Trace Fossils in Coal-Bearing Sequences

Journal of the Geological Society, London, Vol. 145, 1988, pp. 339-350, 5 figs. Printed in Northern Ireland

Trace fossils in coal-bearing sequences

J. E. POLLARD Department of Geology, University of Manchester, Manchester M13 9PL, UK

Abstract: Inthe past decade trace fossils have been recordedextensively from coal-bearing sediments, differingwidely in facies, age and location.Westphalian or Stephanian‘coal-measures’ in Britain,Europe and EasternCanada contain an ichnofaunaproduced by invertebrates and/or vertebrates in upper delta plain sediments. This contrasts with the marine-related lower delta plain ichnofaunas known from Pennsylvanian rocks of the United States and Permian Gondwana ‘coalmeasures’ of South Africa. Deltaic complexes of Middle Jurassic age inthe North Sea basin and UpperCretaceous age in NorthAmerica contain marine trace fossils and dinosaurfootprints in coastal coal-bearing facies. Thesecase histories illustrate the importance of trace fossils both in facies analysis of coal-bearing sequences and in recording the presence of animals rarely known as body fossils in such clastic sediments.

The recognition and use of trace fossils in the sedimen- to the diverse studies presented at the Egham Symposium tological and palaeontological analyses of coal-bearing and appearing in its publications. (Scott 19870, 19876). sediments is a major advance of the past decade. Although distinctive burrows, tracks and trails have been known from Silesian coal measures Euramerica Westphalian rocks since the mid 19th century, for example of fromLancashire (Binney 1852) and the Maritime Provinces, Canada (Dawson 1868), little notice was taken of Westphalian coal measuresof Britain them until the 1950s and 1960s. Over this period German Atthe previous British symposium on ‘Coal and workers established that trace fossil associations in the Ruhr Coal-BearingStrata’, Calver (196th) reviewed the trace CoalMeasures paralleled fauna1 and sedimentary cycles fossils then recognized in the Coal Measures of Britain. He (Jessen 1949; Jessen et al. 1952; Seilacher 1963, 1964). concluded that they demonstratedthe presence of the From this beginning we may recognize the two-fold Planolites ophthalmoides assemblage inmarine bands and development of moderntrace fossil studies, firstly in the P. montanus assemblage in non-marine strata as known conjunction with sedimentological or facies analysis of from theRuhr (Seilacher 1964), but totalling only four coal-bearing sedimentsand secondly, in recording the ichnogenera. More recent work in the Lancashire, presence,behaviour patternsand environmental distribu- Yorkshire and East Midlands coalfields (Eagar et al. 1985; tions of the animals producing the traces. Guion 1985; Pollard 1986) established the presence of more Detailed stratigraphical and sedimentological analysis of than fifteen ichnogenera, in five or morerecurring Silesian deltaicsequences of both Britianand North associations; these associations may be correlated with America in the past decadedemonstrated a close sediment type and specific sedimentary environments (Fig. relationship between trace fossil assemblages, sediments and 1). Thesediverse ichnofossils form part of aseries which environmental models (Hakes 1977; Archer & Maples 1984; first appeared in interchannel strata of late Namurian sheet Eagar et al. 1985). Comparable results are being obtained by deltas of the Pennine area (Eagar et al. 1985, figs 5 and 8) workers on Gondwanacoal-bearing facies andNorth and they constitute a distinct lower delta-plain ichnofauna of American Cretaceous sequences (see below). Recent major largely Westphalian Aage. Figure 1 relatesthese trace publications on ichnology of marginal marineand fossils to specific subaqueous delta plain environments. The continental environments (Miller et al. 1984), on the use of traces reflect the variety of producersand differing biogenic structures in interpretingdepositional environ- behavioural responses to sedimentation conditions in the ments (Curran 1985) and on trace fossils in cores respective environments. (Chamberlain 1978; Ekdale et al. 1984) all contain Muddy sediments of marinea band contain both coal-bearing sediment case-studies. The relative abundance Ophthalmidium [‘Planolites’] ophthalmoides (Calver 1968b; of good quality corematerial produced during coalfield Rippon 1984) and newly recognized Lingula burrows, exploration can reveal a wealth of trace fossil information to Lingulichnus, in the regressive phase of the vanderbecki the trained observer. MarineBand in Lancashire (Fig. 2a). The ichnofauna of Palaeontologistsinterested in the ethological aspect of laminated muddy sediments of the interdistributary bay, or trace fossils have produced new information on diversity and floodbasin lake in higher Westphalian sediments, is the P. environmental distribution of both arthropods and tetrapods montanus association, (Fig. 2b) including Cochlichnus, by the detailed analysis of their trackways. Such ‘Gyrochorte’ carbonaria, Arenicolites, bivalve resting traces information is complementary tothe very limited data (Pelecypodichnus), and rare arthropod resting or crawling available from body fossils of these animals in clastic traces (Selenichnus and Acripes) (Rippon 1984; Guion 1985; coal-bearing sequences. Eagar et al. 1985; Pollard 1986; Romano & Whyte 1987). This paper reviews briefly some of these discoveries and The greatest diversity of traces is recorded from crevasse developments of the past decade particularly in relationship splay sheet sands (Fig. 1) wheretraces frequently show a

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Trace fossil associations T I

Lithology and I sedimentaryenvironment

I T

I I I

I I 7 I l Crevasse splay I I -1-

mdallt l m ..l fig. 1. Diversity, associations and facies relationships of trace fossils in Westphalian Coal measures of Britain. 1, Ophthalmidium association; 2, Planolites montanus association; 3, Arenicolites carbonarius association; 4, Pelecypodichnus association; 5, spreiten burrows; 6a, aquatic arthropod traces; 6b, terrestrial arthropod traces; 7, vertebrate tracks. Key to lithologies, see Fig. 4.

