Anda di halaman 1dari 7



Early Permian Xiphosurid Trackways from India


Department of Geology, Jogamaya Devi College, 96 S. P. Mukherjee Road, Kolkata - 700 026 2 Department of Geology and Centre of Advance Studies in Precambrian Geology, Presidency University, 86/1 College Street, Kolkata - 700 073 Email:

Abstract: Present work is the first report of Early Permian xiphosurid trackways from India. Surface trackways and undertracks, preserved within ice-marginal storm-affected, shoreface sediments of the Talchir Formation, Jayanti coalbasin closely resemble Kouphichnium Nopsca 1923. Trackways reveal distinct heteropody with foliate pusher impressions, paired series of semi-circular, oval, V- or Y-shaped chilate (walking) leg impressions. Varied ventral impressions also include telson drag marks and telson dents; spine drag marks, book -gills impression, genal gouges as well as symmetric, paired lateral genal furrows, partial cubichnial impressions of prosoma and spinose opisthosoma. A tentative early Permian age (Asselian/Sakmarian) and marginal marine facies association points towards a paleolimuloidae affinity for the trace makers, which is in conformity with their inferred widths of exoskeleton, sub-rounded prosoma and subtriangular spinose opisthosoma with predictable movable spines. Flexed pusher impressions and co-dominant pincer impressions as well as, other ventral impressions in abundance, confirm crawling as their dominant mode of locomotion. Inferred presence of juvenile crawlers in the community suggests a probable shoreline or marginal marine ecospace. Keywords: Trackways, Xiphosura, Kouphichnium, Permian, Marginal Marine.


Heteropod trackways and their trace makers have attracted many workers for their palaeoecological and evolutionary implications (Caster, 1938, 1944; Goldring and Seilacher, 1971; Briggs and Rolfe, 1983; Chisholm, 1983; Siveter and Selden, 1987; Selden and Siveter, 1987; Anderson, 1997, 1998; Anderson and Selden, 1997; Braddy and Dunlop, 1997; Racheboeuf et al. 2002; Romano and Whyte, 2003; Pirrie et al. 2004; Weber and Braddy, 2004; Mickelson et al. 2006; Rudkin et al. 2008). Initially, these traces were interpreted variously as the activity of pterodactyls, bipedal vertebrates or amphibia. The ichnogenus, Kouphichnium Nopsca 1923 was assigned to these various traces. Much later, Caster (1938) convincingly identified these trace fossils as arthropod trackways of limulid origin. Till date, there is no report of xiphosura from geologic past in India beyond the tethyan segment. Inferred eurypterid (Palmichnium) trackways (Draganits et al. 1998; Draganits et al. 2001) and trace fossils of probable xiphosurid (chelicerata) origin (Bhargava and Bassi, 1998) from the Lower Devonian Muth Quartzite of Tethyan Spiti Basin are the only eurypterid activities, reported from India. The present work is the first report on early Permian xiphosurid

traces from India. Incidentally, these are the oldest record of xiphosurid activity from the peninsular part of India.

Trackways reported in the present work, occur within the Sandstone-Mudstone Alternation Facies (Facies E, Table 1), representing the glaciomarine shoreface sediments within the Talchir depositional sequence that are exposed along the banks of NNE-SSW flowing Pathorjor tributary of Jayanti River. Geographic locality of trace fossil occurrences (2410'8.82"N; 8641'45.58"E) and their stratigraphic position in the depositional sequence is shown in Fig. 1. The Talchir facies types in this basin lie along a fault contact with the Precambrian granitoids, and are unconformably overlain by trough cross-stratified sandstones of the Karharbari Formation (Table 1, Fig. 1). Similar facies association of the Talchir Formation has been interpreted as a marginal marine storm deposit in the adjoining parts of Deoghar Basins and Damodar Valley Basins (Casshyap and Srivastav, 1987; Bose et al. 1992; Mukhopadhyay and Bhattacharya, 1994; Acharyya, 1997; Bhattacharya et al. 2002; Bhattacharya, 2003; Bhattacharya et al. 2004; Bhattacharya et al. 2005; Chakraborty and Bhattacharya,

0016-7622/2012-80-1-129/$ 1.00 GEOL. SOC. INDIA



Fig.1. Plate showing: (a) Geographic location of the Jayanti Basin in the map of India; (b) Geological map (modified after Raja Rao, 1987) of the Jayanti Basin, Jharkhand marked with fossil location and (c) Distribution of grain size, lithology, depositional fabric and bioturbation at distinct stratigraphic levels.

