Available Online at http://www.recentscientific.com International Journal of CODEN: IJRSFP (USA) Recent Scientific

International Journal of Recent Scientific Research Research Vol. 11, Issue, 07 (B), pp. 39177-39191, July, 2020 ISSN: 0976-3031 DOI: 10.24327/IJRSR Research Article

SOFT–BODIED EDIACARAN METAZOANS AND ICHNOFOSSILS FROM THE MARWAR SUPERGROUP, WESTERN RAJASTHAN: EMERGENCE OF EARLIEST MULTICELLULAR, COMPLEX MEGASCOPIC ANIMAL LIFE

Purnima Srivastava

Centre of Advanced Study in Geology, Lucknow University, Lucknow, 226020

DOI: http://dx.doi.org/10.24327/ijrsr.2020.1107.5457

ARTICLE INFO ABSTRACT

Article History: Well preserved ediacaran fossils have earlier been discovered from the basal part of the Marwar Supergroup, western Rajasthan. The assemblage reported till date comprised of medusoids and Received 06th April, 2020 microbial mats only. Ediacaran assemblage with fronds viz. Charniodiscus and trace fossils are Received in revised form 14th reported for the first time from the Jodhpur Sandstone formation of the Marwar Supergroup. In May, 2020 Jodhpur assemblage, discs of variable sizes and morphological structures are associated with un- Accepted 23rd June, 2020 oriented, haphazardly distributed mesh of irregular bodies comparable to algae. These discs exhibit a Published online 28th July, 2020 wide size range from a few millimeters to 75 centimeters in diameter. Exceptionally large size of the

discs in present assemblage represents largest discs reported so far from any Ediacaran assemblage. Key Words: Although, larger medusoid discs have been reported from USA, but they are from the Middle Ediacaran, Marwar Supergroup, Metazoans, Cambrian and even younger rocks. Presence of microbial mats and weed-like structures with well Multicellular preserved hold fasts and horizontal rhizome like structures in association with some of these large- sized discs support their animal affinity, which probably feed on this weed- like vegetations. This association also support their benthic habitat. Unlike general trend of sudden increase in size of organisms in Ediacaran period and further decrease in size during Cambrian, these discs continued increasing in size in Cambrian also. Palaeoecological distribution of such well preserved morphologies are certainly an indication of ecosystem, where algal blooms enhabited the Ediacaran organisms represented by medusoid discs and possibly serve as their food. Moderately diverse Ediacaran body fossils, traces, burrows and giant sized algal fossils occur sporadically in Jodhpur Sandstone Formation, which are well exposed in open mines in and around Sur Sagar and Artiya Kalan area of Jodhpur district (see Srivastava, 2012-c and 2013, 2015). Here in this paper we deal only with Ediacaran metazoan body fossils (impressions) and ichnofossils.

Copyright © Purnima Srivastava, 2020, this is an open-access article distributed under the terms of the Creative Commons

Attribution License, which permits unrestricted use, distribution and reproduction in any medium, provided the original work is

properly cited.

INTRODUCTION related to modern jellyfish and corals, representing first megascopic animals known in fossil record. In recent years The Ediacaran period was a distinctive stage of biological these fossils have been compared with plants, giant single- evolution and Ediacaran fossils represent the earliest known celled organisms and even a failed evolutionary experiment macroscopic and complex life forms on the planet earth. They completely separated from all known kingdoms of life occur in various parts of world for which more than thirty (Seilacher, 1992). Most of the Ediacaran fossils are preserved species have been identified so far. Reconstruction of as casts and molds in relatively coarse siliciclastic sedimentary biological evolution in the fossil records, only reflect the wide rocks (Narbonne, 2005). Their unusual morphology resulted in expansion of metazoans with a mineralized skeleton. However various interpretations regarding their affinities like lichens, the timing of appearance and early radiation of extant high rank which covered much of the earth during Precambrian times taxa is still a matter of debate for many reasons. The possible (Retallack, 1994), fungus-like organisms (Peterson et al., absence of skeletons in majority of Vendian organisms may be 2003), prokaryotes (Steiner and Reitner, 2001), stem or crown- one of them (Serezhnikova, 2014). Ediacaran fossils are group animals (Glaessener, 1984; Conway Morris, 1993; considered the black holes of palaeontology so inscrutable, Fedonkin and Waggoner, 1997) or vendobionts that are which swallow up every theory thrown at them. Conventionally different from animals (Seilacher, 1992). Problem of large-size they have been interpreted as the impressions left by animals