behavioural response to sedimentation. Burrows of suspen- 1983, 1985; Hardy 1970a; Eagar et al. 1985) (association sion feeders, Arenicolites carbonarius and Polykladichnus 6a). The sinuoussurface trail or burrow Cochlichnus has are usually initiated from the topof the crevasse splays (Fig. recently been the subject of behavioural and mathematical 2c, e). Resting burrows Pelecypodichnus, preserved on the analysis (Elliot 1985). It was probably formed by nematode base of the crevassesplay sands frequently show current worms duringlocomotion or feeding,sometimes on response orientation or are associated with vertical escape decaying bivalves (Hardy 1970b). shaftsindicating behavioural response to seasonal flood The diverseichnofauna of the WestphalianCoal sedimentation (Hardy & Broadhurst 1978; Broadhurst et al. Measures of Britainillustrates clearly both the facies 1980) (Fig.2d). In channel fill sequencesbioturbation is significance and behaviouralaspects of trace fossils. limited to finer sediments although Arenicolites, Pelecypod- However it is remarkably different from contemporaneous ichnus and Planolites arerecorded (Guion 1985; Fielding ichnofaunas of coal-bearingsediments of the Canadian 1986; H. Williams pers.comm. 1986), perhaps associated Maritime Provinces. (Fig. 3). with starvation of active channel fill. Rare horizontal U-shapedor spreiten burrows (association 5) have been recordedfrom channel floor deposits or leveesediments Westphalian sequencesof eastern Canada (Hardy 1970b; Eagar et al. 1985) (Fig. 2f). The classic 'Joggins Section' of coal-bearing strata of the Trackways of terrestrialarthropods Diplichnites, Cumberland Basin of Nova Scotia has long been famous for Paleohelcura (association6b) tetrapodor footprints its fossils, especially vertebrates and upright trees. However (association 7)are not known from CoalMeasures of it also contains a variety of arthropodand vertebrate Britain below the Keele Beds (Westphalian D or trackways figured by Dawson (1868, 1873), suggestive of a Stephanian)but diversebehaviour patterns of aquatic or diversetrace-producing freshwater and terrestrial fauna. semi-aquatic belinurids (limulids) including burrows, resting Thevertebrate footprints show a predominance of traces and locomotion trackways are known (Chisholm amphibians,microsaurs (but also rhachitomes),and rarer

Fig. 2. Burrows from British Westphalian sediments (a-e are cut vertical sections or core section). (Scale bars are 10 mm long.) (a) Lingulichnus sp. Vertical burrows with variable diameter and thick mud-lined walls inlaminated siltstone. uanderbecki Marine Band, Westphalian A/B, Lancs. MGSF 100 (Manchester Geology Department Special FossilCollection) (b) Planolites rnontanus (Richter). Sand-filled irregular horizontal burrows in laminated siltstone. Westphalian A, Lancashire. MGSF 101. (c) Arenicolites carbonarius (Binney). Mud-filled, small U-shaped burrows without spreiten. Westphalian A, Elland, Yorkshire. Manchester Museum L6211. (a) Pelecypodichnus and bivalve escape shaft. Cross section (V-shaped) of bivalve resting burrows which are repeated vertically by upward escape. Westphalian A, Up Holland, Lancashire. MGSF 102. (e) Polykladichnus sp. Small upward Y-branching burrows in ripple cross-laminated fine standstone. Westphalian A, Up Holland, Lancashire. MGSF 103. (Q Rhizocorallium cf. R. jenense. Tongue- or pouch-shaped or short oblique U-burrows with scratched wall mouldings. Westphalian A, Goyts Moss, Derbyshire. MGSF 89.

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~ ~~ S. AFRICA Permian' X X xxxxxx ?

FRANCEStephanian' X XL.S X X

BRITAIN Stephanian xs X X

BRITAINWestphalian4 xxxxxxxx X xxx S

RHUR Westphalian' X X xx

CANADA Westphalian6 xx X X XL,SX X X

~ TENNESSEE X xx X X xxxx ~ennsy~vanian' INDIANA xx X X xxx X Pennsylvanian' 1 KANSAS ~ennsy~vanian~xxxxxx xx X xxx X

fig. 3. Diversity and occurrence of trace fossils in Upper Carboniferous and Permian coal-bearing sequences. Authors cited: (1) Stanistreet et al. 1980; Mason 1985, (2) Langiaux & Sotty 1977, (3) Author's observation, (4) Eagar et a[. 1985 and author, (5) Seilacher 1964, (6) Author's compilation, (7) Miller & Knox 1985, (8) Archer & Maples 1984, (9) Hakes 1976. L, S, large or small varieties of trackways.

reptiles, possibly cotylosaurs and pelycosaurs (Haubold ichnofaunal diversity of Stephanian intermontane basins of 1984, figs 43, 45, 46). They compare closely with the skeletal Europe (Fig. 3). fauna recovered from the lycopod tree stumps (Carroll et al. 1972), suggested by Milner (1987) to be a terrestrial fauna. Dawson (1873) and Ferguson (1966, 1975) both figured large Stephanian of Europe trackways of Diplichnites from Joggins, probably made by Diverse arthropodand tetrapod trackway ichnofaunas are the largemyriapod Arthropleura. A similar sinuous known from two Stephaniansequences in Europe, trackwayfrom contemporaneous sediments of Gardener Blanzy-Montceau (in the Massif Central of France) and the Creek, New Brunswick was shown by Briggs et al. (1984) to Keele Beds of the English Midlands. The Blanzy-Montceau have been made by an individual Arthropleura about 1.8 m coal-bearing succession is in an intermontane basin bounded long winding its way between erect Calamites stems on a by active faultmargins, which produce synsedimentary vegetated subaerially-exposed crevasse splay. debris flows. Coals, lacustinedeposits and alluvial plain Other arthropod trace rossils from Westphalian rocks of deposits with meandering channel fills (Courel 1983, 1988) the Maritimesinclude possible terrestrial scorpionid or are also known. A very rich fauna of 'Mazon Creek' type eurypteridtrackways (Paleohelcura), diverse aquatic crus- including diverse arthropods,vertebrates (both fish and tacean traces (Zsopodichnus, Acripes, cf. Protichnites) and tetrapods), worms and bivalve molluscs occurs in ironstone belinurid walking trackways (Kouphichniurn) (Dawson nodules from the lacustine beds at this locality (Rolfe et al. 1868; Matthew 1910; Copeland 1957; Pollard 1985). 1982). Burrows are rare in Westphalian coal-bearing sediments Arthropod trackways include large and small of the Maritimes (Fig. 3). althoughlarge Beaconites 2nd Diplichnites, cf. Protichnites and Paleohelcura (Langiaux & small Planolites occur in overbank sediments of the Port Sotty 1977) reflecting arthropleurid and scorpionid Hood Formation (E. S. Belt pers. comm. 1982). No marine producers, the presence of which is confirmed by the nodule burrows, bivalve resting or escapetraces are known fauna. Diverse tetrapod footprints include amphibians (compare British andCanadian Westphalian diversity on (microsaurs,eryopsoids and dissorophoids)and reptiles Fig. 3). These absences are reflected by the body fossils as (romeriid cotylosaurs and possibly pelycosaurs; Langiaux & well, for no marine faunas are known from the Maritimes Sotty 1977; Gand 1975; Haubold 1984, figs 52-54). andthe anthracosiid bivalves which produce mostescape Although full analysis of trace fossil to sediment shafts, die out early in Westphalian A (Rogers 1965; Vasey relationships in this sequence has not been carried out, the 1984). The ichnofauna reflects a basinal environment, trackways of both terrestrial arthropods and tetrapods occur predominantly upperdelta plain or alluvial plain depo- in thinly-laminated siltstone or fine sandstone which formed systems in small basins, with axial-meandering fluvial in a lake margin environment. systems and marginal alluvial fans(J. H. Calder pers. Trace fossils from the Keele Beds were collected from a comm. 1986). There is a remarkable similarity tothe quarry at Alveley,Shropshire by F. W. Raw in the 1920s