EARLY PERMIAN XIPHOSURID TRACKWAYS Table1. Brief description and interpretation of the facies types of the Talchir Formation in Jayanti Basin Formation Karharbari Facies Trough Cross-stratified Sandstone Wave-Rippled Sandstone (Facies F) Sandstone- Mudstone Alternation Facies (Facies E) Talchir Thinly Bedded Sandstone- Mudstone Interbedded Facies (Facies D) Dropstone- bearing Mudstone Facies (Facies C) Interpretation Braided Channel deposits. Wave reworked, rip current submerged channel fill sediments. Shallow, upper shoreface sediments that recorded alternating current and slack phases. Shoreface-Lower foreshore environment extending up to offshore. Above wave-base storm deposits. In the context of associated facies, this mudstone facies is interpreted to be basin marginal mud (Syvitski et al. 1985) deposited in a marginal-marine association with floating icebergs. Disparate, patchy distribution of this facies, overlying Facies B indicates occasional iceberg dumping. Storm-laid near shore sedimentation above wave-base.


Bioclast bearing Matrix- Supported Conglomerate Facies (Facies B) Massive Sandstone Facies (Facies A) Precambrian Granitoids

2005). Host sediments (Facies E) containing the trackways overlie a bioclast bearing, matrix-supported conglomerate facies (Facies B, Table 1). Similar fragmentary bivalve shells in the marine biota have been assigned a tentative basal Permian or Permo-Carboniferous age elsewhere in the peninsular gondwanan basins (Reed, 1928; Acharyya and Shah, 1975; Shah and Sastri, 1975; Waterhouse and Ranga Rao, 1989; Archbold et al. 1996; Sengupta et al. 1999; Ghosh, 2003 and references therein). In this context, xiphosurid traces preserved in still higher stratigraphic interval (i.e., in Facies E) may be assigned an early Permian (Asselian/Sakmarian) age.

consisting of two kinds of imprints: (1) two chevron like series each of four oval or round holes or bifid V-shaped impressions or scratches, forwardly directed (made by anterior four pairs of feet); and (2) one pair of digitate or flabellar, toe-shaped or otherwise variable imprints (made by bird-foot like pushers of fifth pair of feet, with their four or five leaf-like movable blades); track with or without median drag mark. Kouphichnium isp. (Figs. 2-5) Materials. K/00-1, K/00-2 and additional field observations in Facies E Description. Heteropod, prevalently circuitous trackways; consisting of continuous and discrete surface features, preserved as paired and juxtaposed, low relief, epichnial troughs and hypichnial ridges; commonly with three (Figs. 2, 3), or rarely four (Fig. 4) flabellar lobes, are recognized as ectognath (pusher) imprints (Pu). Some conical mounds (Pbm) present along the length of trackway are truncated at one end (Fig. 4). Associated tiny, sub-circular or elliptical imprints and V or Y-shaped scratch marks (Figs. 2-4) are recognized as imprints of anterior walking legs or pincers (Pi). Paired series of pincer impressions delineate a V-shape (Fig. 4, between stippled lines). Maximum digit lengths of individual ectognath impressions vary between 6.8mm and 14.95mm. Varied ventral impressions (Figs. 2, 3) include arrays of tiny, concave out indentations recognized as genal gouges (Gg), and continuous paired genal drag marks that define lateral trackway margins (Lm). Pusher impressions lie between the trackway margins except in sharply bent portions of the trackways, where they lie beyond it. In some instances, these imprints are modified with a