*Corresponding author: Purnima Srivastava Centre of Advanced Study in Geology, Lucknow University, Lucknow,226020 International Journal of Recent Scientific Research Vol. 11, Issue, 07 (B), pp. 39177-39191, July, 2020 of ediacarans in Precambrian time is explained by Seilacher as Geological Setting bodies of Ediacaran organisms was comprised of fluid-filled The Marwar Supergroup was previously known as Trans compartments just like an air-mattress. Among quilted body Aravalli-Vindhyans where sedimentary rocks occurring on structures, a large surface area was maintained across which western side of NE-SW trending Aravalli Ranges in parts of nutrients and gasses could diffuse. These organisms must have Bikaner, Churu, Ganganagar, Jaisalmer, Jodhpur, Nagaur and absorbed nutrients from sea-water or harbored symbiotic Pali districts of Rajasthan. Khan (1971) designated these rocks microbes that provided nourishment through photo synthesis or as Marwar Superroup and gave the startigraphic succession. chemosynthesis. Absence of proper mouth and anal structures The supergroup is further divided into three groups, in in Ediacaran fossils also favor the plant affinity to some extent, stratigraphic order these are; Jodhpur, Bilara and Nagaur where photosynthetic ability could explain puzzling aspect. Groups (Pareek, 1984). The rocks of Jodhpur Group show The uncertainity of taxonomic position of these organisms has overlapping relationship with the Malani volcanics (Paliwal, led to different opinions on their place in the evolution. Despite 1998).which is dated as 779- 681 Ma (Rathore et al. 1999). The conflicting theories and reconstructions, some Ediacaran taxa Jodhpur Group is further divided into three formations; these are unanimously assigned to basal group of metazoa are Pokran Boulder Beds, Sonia Sandstone Formation and (Glaessner, 1984; Fedonkin 1985; Seilacher et al., 2003, Girbhakhar Formation. Chauhan et al., 2004, merged the Sonia Fedonkin et al., 2007; Xiao & Laflamme, 2009). Their views Sonia and Girbhakhar Sandstone Formations and named them are not in contrast with present day palaeontological and Jodhpur Sandstone Formation (Fig 2). Ediacaran fossils have molecular genetic reviews, according to which most been reported from the outcrops of Jodhpur Sandtone exposed Phanerozoic animal phyla formed in the late Precambrian in numerous quarries at and around Sur Sagar and Artiya Kalan (although very little evidence is provided by the fossil record). areas of Jodhpur district (Fig. 1). The rocks are least disturbed, It is also believed that in Cambrian, animals became diversified almost horizontal and exhibit well preserved sedimentary and many bilaterian taxa acquired skeletons (Erwin et al., structures like ripple marks of various scales and alignments, 2011). Very recently well preserved ediacaran fossils have been cross bedding, internal primary sedimentary structures, range discovered from the basal part of the Marwar Supergroup, from trough cross bedding, small-scale cross beddings etc. western Rajasthan (Raghav et al., 2005, Kumar et al., 2009, Lithology at Artiya Kalan, chocolate brown sandstone Kumar and Pandey, 2009, Srivastava, 2012 a,b,c). The interbedded with siltstone, shale and conglomeratic sandstone assemblage reported till date comprised of medusoids and and at Sur Sagar can be represented by coarse to fine grained microbial mats only. Ediacaran assemblage with fronds viz. brick red siliciclastic sandstone (Fig. 2-a,b respectively). Charniodiscus and trace fossils are reported for the first time from the Jodhpur Sandstone Formation of the Marwar Supergroup. In Jodhpur assemblage, discs of variable sizes and morphological structures are associated with un-oriented, haphazardly distributed mesh of irregular bodies comparable to algae. These discs exhibit a wide size range from a few millimeters to 75 centimeters in diameter. Exceptionally large size of the discs in present assemblage represents largest discs reported so far from any Ediacaran assemblage. Although, larger medusoid discs have been reported from USA, but they are from the Middle Cambrian and even younger rocks. Presence of microbial mats and weed-like structures with well preserved hold fasts and horizontal rhizome like structures in association with some of these large-sized discs support their animal affinity, which probably feed on this weed- like vegetations. This association also support their benthic habitat. Unlike general trend of sudden increase in size of organisms in Ediacaran period and further decrease in size during Cambrian, these discs continued increasing in size in Cambrian also. Palaeoecological distribution of such well preserved morphologies are certainly an indication of ecosystem, where algal blooms enhabited the Eiacaran organisms represented by medusoid discs and possibly serve as their food. Moderately Figure 1 Geological map of Marwar Supergroup exposed in Jodhpur District, marking the diverse Ediacaran body fossils, traces, burrows and giant sized fossil localities in Sursagar and Artiya Kalan areas (after Srivastava,2012) algal fossils occur sporadically in Jodhpur Sandstone Formation, which are well exposed in open mines in and around Sur Sagar and Artiya Kalan area of Jodhpur district (see Srivastava, 2012-c and 2013, 2015). Here in this paper we deal only with Ediacaran metazoan body fossils (impressions) and ichnofossils.