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and are preserved in the Geological Sciences Department limestones (see Hakes 1976, fig. 2). The ichnofaunas of the Museum of Birmingham University. They were studied by grey shalemembers of four stratigraphically-separated Haubold & Sarjeant (1973) and contain at least six footprint megacyclothems differ in composition,although they are ichnogenera attributableto severalamphibians and a predominantly of marginal marine character (Fig. 3). They herbivorous pelycosaur. Arthropod trackways are of Acripes are notanalogous to the repetitive cycles of body fossils (Diplichnites?) type and Paleohelcura, which may indicate recordedfrom these megacyclothems (Moore 1966). freshwater crustacean and scorpionid producers. Furthermore, the trace fossils indicate that coal formation In a sedimentological analysis of the Keele Beds, Besly occurred on emergence within a regressive sequence prior to (1988) concluded that they are alluvial red beds with large a broad marine transgression and deposition of the overlying channel-fill sandbodies, alluvial palaeosols rooted but fossiliferous limestone (Hakes 1977). The only non-marine without coals, which gave way to caliche horizons. The trace fossils are crustacean tracks ('Diplichnites') and limulid ichnofauna is preservedas hypichnial trackways on fine tracks (Kouphichniurn; Fig. 3) which occur in the sandstone, formed in temporaryponds or channels which TonganoxieSandstone Formation, a fluvial or estuarine were subject to desiccation in a changing tropical sub-humid channel sand (Bandel 1967b; Hakes 1976). to semi-arid climate. The presence of large pelycosaur In a study of Pennsylvanian formations in south-western footprintsinthese beds is consistent with the first Indiana, (Archer & Maples 1984) documented the detailed occurrence of a reptilian fauna which later dominated the relationshipsbetween diversity and abundance of trace arid red beds of the succeeding Permain Rotliegendes facies fossils, sedimentsand interpreted environment (Fig. 4). (Haubold 1984). Trace fossils areabundant, but of limited diversity in limestones and shales of interdistributary bay origin, where fusulinids, brachiopods and bivalve molluscs occur. Similar Pennsylvanian coal basinsof the United States characteristics are shown by the ichnofaunas of tidal flats Information ontrace fossils occurring in Pennsylvanian anddistributary mouth bars, where resting and escape cyclothems of theUnited States is of two main types. structures predominate. The trace fossil sediment relation- Records and analyses of tetrapod footprints mostly from ships of these mouth-bar sandstones closely resemble those east coast and eastern interior coal basins date largely from in Namuriansheet-delta deposits in Britain (Eagar et al. the late 19th century. By contrast, very recentintegrated 1985). The greatest diversity of traces occurs in the trace fossil and sedirnentological analyses have been carried proximal flood-plain or lacustrinesediments (Whetstone out independently by severalworkers in widely-separated facies), which are overlain by seat-earthsand coals interior basins, such as in Kansas, Indiana and Tennessee frequently associated with rooted trees (Fig. 4; Archer & (Fig. 3). The tetrapod footprint record closely parallels that Maples 1984). However, these trace fossils are surface trails from Westphalian C to Stephanian C strata in Europe and and trackways of freshwater invertebrates, (predominantly Nova Scotia, showing a diversity anddominance of arthropods) and they show some common ichnogenera with amphibianfootprints but without any facies relationships the Dwyka glacial lake deposits of South Africa (Figs 3 & 4 being established (Haubold 1984). and see below).They may representa distinct, yet rarely Studies of sedirnents and trace fossils inKansas have preserved, late Palaeozoic freshwater ichnocoenosis. beencarried out by Bandel (1967a, 19676) andHakes The apparentlyanomalous occurrence in thesefresh- (1976, 1977, 1985). Trace fossils occur in both shaleand watersediments of 'deep marine'ichnogenera 'Paleo- sandstonemembers of classic Pennsylvanian cyclothems dictyon' and 'Spirodesmos' can beexplained as surface (Moore 1936, 1966) where coals with seat-earths occur in trails produced by superficially similar behaviour patterns in sequence with siltstones, black shales and fusulinid broadly similar substrate types and low energy environmen- ISedirnents

- .. Fig. 4. Diversity and facies relationships of trace fossils in Pennsylvanian sediments of Indiana, U.S.A. (Archer & Maples 1984).