Kouphichnium trackways were present in the field both as epichnial hollows (Specimen No. K/00-1, in situ block) and hypichnial ridges or undertracks (Specimen No. K/ 00-2). Computer aided measurements were made on scanned images of collected specimens, field photographs and published photographs to document and compare features of xiphosuran trackway attributes of same and varied geological age. Kouphichnium trackways were described following the terminology of Hntzschel, 1975. The specimens studied are preserved in the Department of Geology, Presidency College, Kolkata, India.

Ichnogenus Kouphichnium Nopsca 1923 Diagnosis. (following Hntzschel 1975, p. W75). Hetropodous tracks of great variability; complex track



parallelism between individual opisthosomal spine impressions and attendant feeble sinuosity in the spine drag marks is also discernible (Fig 3, arrow marked). Spine drag marks (Spd) are elsewhere present along the trackways as thin but sharp, discontinuous and non-parallel scratches. A less conspicuous falcate impression (34mm between extremities) probably represents the anterior marginal impression of the prosoma (Pm). Average track widths in the measured trackways (trackways 1, 2 and 3) are respectively 11.60 mm, 28.99 mm and 24 mm. Discussion. A definite xiphosurid origin may be suggested for the trace Fig. 2. Field photograph of Kouphichnium trackways on the bedding surface (Facies E). Note the circuitous pattern of the trackways and the various ventral impressions makers of Jayanti Basin, from the heteropodous trackways with well(Specimen No. K/00-1; coin diameter 2.4 cm). differentiated pusher (ectognath) and pincer (waking leg) impressions, movable telson impressions and forward pointing V-shaped arrangements of paired pincer impressions (cf. Hntzschel, 1975). Codominant pincer and pusher impressions in the trackways; genal and telson drag marks, gill-book impressions as well as partial cubichnial traces of prosoma and opisthosoma present along the trackways, indicate nearly continuous body contacts of the trace makers with the substrate. A probable crawling mode of locomotion for the trace makers is inferred thereof. Arrays of discontinuous genal gouges and Fig.3. Interpretive line drawing of the discernible trackway features (from Fig. 3) showing telson dents perhaps indicate occasional impressions of lateral margin (Lm), prosoma anterior (Pm), book-gills (Gb), halts amidst a clinging up-slope gait. buckler spine (Bs) and spine drag marks (Spd), genal gouges (Gg), varied telson Outstretched, dragged Y-shaped impresmarks (Tm) and telson dents (Td) as well as, pincer (Pi) and pusher impressions sions in this context indicate inward (Pu). Also note the inferred direction of movement of the trace makers along the dragging of pincers on soft sediment in trackways (long arrow). an effort to stabilize and anchor (cf. dragged, incurved extension. Discontinuous, truncate ridges Romano and Whyte, 2003). V-shaped alignment of paired and furrows between and beyond the lateral margins are pincer impressions, truncated mud mounds along the recognized as aspects of movable telson impressions (Tm) trackway and complimentary orientations of the ectognath that run parallel or oblique to trackway axis. These are often blades help to identify the polarity of movement (Fig. 4, associated with high angle telson dents (Td). Series of long arrow) of the trace makers (cf. Caster, 1938; Osgood, transversely parallel, sinuous, low relief discrete ridges 1970). Preponderant pusher and pincer impressions were between lateral margins along the Trackway 3 (inset, Figs. recorded, respectively in the Carboniferous and Triassic 2, 3) probably represent book-gills (Gb) impression. Some trackways (Goldring and Seilacher, 1971; Seilacher, 1978). arcuate impressions with posterior spinose projections Jayanti basin trackways, however, do not show any distinct probably represent a partial cubichnia preserving spinose bias towards either of these. The present authors prefer to opisthosoma with buckler spines (Bs). Subtle nonrelate co-dominant pusher and pincer impressions in the