39178 | P a g e Purnima Srivastava., Soft–Bodied Ediacaran Metazoans and Ichnofossils from the Marwar Supergroup, Western Rajasthan: Emergence of Earliest Multicellular, Complex Megascopic Animal Life

Age of the Marwar Supergroup

Age of the Jodhpur Group has been suggested, younger than 681 Ma, as it unconformably overlies the Malani Igneous Suite (age given by Rathore et al., 1999, between 779-681 Ma). On the basis of medusoid body fossil Marsonia (although poorly preserved), from the Jodhpur Sandstone, Vendian (Ediacaran) age has been suggested (Raghav et al., 2005). Kumar and Pandey, 2008-b reported Aspidella and supported Ediacaran age for this stratigraphic unit. Recently reported Treptichnus pedum; an index trace fossil for Ediacaran-Cambrian boundary and fossils of an organisms, responsible for the construction of pedum burrows (Srivastava, 2012-a, b) suggested the P/C boundary at Nagaur Group. Earlier the PC/C boundary was suggested in Bilara Group on the basis of isotopic data (Pandit et al., 2001; Mazumdar and Strauss, 2006). Presence of trilobite traces was reported (Kumar and Pandey, 2008-a) from the Nagaur Group. Kumar, emphatically suggested Lower Cambrian age for the same horizon from where pedum burrows were reported in Dulmera area, Nagaur Group.

Figure 2 General lithostratigraphy of Marwar Supergroup, western Rajasthan Environment of Deposition

The sedimentation of Marwar Supergroup was supposed to be initiated during Late Proterozoic glaciation evidenced by Pokran Boulder Bed at the base. After deposition of fluvio- glacial deposition of these boulder beds, the climate of the region became semi arid evidenced by deposition of pre- dominantly yellowish brown, reddish brown arenaceous sandstone beds (Kumar, 1999). Hypersaline lagoonal, marginal marine and shallow water fair–weather wave base environment of deposition has been suggested for the fossil bearing Jodhpur Sandstone (Sarkar et al., 2008, Pandey and Tej Bahadur, 2009). Chauhan et al., (2004), interpreted ripple bedded middle part of the Jodhpur Sandstone deposited in shore-face-back shore setting of beach environment of a NNW-SSE trending wide shallow intracratonic sag basin.

Fossil assemblage from the Jodhpur Sandstone

Figure-2-a Lithocolumn of Artiya Kalan area comprising fossil The fossils are well preserved and exhibit diagnostic features of bearing beds (after Srivastava, 2012 ) their corresponding genera and species. In addition to identified ediacaran fossils, there are number of unidentified forms, informally reported here. Morphologically the present macrofossil assemblage can be broadly categorized in four, following Brigg et al., 1994, irrespective of their evolutionary relationship.

1. Most abundantly occurring category is circular impressions plausibly reminding their affinity towards jellyfish and similar medusoids cnidarians. The assemblage exhibits extensive diversity in size and morphology ranging from very small to big and from simple to little complex forms. Some of them represent the hold fasts of the frond like ediacarans. 2. Least abundant, but most significant attached, frond – like forms are Charniodiscus, first time reported from any Indian assemblage. Affinity of these forms has been suggested towards sea-pens or other soft-corals. 3. Traces and burrows probably made by bilaterians is another category. Although evidences of such activities Figure 2 b Lithocolumn of Sursagar area comprising fossil bearing are very few, but their presence can not be ruled out. beds (after Srivastava, 2012 ).

39179 | P a g e International Journal of Recent Scientific Research Vol. 11, Issue, 07 (B), pp. 39177-39191, July, 2020

4. There are number of forms very difficult to assign presence of outer rim are the features observed in different taxonomic position and are not comparable with known specimens of present assemblage. Diameter of discs varies living or fossil forms are, informally described among between 1- 10 cms (6 specimens measured). Exceptionally well problematic forms. preserved largest specimen of the present assemblage exhibits outer diameter of 75 cms (Fig 1-B). Taxonomic description

Genus- Heimalora, Fedonkin, 1992 Discussion-Hofmann, et al., (2008) considered Aspidella as cf. Heimalora sp. taphonomically variable remains, Gehling et al., 2000, interpreted these forms or discs as casts of the hold fasts or (Fig. 1-C,E; 2-B) basal impression of collapsible, hollow bulb shaped organism.