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tal situations. Triassic fluvio-lacustrine sequences in Bntain montanus type and possible escape structures (Salamuni & also containbehavioural variants of Treptichnus and Bigarella 1967). Similar Planolites bioturbation, together Plantichnus which resemble ‘Paleodictyon’ or ‘Spirodesmos’. with Skolithos and possible bivalve escapeshafts are Hardy (1970~)recorded crude polygonal networks resem- illustrated by Kar & Chaudhuri (1981) fromthe Barakar bling ‘Paleodictyon’ produced by intersection of fine Formation of the Lower Gondwana age Raniganj Coalfield Cochlichnus trailsBritishin Westphalian non-marine of India. Such subjective assessments suggest that broadly sediments. Such occurrencesillustrate thedanger of similar trace fossil and facies relationshipsas described uncritical assessments of ichnofaunal lists or tables. above forEuramerican Westphaliansequences may exist The diversity and distribution of trace fossils in widely in Gondwana coalmeasures sequences. Such Pennsylvaniancoal-bearing rocks of Indiana contrast with relationshipshave been demonstrated recently from the those of Tennessee. In Tennessee the coals were formed in Karoo Supergroup of South Africa. back-barrierenvironments orabandoned tidal or dis- tributarychannels andthe overlying trace-fossil-bearing clastics were deposited in a wave or tide dominated deltaic Ecca Group of South Africa system (Miller & Knox 1985, fig. 3). Thetrace fossils There is a very distinct difference between the trace fossil presentare mainly marine ichnogenera or facies crossing assemblages of the glacial Dwyka depositsand the types and exclusively non-marine tracesare absent. succeeding coal-bearing Vryheid Formation (=Middle Ecca) However,some marginalmarine to non-marine facies- in the lower part of the Karoo Supergroup of South Africa. crossing forms (Cochlichnus,Palaeophycus, Kouphichnium The fish traces and diverse arthropod trackways of the and Pelecypodichnus) do occur insandstones of the Dwyka varvites (Savage 1971; Anderson 1975, 1976, 1981) overlying Rock CastleConglomerate Formation, which are similar to those from other late Palaeozoic freshwater could have been deposited incoastal low-sinuosity stream lacustrine sequences (cf. Archer & Maples 1984 and below). systems subject to localized incursion of marine waters These are absent from the Vryheid Formation. (Miller 1984). Trace fossils and associated sediments of the Vryheid Despite the local palaeogeographical and sedimentologi- Formation were first studied in northern Natal by Hobday & cal differences between the Pennsylvanian coal-bearing Tavener-Smith (1975), who recognized a changefrom sequences of these three interior basins it is apparent that Nereites ichnofacies to Skolithos ichnofacies in an upward- they show a common character of marine dominance of the coarsening regressive deltaicsequence. Although this trace fossil assemblages, which contrasts with the Westpha- pioneer bathymetric model was too simplistic, it has been lian andStephanian ichnofaunas of Europe (Fig. 3). refined by independent studies in both the Transvaal and However, they are much closer in the ichnofaunas and Natal coal-basins. Stanistreet et al. (1980) showed that sedimentary character to the earlier Carboniferous deltas of distinct trace fossil associations characterize prodelta slope, Yoredaletype (late Dinantian to earlyNamurian) and lower delta-plain and delta abandonment-barrier and marsh Namurian sheet deltas of Britain (Eagar et al. 1985; Pollard facies in the Transvaal coalfields. Most of the ichnogenera 1986). Recent work by George Lees (pers comm. 1986) on are marine-related in terms of comparable European theYoredale cyclothemic deposits of N Englandand S Silesian coal-bearing sequences (Fig. 3). Very similar facies Scotland shows a low diversity of tracesin non-marine relationships have been established in the Natal coal-basin delta-top environments, which are frequently coal-bearing, where Helminthopsis, Diplocraterion,Skolithos, but a high diversity of ichnogenera exists in the marine Siphonichnus, Spirodesmos, Teichichnus, Helminthoida?, influenced lower delta-plain ordelta front environment Planolites, Tigillites and Monocraterion occur in sediments (Pollard 1986; Lees 1986). Many of these ichnogenera are interpreted as having formedin a regressive deltaic also common in American Pennsylvanian sequences. The sequence.This included prodelta, distributarymouth bar, gradual decline of marine influence and development of interdistributary bay and distributary channel environments facies-related delta-plain ichnofaunas recently demonstrated (Turner et al. 1981; Mason et al. 1983; Mason & Christie in the Namurian to Westphalian Pennine Basin sequences in 1986; T. R. Mason pers.comm. 1986). Coalformation Britain (Eagar et al. 1985; Pollard 1986) hasnot been apparentlyoccurred eitheronthe lower delta plain recognized in the Pennsylvanian basin sequences of the associated with restricted bay environments or on alluvial United States. As yet there are few common characteristics plain associated with fluvial flood basin deposits. Distinct apparent in the trace fossils of crevasse splay, flood basin trace fossils have not been recognized from fluvial deposits lake and fluvial channelsediments, often associated with exceptfor limited bioturbation (I.G. Stanistreet pers. coals, in respectively the British Westphalian and American comm. 1986). Pennsylvanian sequences (cf. Figs 1 & 4). The abovestudies havebeen of considerable importance in establishing the probable marine nature of the water in the Karoo Basin in Permian times, in the absence Late Carboniferous-Permian coal measures of of body fossil evidence. Using everydetailed analyses of Gondwana Diplocraterion populations Mason & Christie (1986) even The major exploration and exploitation of the Gondwana claimed todetect colonization and extinction of infaunal coal measures of the southern hemisphere continents in the burrowers related to salinity changes and flood sedimenta- past decade hasled tothe discovery of trace fossils. tion in shoal water delta complexes. Here again there is a Surprisingly littlehas been recorded yet fromAustralia remarkable similarity of ichnogeneric composition between (McCarthy 1979; Johnson 1984; Hobday 1987). sediments of the ‘Yoredales’ of Britainand the Vryheid Although not analysed in detail, cores of interlaminated Formation of SouthAfrica, interpreted as having been fine sandstoneand siltstone fromthe Rio Bonito Coal deposited in prodelta, mouth barand lower delta-plain Measures of Brazil apparently show burrows of Planolites environments (compareTurner et al. 1981; Mason &