Caster, 1938). Trackways, varying in average track widths probably suggest trace-making instars of varying ontogenic phases within the community. (cf. Chisholm, 1983; Tyler, 1988). Status of circuitous crawling traces of xiphosura as an environment indicator is intriguing. Goldring and Seilacher (1971) reported walking traces from sediments deposited at moderate depths while other workers insisted on their subaerial or shallow-water origin; or their formation in the absence of traveling waves (Caster, 1938; Rudloe and Herrnkind, 1980; Mickelson et al., 2006). Shoreline attachments of limuloidae instars have been documented by many workers (Botton et al., 2003; Braddy 2004 and references therein). Permian paleolimuloids are so far described as marine forms (cf. Babcock et al., 2000). A definite marine link is therefore suggested for these trace fossils, based on their sedimentary facies association and ethology.
Fig.4. Hand specimen photograph of Kouphichnium trackway features showing crawling gait of xiphosurid trace maker. Note the V-shaped arrangement (between stippled lines) of pincer impressions (Pi) and comparable abundances of oriented pusher impressions (Pu); and truncated push -back mud mounds (Pbm) defining the direction (long arrow) of movement of the trace-maker (Specimen no. K/00-2). CONCLUSIONS

Jayanti Basin trackways to their crawling mode of locomotion (cf. Chisholm, 1983; Romano and Whyte, 2003). A tentative link of these early Permian trackways with Paleolimulidae trace-makers seems probable (cf. Anderson and Selden, 1997) from their inferred movable opisthosomal spines (cf. Selden and Siveter, 1987), narrow opisthosoma and rounded prosoma (cf. Ambrose and Romano, 1972; Anderson and Selden, 1997; Babcock et al., 2000; Hagadorn, 2002; Allen & Feldmann, 2005; Buta et al., 2005; Rudkin et al., 2008 and references therein). Widths of exoskeletons (11.60mm-34.63mm) of the trace-makers, inferred from the track widths, measured between pusher impressions and genal drag marks, overlap with those (22mm-135mm) of reported Paleolimulids (cf. Selden and Siveter, 1987; Babcock et al., 2000; Hagadorn, 2002; Allen and Feldmann, 2005; Lucas and Lerner, 2005; Moore et al., 2007; Rudkin et al., 2008) probably indicating the presence of at least few juvenile trace makers in the community, a fact, further supported from their circuitous trackways (cf.

Kouphichnium trackways in Talchir sediments of Jayanti Basin record Early Permian xiphosurid activities from Indian subcontinent. A paleolimuloid affinity seems probable from their age constraints, inferred subtriangular opisthosoma with movable spines and broadly rounded prosoma. Circuitous trackways of distinctly varying average track widths suggest that some of the trace-makers in the community were juvenile instars. Varied ventral impressions including incomplete cubichnia, presence of co-dominant pusher and pincer impressions along the trackways indicate aspects of surface crawling of the trace-makers. A definite marine link is emphasized for these trackways based on their ethology and association with shoreface sediments. Acknowledgements: The authors are thankful to the Department of Geology, Presidency University, Kolkata for providing infrastructure facilities during the work. Authors are grateful to L.I. Anderson and J.E. Pollard for their helpful suggestions. Biplab Bhattacharya, Hooghly Mahosin College is acknowledged for his help during the work. We thank University Grants Commission, Government of India, for partial financial support (Grants No. F.5-14/2001 SR-1; F.PSW-046/05-06-ERO).

References ACHARYYA, S.K. (1997) Evolutionary characters of the Gondwanic crust. Indian Minerals, v. 51(1-2), pp.124-127. ACHARYYA, S.K. and SHAH, S.C. (1975) Biostratigraphy of marine fauna associated with the diamictites of the Himalayas. Bull.

Indian Geol. Assoc., v.8(20), pp. 9-23. ALLEN, J.G. and FELDMANN, R.M. (2005) Panduralimulus babcocki n. gen. and sp., A new Limulacean Horseshoe Crab from the Permian of Texas. Jour. Paleont., v.79(3), pp.594-600.