Ediacaria is also a disc-shaped fossil form with exceptional Description- Circular disc-like structure with a number of large size and bipartite to tripartite organization. It is radiating tentacles or processes of variable length and width. considered to be the largest discoid fossil of ediacaran Fossils occur on bedding plane as molds. Centrally located assemblages, which was previously interpreted as a medusoid inner disc is well preserved in few specimen (Fig.I-C,E,2-B). (Sprigg 1947), Glaessner and Wade (1966). Fedonkin (1985) Innermost subspherical body with an outer ring and uneven and Jenkins (1989) interpreted it as an attached polyp. Gehling radiating rays, appendages or processes of variable lengths are et al., (2000) have interpreted the form as cast of the basal there. The processes are straight to sinuous with .5- 1.5 mm impression of collapsible or hollow bulb shaped organisms. width, length varies between 1- 4 cms. Diameter of circular O’Brien and King (2004), merged Aspidella, Ediacaria and body vary between 1.5-4.cms., and size of complete specimens Spriggia and interpreted these forms as morphological and vary between 6-12 cms (12 specimens measured). The preservational variants. Kumar and Pandey (2009) briefly processes are sinuous or irregularly inclined. They are described this form from the Jodhpur Sandstone along with generally unbranched, but in few specimens, bifurcation can be other forms. seen. Tapering at rear ends of the processes is a common feature. Very recently, a moderately diverse and heterogenous Ediacaran discoidal assemblage has been reported by the author Discussion-Moderate preservation of Jodhpur specimens from the Jodhpur Group (Srivastava, 2012-c). Morphological assignable to Heimalora make it diffiult to identify them at diversity among these discs is biological or governed by specific level. O’Brien and King, 2004, described the discs taphonomic interplay has been discussed in that paper. with projections as tentaculated discs and inferred that the Gigantism or sudden increase in size of the Jodhpur discs and organisms may have been attached to the sea floor by these its significance has also been discussed in that paper. projections or they have functioned as tentacles comparable with those in a sea anemone or in a jelly fish. Similar Aspidella is a common and globally distributed member of the morphological forms occur in he Mistaken Point assemblage Ediacaran assemblages. At some places its association with (Anderson and Conway Morris, 1982) with possible affinities stalks, fronds and Textured Organic Surfaces (TOS) has led to to Heimalora, from the White sea region of Russia (Gehling et the interpretation of Aspidella as the hold fast of a frondose al., 2001). The overall morphological features are comparable :Charniodiscus like organisms. This organism is supposed to to the H. stellaris described from the Newfoundland, Canada live with holdfast secured within or under the sandy microbial by Hofmann et al., (2008), however Marwar forms are much mat and its stalk and frond protruding above the substrate and larger in size. Laflamme, (2010) reported discs with process also into the water column (Tarhan et al., 2015). like structures from South Australia and interpreted them as the Cyclomedusa, radiata, Crimes and Mcllroy, 1999 oldest sponges. (Fig.4-D,I) Taphonomic and morphological observations indicate that the fossil is just a part of unknown organisms (Patricia Vickers, Description: It is very common and widespread Ediacaran 2007). Discovery of a frond like structure attached to some disc. It has one of the largest size range, ranging from a few Heimalora has added an entirely different interpretation., that it millimeter to about a meter in diameter. In Ediacaran represents the hold fast of a larger organism. This interpretation assemblage in Jodhpur Sandstone Formation, diameter of discs would stand against its original classification in the medusoid ranges from 5-20 cms (12 specimens measured). Radial lines cnidarians or it would also place Heimalora among are very prominent, however, concentric lines are not well Chondrophores. Studies regarding this aspect resulted in preserved. possibilities that fragile tentacles may and may not be preserved. Discussion-Initially the form was thought to be a representative of planktic jellyfish, but now it is considered by many workers Aspidella, Billings, 1872 to have been like a benthic (bottom dwelling) polyp, somewhat like sea anemone.It reproduced by division in two or by (Fig.1- A,B: 2-C,HI) budding.

Description- Circular to elliptical discs on bedding plane with Nimbia Fedonkin, 1980 convex hyporelief. Discs in present assemblage exhibit variable (Fig.3-D,K) morphologies, some of which may be taphonomic. At the same time there is a possibility that they may belong to different Nimbia occlusa Fedonkin, 1980 affinities. Presence of outer sheath, occasional multilamellate Synonyms-Circulichnus monatus, Gureev, 1986 structure, prominent inner body, ring like structures and Annulusichnus regularis, Zhang, 1986 39180 | P a g e Purnima Srivastava., Soft–Bodied Ediacaran Metazoans and Ichnofossils from the Marwar Supergroup, Western Rajasthan: Emergence of Earliest Multicellular, Complex Megascopic Animal Life