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Christie 1986 with Pollard 1986; Lees 1986). The absence of (Sarjeant 1974) shows several types of tridactyl dinosaur trace fossils from the fluvial sediments of the Ecca Group footprintsin bioturbated sandstones and shales associated may be related tothe immaturity of thesesediments, or with desiccation cracks. Even here however, the sediments perhaps tothe low temperature of these ephemeral yield marinemicroplankton (acritarchs and dinocysts) post-glacial environments. At higher horizons in the Karoo suggesting that they formed in either a saline swamp on a Supergroup(Beaufort Groupand Stormberg Group river-dominated delta plain (Fisher & Hancock 1985) or in a Triassic) both invertebrate and vertebrate traces are known coastal alluvial plain subject to tidal flooding (Leeder & fromsediments diagnostic of semi-arid fluvial and lake Alexander 1985). Similar dinosaurfootprints have been environments (Van Dijk et al. 1978; Haubold 1984; Ostrom discovered recently in both the braided channel facies of the & Galton 1984; Mason 1985). Within theKaroo Moor Grit Member and the meander belt facies of the Long Supergroup of South Africa, therefore, the study of trace Nab Member of the Scalby Formationat other localities fossil assemblages in relationship to sedimentology is playing (Delair & Sarjeant 1985). Broadly similar facies associations a major part in the reconstruction of faunal, environmental of dinosaur tracks have been described from the Cretaceous and palaeoclimatic changes of Gondwana and its borders. DakotaGroup of Colorado (Lockley 1985a, 19856;see below). Inthe past decade coal-bearingsediments of Middle Mesozoic coal-bearing sequences Jurassic age have been discovered to occur in sedimentary basins of the Northern North Sea especially in the Brent Group of the Viking Graben-EastShetland Basin. The Middle Jurassic of the North Sea basin and its margins sedimentaryenvironments of thesesequences have been Although coal has been known for more than two centuries deduced by analysis of their sedimentology and palynofacies in the MiddleJurassic deltaic rocks of Yorkshire, closely (Dennison & Fowler 1980; Proctor 1980; Handcock & similar sequences of coal-bearing strata have been Fisher 1981; Parry et al. 1981; Budding & Inglin 1981; discovered recently in several offshore areas of the North Johnson & Stewart 1985; Richards & Brown 1987). Sea basin. At least five geographically discretedeltaic Initially trace fossils wererecorded from the cores of complexes are present with different sedimentary these sediments in terms of general bioturbation or ‘worm sequences,but all broadlycontemporaneous. To aid the burrows’, or as ‘worm and shrimp burrows’. More recently interpretation of these deltaiccomplexes, there has been they have been used as a tool in more precise environmental renewedinterest in the classical Yorkshire coast sections, analysis, especially by Norwegian workers. R. Lundstrom et mainly from a sedimentological and palynofacies approach, al. (pers.comm. 1986) reportedon thesecoal-bearing with only minimal attention paid tothe containedtrace sequences and recorded detailed trace fossil distributions in fossils. However, the most recent work on coal-bearing relationship to sedimentaryenvironment in the Viking sequences from the Norwegian offshore areas has included Graben and Hammerfest Basin. In the Sleipner Formation some detailed recording and assessment of trace fossils in (Bajocian-Bathonian)they recorded thick coal seams of relationship to lithofacies and environmental interpretation. coastal ordelta plain origin and ichnofaunas including The Ravenscar Group (Middle Jurassic) of Yorkshire Ophiomorpha, Skolithos, Diplocraterion and Asterosoma contains coals, or at least rooted seat-earths, in most of the associated with facies interpreted as stacked barriers, tidal non-marine beds, although they were mined in the 18th and inlets and minor transgressive-regressive cycles. More than 19th century only fromthe Saltwick Formationand twenty ichnogenera were recorded from the Ness Formation GristhorpeMember (Hemingway 1974). The coals were of theOseberg and Gullfaks areas, although they were prospected by William Smith (Hemingway & Owens 1975). largely marine-related ichnogenera occurring in lagoonal or Thefour non-marinesequences separated by marine bay sediments. The Nordwela Formation (Lower Jurassic) incursion strata have long been known to contain marsh in the Troms I area of the Hammerfest Basin also contains and channeldeposits but recent sedimentological and coal-bearingsediments together with channelsandstones, palynofacies analyses (Livera & Leeder 1981; Hancock & coastal plain-tidal flat sediments and trace fossils including Fisher 1981) demonstrated a wide variety of deltaic Diplocraterion, Skolithos and Teichichnus.. environments. These includecoastal shallow marine, Work on coal-bearing sediments and trace fossils in the coastal-lower delta plain, upperdelta plain, fluvial and Brent Group in the British sector of the Viking Graben is marsh deposits. either at an early stage or not yet released. The ichnofauna Trace fossils have been described in detail only from the of the Rannock and Etive Formations includes Skolithos, marine members such as the Eller Beck Formation and the Teichichnus Terebellina and Palaeophycus in highly Scarborough Formation, although bioturbation is frequently bioturbated laminated sandstones which are interpreted as referred to from levee and upper delta plain sediments. Two shoreface storm deposits of a prograding barrier coast broadly defined trace fossil-facies associations are currently (Richards & Brown 1986, 1987). The coastal and delta plain recognized in marine or coastal sediments. The Scarborough complex of the Ness Formation comprises a wide range of Formationcontains Thalassinoides-Rhizocorallium-Teich- coal-bearing sands and muds with rootlets(Budding & ichnus-Asterosoma in a littoral to offshore association, Inglin 1981; Johnson & Stewart 1985; Brown 1985). More and an Arenicolites ichnofauna in shoreline or beach sands than seven facies associations havebeen identified and of the incursion phase(Farrow 1966). The Eller Beck several recurrenttrace fossil/sediment associations have Formation and Yons Nab beds are intensely bioturbated been observed personally in released cores. Diplocraterion locally by Diplocraterion and/or Chondrites. Non-marine polyupsilon occurs widely inlagoonal or interdistributary sediments rarely contain identifiable trace fossils except for bay muds or silts. The form of this U-burrow is remarkably dinosaur footprints which are particularly abundant in the similar to its expression in both Permian Vryheid Scalby Formation. The famousBurniston FootprintBed Formation of S. Africa (Mason & Christie 1986) and Lower