AMBROS, T. and ROMANO, M. (1972) New Upper Carboniferous Chelicerata (Arthropoda) from Somerset, England. Palaeontology, v.15(4), pp.569-578. ANDERSON, L.I. (1997) The xiphosuran Liomesaspis from the Montceau-les-Mines Konservat- Lagerstatte, Massif Central, France. Neues Jahrbuch fr Geologie und Palaontologie, Abhandlungen, v.204, pp.415-436. A NDERSON , L.I. (1998) A new specimen of the Silurian synziphosurine Cyamocephalus. Proc. Geologists Assoc., v.110, pp.211-216. ANDERSON, L.I. and SELDEN, P. A. (1997) Opisthosomal fusion and phylogeny of Palaeozoic Xiphosura. Lethaia, v.30, pp.19-31. ARCHBOLD, N.W., SHAH S.C. and DICKINS, J.M. (1996) Early Permian brachiopod faunas from Peninsular India: Their Gondwanan relationships. Historical Biology, v.11(1), pp.125135. B ABCOCK , L.E., M ERRIAM , D.F. and W EST , R.R. (2000) Paleolimulus, an early limuline (Xiphosurida), from PennsylvanianPermian Lagerstatten of Kansas and taphonomic comparison with modern Limulus. Lethaia, v.33, pp.129-141. BHATTACHARYA, B. (2003) Storm event beds in Talchir Formation, Jayanti Basin, Jharkhand, India. Indian Jour. Earth Sci., v.30(14), pp.37-41. BHATTACHARYA, H.N., BHATTACHARYA, B., CHAKRABORTY, I. and CHAKRABORTY, A. (2004) Sole Marks in Storm Event Beds in the Permo-Carboniferous Talchir Formation, Ranigunj Basin, India. Sedimentary Geol., v.166, pp.209-222. BHATTACHARYA, H.N., CHAKRABORTY, A. AND BHATTACHARYA, B. (2005) Significance of Transition between Talchir Formation and Karharbari in Lower Gondwana Basin Evolution- A Study in West Bokaro Coal Basin, Jharkhand, India. Jour. Earth System Sci., v.114(3), pp.275-286. BHATTACHARYA, H. N., GOSWAMI, A. and CHAKRABORTY, A. (2002) Sedimentary facies analysis of a Permo-Carboniferous terminoglacial succession, Saharjuri Basin, Jharkhand, India. Jour. Geol. Soc. India, v.60, pp.401-410. BHARGAVA, O.N. and BASSI, U.K. (1988) Trace fossils from the Palaeozoic-Mesozoic sequence of Spiti-Kinnaur (Himachal Himalaya) with comments on palaeoenvironmental control on their frequency. Jour. Geol. Soc. India, v.32(3), pp.227238. BOSE, P.K., MUKHOPADHYAY, G. and BHATTACHARAYA, H.N. (1992) Glaciogenic coarse clastics in a Permo-Carboniferous bedrock trough in India: A sedimentary model. Sedimentary Geol., v.76, pp.79-97. BOTTON, M.L., LOVELAND, R.E. and TIWARI, A. (2003) Distribution, abundance, and survivorship of young-of-the-year in a commercially exploited population of horseshoe crabs Limulus polyphemus. Marine Ecology Progress Series, v.265, pp.175184. BRADDY, S.J. (2004) Ichnological Evidence for the Arthropod Invasion of Land. Fossils and Strata, v.51, pp.136-140. BRADDY, S.J. and DUNLOP, J. (1997) The functional morphology of mating in the Silurian eurypterid Baltoeurypterus