Description- Discoid impression in convex hyporelief like Charniodiscus, but with an unusual feathery structure. At with almost centrally located smooth flat inner body, the bottom is another distinctive Newfoundland form, a disc- surrounded by annular marginal depression. Most of the shaped fossil divided into irregular lobes. This form is also specimens do not comprise any sort of ornamentation on their similar to a Charnwood Forest fossil specimen: Ivesia and surfaces. Diameter ranges from 4-13 cms. Number of appears lying on top of a "spindle" as it was buried. Jenkins and specimens measured are 22. Gehling (1978) and Glaessner (1979) regarded the frond as the most diagnostic part of the fossil. Marwar specimens are Discussion-The form is relatively common, general comparable to the fossils reported by Narbonne and Hofmann morphology is comparable to the N. occlusa from from Meitte (1987). Group of Mount Fitzwilliam area, British Columbia reported by Hofmann et al., 1991. Size of Marwar specimens is larger than Charniodiscus represents morphologically comparable to the Meitte specimens, but it is more or less comparable with the some extent a holotype specimen C. concentricus from White Sea specimens (Fedonkin, 1980), which were also Charnwood Forest (Ford, 1958). That specimen is dissimilar in preserved in convex hyporelief. Feodonkin interpreted this its branching structure to the various Charniodiscus species, fossil form as comparable with modern Trachylinid solmissus, subsequently described from the Newfoundland (Laflamme et which possess slender tentacles. al., 2004). Presence of high relief discoid holdfast connected to a clear central stem that runs up the length of the frond in Tirasiana Palij, 1976 present assemblage. Shared alignment of fronds with respect to Tirasiana sp. (Fig,5-F) their discs on many bedding planes indicate that they were Type Species- Tirasiana disciformis Palij, 1976 anchored within the substrate by their holdfasts and aligned by Description- Discoidal structure preserved in positive relief on currents (Wood etal., 2003; Liu et al., 2014). bedding plane surface. The disc consist of main disc, a second Beltanella Sprigg 1947 smaller disc and centrally located small round tubercle like structure. At places discs also exhibit presence of outer sheath (Fig. 3- F,G: 5-F) like structure. Diameter of outer disc ranges between 1.5- 6 Type species- Beltanella gilessi Sprigg 1947 cms, size of inner tubercle ranges between .5-1.5 cms (17 Synonyms- Planomedusites grandis Gureev 1987, Narbonne & specimens measured). Hofmann, 1987

Discussion-Jodhpur specimens assignable to Tirasiana are Description-Discs preserved in convex hyporelief, smooth smaller than the type material, but bigger than the specimens surface with 12- 35 mm in diameter with a prominent centrally reported from the Digermul Penninsula, Arctic Norway located tubercle (18 specimens measured). In number of (Crimes and Mcllroy, 1999). In few specimens of Jodhpur specimens, discs surrounded by a flange with irregularly spaced Sandstone, the morphology is comparable with carbonaceous marginal indentations. discs of Chuaria at megascopic level and Gloeodiniopsis lamellosa (multilamellate spheroidal body of cyanobacterial or Discussion- Marwar specimens assignable to Beltanella are chlorelloidiaceae affinity) at microscopic level. Presence of comparable in morphology with specimens illustrated by such morphologies may suggest the plant affinity for few Narbonne and Hofmann (1987) and Narbonne and Aitken, Ediacaran fossils (IV-C, VI-E). (1990) from the NW Canada. The form is traditionally categorized among pelagic medusoids (Sprigg 1947), but Charniodiscus sp. Ford, 1958 Jenkins (1988) interpreted it as a benthic polyp attached to the sea bottom by a short stalk. Type species- Charniodiscus concentricus, Ford 1958 Beltanelliformis minuta Mcllroy et al., 2005 Description- The morphology exhibited by these fossils is represented by a small stalk with frond or feather like structure (Fig. 5-G) and a holdfast like body. Jodhpur specimens exhibit moderate Description-Small-sized circular to elliptical discs occurring on preservation; hence venation like mesh reported in earlier form top of the bed in a cluster. These discs do not have any radial or is not visible. Diameter of hold fast is 1.9 cms, length of entire concentric markings. Diameter of individual disc ranges specimen is 14 cms. between 2- 3.5 mm (38 specimens measured).

Discussion-Charniodiscus is one of the frond-like Ediacaran Discussion-Marwar specimens are well comparable with B. fossils considered by some to be a ‘conventional’ cnidarian, minuta reported by Mcllroy and Walter (1997), from the possibly a pennatulacean affinity. It was earlier reported from Longmyndian Supergroup, U.K. The form is earlier reported the Ediacaran Member, Rawnsley Quartzite, Bunyeroo Gorge, from the Vindhyan Supergroup, India (Kumar and Pandey, Flinders Ranges, South Australia. Maximum length reported 2008-b). earlier is 40 cm. Jodhpur specimens are smaller in size and due to moderate preservation, minor details of morphology are not Zolotytsia, Fedonkin, 1985 preserved. This form is reported for the first time from any Zolotytsia sp. (Fig. 4-A) Indian fossil assemblage. A very similar and probably identical to a fossil named Bradgatia from the late Precambrian rocks of Description- Vermiform marking on medium to coarse grained Charnwood Forest, England (Boynton and Ford 1995). In the pinkish brown sandstone, is well preserved as convex epirelief middle we can see an unnamed fossil with a stalk and holdfast in present assemblage. Specimen is 45 cms long and 6.5.cms 39181 | P a g e International Journal of Recent Scientific Research Vol. 11, Issue, 07 (B), pp. 39177-39191, July, 2020 wide (single specimen). Hook like structure appears to be the head of organism and rear end is slightly tapered (probably the posterior end).