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Carboniferous deltaicsediments (Knox 1973). Planolites since the 1920s (Peterson 1924), they have been the subject montanus occurs in lagoonal or lacustrine shales and escape of a recent fascinating analysis by Lockley et al. (1983) and burrows with Pelecypodichnus profile in crevasse splays or Lockley (19866). Not only do the footprints show that the mouth bar sands. Such ichnofaunas appear closely similar to late Cretaceous peat swamps were extensively trampled by the P. montanus and Pelecypodichnus associations of dinosaurs (Lockley 1986a), but also that the producers were Westphalian deltas (Fig. 1). slow-moving hadrosaurs, travelling in herdsperhaps for Thus whilst trace fossils occur in association with Middle mutualprotection from contemporary carnosaurs, whose Jurassiccoal-bearing sediments both onshore and offshore footprints are associated with those of the herbivores. their detailedichnofacies relationships havebeen only Similar palaeobiological and palaeoenvironmental relation- partially analysed. The widespreadoccurrence of such ships may await discovery in dinosaur footprint beds of the marinetrace fossils as Diplocraterion, Skolithos, YorkshireJurassic (Delair & Sarjeant 1985) or in the Ophiomorpha, Teichichnus and Asterosoma suggests per- Westphalianand Stephanian amphibian and reptilian hapsclosera similarity tothe ichnofaunas of the tracksites of Europe or the Maritime Provinces (see above, marine-influenced Carboniferous deltas (e.g.Yoredales, Haubold 1984; Fichter 1982). Lees 1986) or PermianVryheid Formation ratherthan to Several major coal seams have been mined in the past the well-known Westphaliancoal measures (Fig. 2). century Horseshoefromthe Canyon Formation However, such conclusions may be premature aslittle (Campanian-Maestrichtian, Upper Cretaceous) of the detailed analysis is published and Westphaliananalogues EdmontonGroup in the Drumheller area of Alberta, occur. Canada (Allan & Sanderson 1945). Recentstratigraphical andsedimentological analyses (Shepheard & Hills 1970; Rahmani 1981) suggest that this formation was deposited in Upper Cretaceous of Western North America a tidally dominated delta-complexprograding south- Coal-bearing strata of late Cretaceous age have long been eastward fromthe uplifting Rocky Mountains intothe known and mined in the western United States (e.g. Utah epicontinentalseas of the Western Interior. The sequence andColorado) and Alberta in Canada.Recent detailed coarsensupwards and becomes fluvially dominated in the sedimentological analyses of these coal-bearing strata have higherparts with Scoyenia ichnofacies trace fossils included work on trace fossils, for example Utah, Colorado (Nurkowski & Rahmani 1984). The lower coals are and Alberta. Such studies have given new insights into the associated with bay fill, barrier to back-barrier facies and use of trace fossils for environmental and substrate analysis, tidal channel deposits (Bromley et al. 1984), although they and dinosaur locomotion and behaviour. may have formed as far as 40-50 km back from the active The Spring Canyon Member of the Upper Cretaceous shoreline as they are ash free (McCabe 1987). Trace fossils Blackhawk Formation of east-central Utah contains both presentare in two distinct facies associations; open marine and marginalmarine facies sediments,each Ophiomorpha-Teichichnus in barrier island sandstones with distinctive trace fossil associations(Howard & Frey (Rahmani 1981) and a Teredolites ichnocoenosis of bivalve 1984; Kamola 1984). In the marginal marine facies Kamola borings in the upper part of one of the coal seams (Bromley (1984) described sediment/ichnofaunal associations of et al. 1984). channels, deltas and lagoons. Coals are generally developed The Teredolites borings occur in the top 60 mm layer of a on top of channel fills, lagoon fills and delta sequences and coal seam 0.6-1 m thick, where it is eroded by the base of may be overlain by lagoon fills or cut into by channels. an 8 m deep tidalchannel which was subsequently mud Trace fossils in the channel fill sediments include filled. The distinctive borings havebeen moulded by the Ophiomorpha, Skolithos, Teredolites borings indriftwood overlying sandstone and ironstone and show two kinds of and Thalassinoides sueuicus networksmoulded ontothe sculpture, ‘xyloglyph’ or wood grain markings from the base of the channel fill sandstone, where the channelcut woody substrate and ‘bioglyph’ markings resulting from the into open tunnel systems in the underlying peat (coal). The rasping of the shell of the boring bivalve. Theseunique upwardcoarsening delta sequencescontain trace fossils characters of this flexible, combustible, yet biodegradable analogous thoseto in modern tidal flats including woody substratetogether with distinctive forms of the Ophiomorpha, Arenicolites, Pholeus (crab burrows)and borings led Bromley et al. (1984) to erect a new ichnofacies, Teredolites. The ichnofauna of the lagoon fill sediments the Teredolites ichnofacies, for such trace fossil assemblages (OphiomorDha. Thalassinoides, Planolites montanus. Pele- formed on shallow marine ‘woodgrounds’. cypodichnus and Palaeophycus) is likened by Kamola (1984) The significance and validity of this ichnofacies related to to that of modern lagoons or estuaries, but also shows some a peat or coal substrate may be assessed by consideration of traces similar to those in Westphalian lagoonal or bay fill similar communities of bivalve mollusc borings in sub-fossil sequencesdescribed above (Fig. 1). All thetrace fossils Holocene peat beds of theGerman North Sea coasts recorded here are of marine types, many being common in described by Schaefer (1972). Here raised bog, swamp the underlying and overlying open marine sequences. The forest or Phragmites peats, all formedbetween 7000- presence of T. sueuicus in the upper part of the coal seams is 10000 BP, areinterbedded with marinesediments. They analogous to a firmground (Glossifungites ichnofacies) and become compacted, consolidated anddewatered, prior to attests the early compaction and perhaps coalification of the exposure to recent tidal watersin sea cliffs or channels, peat. Teredolites borings also shows the presence of woody wherethey form a major substratefor boring bivalves (xylic) substrates, albeit often asdriftwood preserved in (Petricolu). This Holocene occurrence confirms the se- channel infills. quence of events necessary to form the Cretaceous Although dinosaur footprints moulds have been known Teredolites ichnocoenosis. Coastal swamp growth andpeat fromthe roofs of coal mine tunnels in the Mesaverde accumulation precedes burial, dewatering and compaction, Formation(Campanian, Upper Cretaceous) of Colorado before exposure following erosion by marine currents allows

