Lisa Masini, Kate Parker,Sarah and Hamish Frankland, Gabrielle Archard, Caro-line Northward, Brian Woods, Kat Pawley, Travis Duka,Mike Sparg, Owen Welsh, Jacob Fitzgerald, MonicaYeung and Bruce Loomes of ‘Gondwana Dreaming’,and Andrew Hoogenraad and students from CamberwellGrammar School for assisting JMG with fieldwork. Weare grateful to the Commerford families for access totheir properties. We thank Jordi de Gilbert for providingreprints to us of his research on Undichna, and AnneWarren for the photo of Undichna from the MansfieldBasin. Two anonymous reviewers and Anne Warren arethanked for their comments, which much improved thismanuscript. JMG acknowledges support from a LaTrobe University Postgraduate Scholarship.

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