Discussion- As mentioned by Hofmann et al., (1991), the morphology appears to be of some soft bodied elongated, large sized organism, rather than trace fossil. The nearest analogue of present morphology is a vermiform Vendian (Ediacaran) fossil of North Carolina (Cloud et al., 1976). Laterally stacked crescentic internal features (suggestive of trace fossils back fill pattern) are not evident in present specimen. The structure is also comparable to some extent with Cylindrichnus (Glaessener 1969).

Cf. Wiwaxia sclerites Walcott, 1911

(Fig.4-E; 7-B,D)

Description- A globular or balloon shaped body with a notch or spine like structure. Maximum diameter of balloon- shaped structure is 4.5- 20 cm, length of stipe or stalk- like structure ranges between 5-17 cms (3 specimens measured).

Discussion-It is a problematic form or a fossil worm from the Burgess Shale, Walcott (1911). It was compared with modern polychaetous scale worms (Butterfield (1990). The form is not directly comparable to any modern polychaete, lacking both ventral neurochaeteae and internal acicula, the feature would be expected to preserve under Burgess Shale type conditions. It was earlier suggested as a stem group annelid.

cf. Arkarua Figure 3 Ediacaran discs from the the siliciclastic rocks of the Jodhpur Phylum- Echinodermata Sandstone in Sursagar locality. Scale in all figures represented by hammer, Class-Edrioasteroidea length of pen is 6 inches, one and two rupee coins, with diameters of 2.4 and 2.6 cms, Length of hammer is 14 inches. (Figure I- J, III-E, IV- A,B) A,B,I Aspidella sp. C,E Heimalora sp. D, K Nimbia occlusa F,G Beltanella gilessi H Tirasiana sp J cf. Arkarua sp. Description-Discoidal to hemispherical disc like organism with a pentameral star or flower-shaped dipression on surface Description- Discs ranging in size from 1-3.5 cm (6 specimens (interpreted as oral surface). When pentameral star is embossed measured) with well preserved quadriradial canal like structure that side may be the peristomal region. It comprises ambulacral and four fold symmetry are the diagnostic features. Almost groove-like structures meeting in a centrally located dipression circular in shape and margins are more or less smooth. or convex peristomal region. Polygonal to oval outlines and morphology of aboral surface is unknown. Diameter of entire Discussion- Raghav et al, 2005, first named it Marsonia, on disc ranges from 1.5 to 5 cms (4 specimens measured). the basis of its occurrence in Sonia Sandstone (name given to Jodhpur Sandstone, as one outcrop is located near Sonia Hill in Discussion- Preservation of Jodhpur specimens is Jodhpur district) of the Marwar Supergroup. It was also moderate to poor and finer details of morphology are not reported by Srivastava 2013, and assigned Scyphozoan affinity. visible, hence identification upto specific level is not possible. Quadriradial canals are the distinguishing features. Kumar et al., 2012 have also reported a giant-sized fossil Echinoid from the Jodhpur Sandstone Formation. The fossils Variable morphologies among discs preserved in siliciclastic indicate initiation of water vascular system during Ediacaran shale beds from the Artiya Kalan area, support scyphozoan time, which was till date believed to have occurred during affinity (earlier named Marsonia from the same locality), Cambrian. cnidarian affinity for small discs comparable with Funisia : considered being the first animals having sexual reproduction cf- Marsonia artiyansis (Fig.5- D). Disc’s morphologies also suggest ephyra stage (Fig. 5-B) (Fig. 8- F) of Aurilia: a cnidarian form, budding stages of

Phylum- Cnidaria some fungi, like Germinosphaera ( Fig. 6, 8- D), which are Class – Scyphozoa multicellular benthic fungal fossils and no carbonaceous discs with well preserved wrinkles and folds (Fig6-C; 8-E), like in Chuaria (Srivastava, 2013).

Unidentified Form‘A’ (Fig. 7-J)

39182 | P a g e Purnima Srivastava., Soft–Bodied Ediacaran Metazoans and Ichnofossils from the Marwar Supergroup, Western Rajasthan: Emergence of Earliest Multicellular, Complex Megascopic Animal Life

Description- Preserved as a multicellular microorganism is damaged. Another specimen: A disc exhibiting six arms is exhibiting morphology of a flat multi-chambered longitudinal difficult to interpret (Fig. V-E). section of a gastropod sheel, a leaf or a flexible sheet like structure with a distinct growth pattern. Maximum length varies between 21-48 cms, width varies between 12-40 cms (11specimens measured).