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colonization by marinewood-borers. Preservation of this similar environmental situation in Cretaceous coals of Utah. ichnocoenosis was only achieved following the death of the Eagar et al. (1985) also described a channel floor infauna, infilling of the borings andrapid burial. Such a ichnocoenosis of Rhizocorallium jenense in a mud substrate unique ichnocoenosis gives us not only a means of from a Westphaliansequence in Cheshire. Todate, recognizing penecontemporaneousexposure of peat and however, no ichnocoenosis of borings in contemporaneous coal to marineconditions but also an indicator of the peat or coal has been recorded from Palaeozoic coal-bearing degree of 'coalification' of thesubstrate. However, it is sediments. questionable whether either the Cretaceous or the Holocene ichnocoenoses shouldbe the basis for a new woodground Teredolites ichnofacies. They may otherwise be regarded as Conclusions firmground boring communities that should be assigned to This review demonstratesthat asin sedimentological and the Glossifungites ichnofacies (Pemberton & Frey 1985). palaeoenvironmental analysis of other environments, the TheAlberta Cretaceouschannel floor ichnocoenosis of advances in recognition and interpretation of trace fossils in Teredolites is closely comparable to a Thalassinoides the past decade have added new dimensions to our suevicus ichnocoenosis recorded by Kamola (1984) from a understanding of coal-bearing strata, theirfaunas and

Asteriacites RB l I Chondrites FB I I Asterosoma F8 l Teichichnus FB I I RhizocoralliumFBlD I D8 I Lingulichnus I Skolithos I l Ophiomorpha D0 I Thalassinoides FBID I Diplocraterion DB I +l 1 1.9 I I l I I I I I DB P Arenicolites b9I 1 b 31 l Polykladichnus DB l I I l 7 31 I OphthalmidiurnFB I I I c, 3.5 I Phycodes F8 I 12.7; I PIanoliks FB 7 I 91 Beaconites FB I I l I Scolicia ' FT I I l=-1 I I I Helminthopsis FT 7 -1 I I I I I Cochlichnus FT Invertebratetrails 7 PelecypodichnusRB/E IsoDodichnus RB Kouphichnium LT -7 I I (Selenichnus) (RB) I I I I I I Paleohelcura LT I I I !.5.f Acripes LT lI I I l I Diplichnites LT I l I I l I Vertebrate tracks I I L Key to strata reviewed: FB-Feeding burrow l-Perrnian. Vryheid Frn. S. Africa 7-Pennsylvanian. Tennessee DB-Dwelling burrow 2-Stephanian, France 8-Pennsylvanian,Indiana EB-Escape burrow 3-Westphalian. Britain 9-Pennsylvanian,Kansas FT-Feeding trail 4-Stephanian. Britain 10-Middle Jurassic, Yorkshire coast LT-Locomotiontrace 5-Westphalian. Europe 1 l-Middle Jurassic. North Sea Basin RB-Resting burrow 6-Westphalian, Eastern Canada 12-Upper Cretaceous. Utah 8 Alberta

'ig. 5. Environmental distributionof trace fossils in Carboniferous to Cretaceous coal-bearing deltaic :diments as recorded by various authors. (Authors cited:(1)-(9) see Fig. 3, (10)-(12) see references ited in text).