tetragonophthalmus. Zoological Jour. Linnaean Society, v.121, pp.435-461. BRIGGS, D.E.G. and ROLFE, W.D.I. (1983) A giant arthropod trackway from the Lower Mississippian of Pennsylvania. Jour. Paleont., v.57, pp.377-390. BUTA, R.J., KOPASAKA-MARKEL, D.C., RINDSBERG, A.K. and MARTIN, A.J. (2005) Atlas of Union Chapel Mine Invertebrate Trackways and other Traces. In: Buta, R. J., Rindsberg, A. K. and Kopaska-Merkel, D. C. (eds.) Pennsylvanian Footprints in the Black Warrior Basin of Alabama. Alabama Paleontological Society Monograph no. 1, pp.277-337. CASSHYAP, S.M. and SRIVASTAV, V.K. (1987) Glacial sedimentation of Late Paleozoic Talchir diamictite, Pench Valley coalfields, Central India. Geol. Soc. Amer. Bull., v.85, pp.749-760. CASTER, K.E. (1938) A restudy of the tracks of Paramphibius. Jour. Paleontology, v.12, pp.3-60. CASTER, K.E. (1944) Limulid trails from the Upper Triassic (Chinle) of the Petrified Forest National Monument, Arizona. Amer. Jour. Sci., v.242, pp.74-84. CHAKRABORTY, A. and BHATTACHARYA, H.N. (2005) Ichnology of a Permo-Carboniferous glaciomarine ice-contact deltaic set-up, Talchir Formation, Saharjuri Basin, India. Ichnos, v.12(1), pp.31-45. CHISHOLM, J.I. (1983) Xiphosurid traces, Kouphichnium aff. Variabilis (Linck), from the Namurian Upper Haslingden Flags of Whiteworth, Lancashire, Rep. Inst. Geol. Sci., No.83(10), pp.37-44. DRAGANITS, E., GRASEMANN, B. and BRADDY, S.J. (1998) Discovery of abundant arthropod trackways in the ?Lower Devonian Muth Quartzite (Spiti, India): implications for the depositional environment. Journal of Asian Earth Sciences, v.16(2-3), pp.109-118 DRAGANITS , E., B RADDY , S.J. and B RIGGS , D.E.G. (2001) A Gondwanan Coastal Arthropod ichnofauna from the Muth Formation (Lower Devonian, Northern India): Paleoenvironment and Tracemaker Behavior. Palaios, v.16(2), pp.126-147. GHOSH, S.K. (2003) First record of marine bivalves from the Talchir Formation of the Satpura Gondwana Basin, India: Palaeogeographic implications. Gondwana Res., v.6, pp. 312320. GOLDRING, R. and SEILACHER, A. (1971) Limulid undertracks and their sedimentological implication. Neues Jahrbuch fr Geologie und Palontologie Abhandlungen, v.137, pp.422442. HAGADORN , J.W. (2002) Bear Gulch: an exceptional Upper Carboniferous PlattenKalk, 167-183. In: D. J. Bottjer and R. K. Bambach (Eds.). Exceptional fossil preservation: a unique view on the evolution of marine life, Columbia University press, New York, NY, 424p. HNTZSCHEL, W. (1975) Trace fossils and problematica. In: C. Teichert (Ed.), Treatise on Invertebrate Paleontology, Part W. Miscellanea, Supplement 1. Geol. Soc. Amer. and Univ. of Kansas Press, W269p. LUCAS , S.G. and LERNER , A.J. (2005) Lower Pennsylvanian