Discussion- Albani et al., (2010), reported more or less similar specimens and interpreted them as multicellular organisms from the 2.1 billion years old sediments of Francevillian Formation, Gabon. Jodhpur specimens are much larger in size in comparison to Albanis specimens.

Unidentified Form ‘B’ (Figure II-K, V-E)

Figure 5 Ediacaran fossils from the the siliciclastic rocks of the Jodhpur Sandstone in Artiya Kalan area. Scale bar in all figures representing 2 cms A-cf. Arkarua , pentameral convex body; B- Marsonia artiyansis, well preserved quadriradial canals, exhibiting fourfold symmetry in Scyphozoan affinity; C- Baloon shaped body prior to budding disc; D-Ephyra stage of Aurelia (dispersal stage); E-Two small convex discs, upper one exhibits pentameral radial ridges cf Arkarua,lower one exhibits radial ridges ,to some extent comparable to Medusinites (a polyp of cnidarians affinity); F-Colony of circular small discs with centrally located embossed tubercles, cf. Beltanella gelessi; G- Very small discs of Beltanella minuta

Discussion- Kumar et al., 2012 have reported a five armed body fossil from the Jodhpur Sandstone and interpreted that a pre-biomineralization stage of echinoderms during Ediacaran

period. As in echinoids are made up of calcium carbonate/ aragonite and present fossils are found in siliciclastic sediments Figure 4 Ediacaran discs from the the siliciclastic rocks of the Jodhpur Sandstone in Sursagar locality. Scale is represented by pen of is 6 (no evidence of calcareous sediment).

inches length and one and two rupee coins, with diameters of 2.4 and Trace Fossils and Burrows 2.6 cms respectively, A- cf. Zolotystia; B- Heimalora sp. ;C-- Aspidella sp.; D, I- Lockeia, James 1879. Cyclomedusa with well marked radial markings; E -cf . Wiwaxia sclerites with (Fig. 9-J) balloon Type species- L.siliquaria (Pelecypodichnus, Seilacher,1953) shaped body and a stipe; F- Deformed disc with sheath; G-Charnio- discus, disc or hold fast with frond; H-cf. Tirasiana; J-Nimbia occlusa Synonyms- Pelecypodichnus amygdaloides with central notch like structure and outer sheath and inner wall; K- Unidentified problematic form ‘B’, a disc with four arms, may be a precursor of Echinodermata.

Unidentified Form ‘B’ (Fig.4-K;7-E)

Description- A spherical disc with four arms like features. Disc is 11.5 cms in diameter, arms are of variable sizes. Preservation of specimen is not so good. There is a possibility that fifth arm

39183 | P a g e International Journal of Recent Scientific Research Vol. 11, Issue, 07 (B), pp. 39177-39191, July, 2020

Figure 7 Ediacaran fossils from the the siliciclastic rocks of the Jodhpur Sandstone at Sur Sagar .Scale in all figures represented by pen-, length =6 inches, one and two rupee coins, with diameters of 2.4 and 2.6 cms respectively., A- Field photograph of Sur Sagar area exhibiting Sandstone mine ; B- cf. Wiwaxia’ in which balloon shaped discs are attached to a string, as leaves to the

stem : C-Didymaulichnus, gently curved bilobate burrows with distinct median Figure 6- Ediacaran fossils from the the siliciclastic rocks of the ridge; D- cf. Wiwaxia, a balloon shaped body with stipe ; E-Problematic form , Jodhpur Sandstone in Artiya Kalan area. Scale bar in all figures central disc with six arms or appendages; F-Disc with tubercle, cf. Tirasiana ; representing 2 cms G-Unornamented disc without inner body; H- Frond like structure without hold fast ;I-Unidentified burrow exhibiting constriction at regular interval; J- Multicellular unidentified problematic form ‘A’ A,B- Circular disc with pentameral radial ridges, cf. Arkarua; C- Chuaria like morphology, exhibiting folds and wrinkles; D- Division stage exhibiting budding like structure ; E- Frond like structure with median ridge and These are simple, smooth, gently curved bilobate trails, about ornamentation 1.5 cms wide and maximum length measured in Marwar

specimens is 12 cms (8 specimens measured). The trails or Description-Small almond shaped oblong bodies preserved in burrows are preserved in convex hypo-relief; parallel to convex hyporelief, tapering to sharp and obtuse points at both bedding surface. Lobes are separated by distinct furrow, which ends. Surface commonly smooth, mostly symmetrical, length may overlap and truncate over one another. varies from 4-10 mm, width ranges between 0.5-3.5 mm (44 specimens measured). Discussion The origin of this form is speculative but - tentatively interpreted as crawling trail of molluscan origin, Discussion-The form was originally interpreted as an alga, similar to molluscan trails to some extent (Glaessner 1969). later on regarded by James (1885) as ovarian capsules of Crimes (1970) considered them traces made by trilobites. The graptolites; now considered resting trails of small burrowing form was earlier reported from the Upper Precambrian pelecypods; which were possibly semi-sessile forms. sediments of North America (Canada), and Vindhyan Didymaulichnus, Young 1972 Supergroup, India. The trace fossil Didymaulichnus miette was introduced as a new genus and species by Young, 1972, who defined it as gently curved, moderately deep, smooth, furrow- like trails, which are bisected longitudinally by a narrow median ridge. In positive epi-relief, it shows a median groove. Young reported it from the upper Miette Group (southern Cordillera of Canada) of the Terminal Proterozoic age.