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environments of formation.Clear relationships exist Carboniferousstrata, with remarks on the Microconchus carbonarius. between ichnofaunas and sedimentary facies in coal-bearing Memoirs of the Literary and Philosophical Society of Manchester, second strata ranging in age from Carboniferous to Cretaceous and series, 10, 181-201. BRIGGS,D. E. G., PLINT, A.G. & PICKERILL, R. K.1984. Arthropleura trails probably Tertiary.The comparison of the environmental from the Westphalian of eastern Canada. Palaeontology, 27, 843-55. distribution of ichnogenera shown in Fig. 5 suggests that at BROADHURST,F. M,, SIMPSON,I. M. & HARDY,P. G. 1980. Seasonal the presenttime it is prematureto define any distinct sedimentation in theUpper Carboniferous of England. Journal of ichnofacies or subichnofacies for coal-bearingsequences. Geology, 88, 639-55. BROMLEY, P.G., PEMEIERTON,S. G. & RAHMANI,R. A. 1984. A Cretaceous However, some recurring associations are recognizable, for woodground:the Teredolites ichnofacies. Journal of Paleontology, 58, example, Diplocraterion polyupsilon ininterdistributary 488-98. bays (Carboniferous,Permian, Jurassic), Pelecypodichnus BROWN, S. 1985. Jurassic. In: GLENNIE,K. W.(ed.) Introduction to the with escapeshafts in distributary mouth bars or crevasse Petroleum Geology of the North Sea. BlackwellScientific Publications, splays (Silesian, Jurassic), in lagoonal Oxford, pp. 133-60. Plunolites montanus BUDDING,M. & INGLIN, H.1981. A reservoir geological model of the Brent or floodbasin lake sediments (Silesian, Jurassic, Cretaceous) Sands in Southern Cormorant. In: ILLING,L. V. & HOBSON,G. D. (eds) andperhaps firmground borers Teredolites in compacted Petroleum Geology of 'the Continental Shelf of North-WestEurope. peator coalexposed tomarine conditions (Cretaceous, Heyden, London, pp. 326-34. Recent). Studies of the diversity and facies distribution of CALVER, M.A. 1968a. Coal Measures invertebrate faunas. In: MURCHISON, D. G. & WESTOLL,T. S. (eds) Coal and Coal bearing Strata. Oliver and both Silesian arthropod trackways and Cretaceousverte- Boyd, London, pp. 147-77. brate tracks in freshwater and terrestrial sediments correlate - 19686. Distribution of Westphalian marine faunas in northern England closely with interpretations derivedfrom body fossil and adjoining areas. Proceedings of the Yorkshire Geological Sociery, 37, evidence. 1-72. CARROLL,R. L., BELT,E. S., DINELEY, D. L.,BAIRD, D. & MCGREGOR,D. There is a need for more detailed sediment/trace fossil C. 1972. Excursion A 59. Vertebrate paleontology of eastern Canada. analysis in non-marine parts of coal-bearingsequences, Guidebook. 24th International Geological Congress, Montreal, 1972. particularly of Mesozoic and Cenozoic age to establish such CHAMBERLAIN, C.1978. K. Recognition of trace fossils in cores. In: BASAN,P. detailed associations as have been shown to exist in Silesian B. (ed.) Trace Fossil Conceptr. Society of Economic Paleontologists and Mineralogists, Short Course No. 5, pp. 119-66. sequences.Studies of trace fossils in cores of Jurassic CHISHOLM,J. I. 1983. Xiphosuridtraces, Kouphichnium aff. variabilis coal-bearing stratafrom offshore North Seaindicate that (Linck),from the Namurian Upper Haslingden Flags of Whitworth, there is considerablepotential for theroutine analysis of Lancashire. Report Institute of Geological Sciences, London, 83/10, trace fossils in cores, particularly in exploration or 37-44. -1985. Xiphosurid burrows from the Lower Coal Measures (Westphalian environmental evaluation of sedimentary basins. A) of West Yorkshire. Palaeontology, 28, 619-28. The relatively high preservation potential of burrows and COPELAND,M. J. 1957. Arthropod fauna of the Upper Carboniferous rocks of trackwaysin clastic coal-bearingsediments indicates their the Maritime Provinces. Geological Survey of Canada Memoir, 286. frequentadvantage over body-fossils interpretingin COUREL,L. 1983. Place du charbon dans le bassin deffondrement Stephanien coal-forming environments and their faunas. In situ burrows de Blanzy-Montceau (Massif Centralfrancais). Memoires Gtologique llniversite' de Dijon, 8, 71-82. often record the response of recognizable producers, many -1988. Intra-montaine Stephanian and Permian coal basins of the French of them soft-bodied or weakly skeletonized, to high stress Massif Central: new sedimentological data. In: BESLY, B.M. & KELUNG, factors in deltaicenvironments. Trackways of active G.(eds) Sedimentation in a Synorogenic Basin Complex: The Upper epifaunal arthropodsor tetrapods reveal not only the Carboniferous of North-West Europe. Blackie, Glasgow. CURRAN,H. A. (ed.) 1985. Biogenic structures: their usein interpreting presence of little-knownproducers in ephemeral aquatic depositional environments. Society of EconomicPaleontologists and marginal or terrestrialsituations, but frequentlyrecord Mineralogists, Special Publication No. 35. fascinating behaviour patterns or organism interrelation- DAWSON,J. W. 1868. Acadian Geology. Macmillan, London. ships, which are otherwiseunknown. In all aspects the - 1873. Impressionsand footprints of aquaticanimals and imitative markings on Carboniferous rocks. American Journal ofScience (Series information obtainablefrom trace fossils in coal-bearing 3), 16-24. sediments should not be ignored. DELAIR,3. B. & SARIEANT,W. A. S. 1985. History and bibliography of the study of fossil vertebratefootprints in theBritish Isles: Supplement 1973-1983. Palaeogeography, Palaeoclimatology, Palaeoecology, 49, References 123-60. DENNISON, C.& FOWLER,F. M. 1980. Palynological identificationof facies in ALLAN,J. A. & SANDERSON,J. 0. G. 1945. Geology of Red Deer and a deltaic environment. In: The Sedimentation of the North Sea reservoir Rosebud Sheets, Alberta. Research Council of Alberta Report No. 13. rocks. Norwegian Petroleum Society, Geilo. ANDERSON,A. M. 1975.Limulid trackways in LatePalaeozoic Ecca EAGAR, R.M. C., BAMES, J. G.,COLLINSON, J. D., HARDY,P. G., OKOLO, sedimentsand their palaeoenvironmental significance. South African S. A. & POLLARD,J. E. 1985. Trace fossil assemblagesand their Journal of Science, 71, 249-51. occurrence in Silesian(mid-Carboniferous) deltaic sediments of the -1976. Fish trails from the Early Permian of South Africa. Palaeontology, CentralPennine Basin, England. In: CURRAN,H. A. (ed.) Biogenic 19, 397-409. structures: their use in interpreting depositional environments. Society of - 1981. The Umfolozia arthropod trackways in the Permian Dwyka and Economic Paleontologists and Mineralogists, Special Publication No. 35, Ecca Series of South Africa. Journal of Paleontology, 55, 84-108. pp. 99-149. ARCHER,A. W. & MAPLES,C. G. 1984. Trace-fossildistribution across a EKDALE,A. A.,BROMLEY, R. G. & PEMEIERTON,S. G. 1984. Ichnology: the marine-to-nonmarinegradient in thePennsylvanian of southwestern use of trace fossils in sedimentology and stratigraphy. Society of Indiana. Journal of Paleontology, 58, 448-66. Economic Paleontologists and Mineralogists, Short Course No. 15. BANDEL,K. 1967a. Trace fossils from two Upper Pennsylvanian sandstones in ELLIOT, R.E. 1985. An interpretation of the trace fossil Cochlichnus kochi Kansas. Paleontological Contributions, University of Kansas. Paper 18. (Ludwig) from the East Pennine Coalfield of Britain. Proceedings of the - 1967b.Isopod and limulid marks and trails in TonganoxieSandstone Yorkshire Geological Society, 45, 183-8. (UpperPennsylvanian) of Kansas. Paleontological Contributions, FARROW,G. E. 1966. Bathymetric zonation of Jurassic trace fossils from the University of Kansas, Paper 19. coast of Yorkshire,England. Palaeogeography, Palaeoclimatology, BESLY,B. M. 1988. Palaeogeographicimplications of lateWestphalian to Palacoecology, 2, 103-51. early Permian red beds, Central England. In: BESLY,B. M. & KELLING, FERGUSON,L. 1966. Therecovery of somelarge track-bearing slabs from G. (eds) Sedimentation in a Synorogenic Basin Complex: The Upper Joggins, Nova Scotia. Maritime Sedimentr, 2, 128-30. Carboniferous of North-West Europe. Blackie, Glasgow & London, pp. - 1975. TheJoggins Section. Ibid 11, 69-76 (= 1976). In: Ancient 200-21. Sedimentr of Nova Scotia. Field Trip Guidebook Eastern Section Society BINNEY,E. W. 1852. One sometrails and holes found in rocks of the of Economic Paleontologists and Mineralogists, 111-18.

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Received 22 September 1986; revised typescript accepted 28 September 1987.

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