Invertebrate Ichnofossils from the Union Chapel Mine, Alabama: A Preliminary Assessment In: R.J. Buta et al., (Eds.), Pennsylvanian Footprints in the Black Warrior Basin of Alabama. Alabama Paleontological Society Monograph no. 1, pp. 147-152 MICKELSON, D.L., HUNTOON, J.E. and KVALE, E.P. (2006) The Diversity and Stratigraphic Distribution of Pre-Dinosaurian Communities from the Triassic Moenkopi Formation In: S.G. Lucas, J.A. Spielmann, P.M. Hester, J.P. Kenworthy and V.L. Santucci (Eds.), Fossils from Federal Lands. New Mexico Museum of Natural History and Science Bull., v.34, pp.132137. MOORE, R.A., MCKENZIE, S.C. and LIBERMAN, B.S. (2007) A Carboniferous Synziphosurine (Xiphosura) from the Bear Gulch Limestone, Montana, U.S.A. Palaeontology, v.50(4), pp.1013-1019. MUKHOPADHYAY, G. and BHATTACHARYA, H. N. (1994) Facies analysis of Talchir Sediments (Permo-Carboniferous), Dudhi Nala, Bihar, India- A glaciomarine model. Ninth Internat. Gondwana Symp., 1, pp.737-753. OSGOOD, R. G., (1970) Trace fossils of the Cincinnati area. Palaeontographica Americana, v.6 , pp. 281-444. PIRRIE, D., FELDMANN, R.M. and BUATOIS, L.A. (2004) A new decapod trackway from the Upper Cretaceous, James Ross Island, Antarctica. Palaeontology, v.47(1), pp.1-12. RACHEBOEUF, P.R., VANNIER, J. and ANDERSON, L.I. (2002) A new Three-Dimensionally Preserved Xiphosuran Chelicerate from the Montceau-Les-Mines, Largersttte (Carboniferous, France). Palaeontology, v.45(1), pp.125-147. RAJA RAO, C. S. (1987) Coal Fields of India. Bull. Geol. Surv. India, Series A, v.45, IV(1) , pp.336. REED, F.R.C. (1928) A Permo-Carboniferous marine fauna from Umaria Coal field. Rec. Geol. Surv. India, v.60, pp.367-398. Romano, M. and WHYTE, M. (2003) The first record of Xiphosurid

(Arthropod) trackways from the Saltwick Formation, Middle Jurassic of the Cleveland Basin, Yorkshire. Palaeontology, v.46(2), pp.257-269. RUDKIN, D.M., YOUNG, G.A. and NOWLAN, G.S. (2008) The Oldest Horseshoe Crab: A New Xiphosurid from Late Ordovician Konservat-Lagerstatten Deposits, Manitoba, Canada Palaeontology, v.51(1), pp.19. R UDLOE , A.E. and H ERRNKIND , W.F. (1980) Orientation by horseshoe crabs, Limulus polyphemus, in a wave tank. Jour. Marine and Freshwater Behaviour and Physiology, v.7(3), pp.199-212. S EILACHER , A. (1978) Use of trace fossil assemblages for recognizing depositional environments. In: P.B. Basan (Ed.), Trace Fossil Concepts. SEPM Short Course 5, pp.167-181. SELDEN, P.A. and SIVETER, D.J. (1987) The origin of Limuloids. Lethaia, v.20, pp.383-392. SENGUPTA, S., CHAKRABORTY, A. and BHATTACHARYA, H.N. (1999) Fossil Polyplacophora (Mollusca) from Upper Talchir sediments of Dudhi Nala, Hazaribagh, Bihar. Jour. Geol. Soc. India, v.54, pp.523-527. SHAH, S.C. and SASTRY, M.V.A. (1975) Significance of early Permian marine fauna of Peninsular India, In: K.S.W. Campbell (Ed.), Gondwana Geology, pp. 391-396. TYLER, D.J. (1988) Evidence and significance of limulid instars from trackways in the Bude Formation (Westphalian), southwest England. Proc. Ussher Society, v.7, pp.77-80. WATERHOUSE , J.B. and RANGA RAO, A. (1989)Early Permian Brachiopod and Molluscan Species from the Bap Formation of Peninsular India. Palontologische Zeitschrift, v.63(1-2), pp.25-39. WEBER, B. and BRADDY, S.J. (2004) A Marginal Marine Ichnofauna from the Blaiklock Glacier Group (?Lower Ordovician) of the Shackleton Range, Antarctica. Trans. Royal Soc. Edinburgh: Earth Sci., v.94, pp.1-20.

(Received: 18 December 2010; Revised form accepted: 13 February 2012)