39184 | P a g e Purnima Srivastava., Soft–Bodied Ediacaran Metazoans and Ichnofossils from the Marwar Supergroup, Western Rajasthan: Emergence of Earliest Multicellular, Complex Megascopic Animal Life

Other Traces and Burrows

There are number of traces and burrows which can be compared with

1. Thin trails of 1-3 mm width, winding in irregular curves of small amplitude, with short lateral thorn-like branches on one or both sides, the form is comparable to Acanthorhaphe (Fig.9-A) 2. Cylindrical tunnel like structure with axial and marginal part forms the axis. Almost similar width all along the length (Fig.9-B). One of the problematic form, which can be an imprints of marine algae, a body fossil of sponge or coral. Closest affinity of this form is, the feeding burrow (Bradley, 1973). 3. Thin elongated burrows-like structures arranged in number of rows in a circular path. Structure is comparable to some extent with Spirorhaphe (Fig.9-C). 4. Regularly meandering smooth trails cf. Cochlichnus like structure, earlier reported from Ediacaran deposits of Australia (Fig. 9-D). 5. Freely meandering simple smooth ridges of extraordinary regular form comparable to some extent with grazing trails of Cosmorhaphe (Fig. 9-E). 6. Tree like trails, straight or curved, side branches are also present, where length varies from specimen to specimen, cf. Dendrotichnium (Fig. 9-F). 7. Narrow intermittent double drag trails, irregular and closely set, interpreted as tracks of limulids or gastropods (Fig. 9-G, H, K).

8. U-shape in cross section and circular, double or triple Figure 9 Traces and burrows from the the siliciclastic rocks of walled structure cf Diplocraterion (Fig. 9- I). In Jodhpur the assemblage only surface view observed. Jodhpur Sandstone (of Ediacaran age), in Sursagar locality. Scale in all figures represented by hammer of 14 inches, pen of 6 inches length, one and two rupee coins, with diameters of 2.4 and 2.6 cms respectively A-cf. Acanthorhaphe, B- cf. Zoophycus, C- cf. Spirohaphe,D-cf. Cochlichnus, E- cf. Cosmorhaphe, F- cf. Dendrotichnium, G,H,K- cf. Protichnites, I- cf. Diplocraterion, J- cf. Pelecypodichnus

Figure 8- Ediacaran fossils from the the siliciclastic rocks of the Jodhpur Sandstone in Artiya Kalan area. Scale bar in all figures representing 2 cms A-Circular disc with tube like projection ;B-Division stage, exhibiting merged wall between two discs ; C- Arumberia like structure ;D- Budding like Str; E- Chuaria like morphology exhibiting folds and wrinkles; F- Ephyra stage of Aurelia (moon jelly fish) ;G- Marsonia, disc with quadriradial canals in

Scyphozoan affinity. Figure 10 Microbial mats and Arumberia banksi from the siliciclastic rocks of the Jodhpur Sandstone (of Ediacaran age), in Sursagar locality, exhibiting different structures.

39185 | P a g e International Journal of Recent Scientific Research Vol. 11, Issue, 07 (B), pp. 39177-39191, July, 2020

DISCUSSION AND CONCLUSION atmosphere) and sufficient supply of water. If this condition is supplemented by low energy hydrodynamic conditions and The Ediacaran organisms remain one of the greatest enigmas absence of mat consumers, cyanobacteria may spread out on within evolutionary palaeobiology. It has now been a general the sediment surface. Resulting organic matter accumulation is agreement that some Ediacaran fossils are indeed cnidarian and balanced by re-mineralization and degradation of the organic bilaterian metazoans but their zoological affinities are still a material via other bacteria by processes like aerobic and matter of debate. There are number of problematic forms in the anaerobic respiration. Within this complex layered mat system, present assemblage, which are very difficult to decide whether the surface layer is of specific importance, concerning the they belong to plant or an animal kingdom. In number of cases formation of mat related structures and their preservation. in the history of palaeontology, these problematic fossils were extensively used in ecological and evolutionary studies before Like other micro or macro organisms, gigantisms is well anyone could be certain of their systematic affinities even at marked among Ediacaran macro organisms from the Jodhpur kingdom or phylum ranks, Waggoner (2003). Sandstone, whether belonging to plant or animal kingdom. Maximum diameter of Jodhpur discs reached up to 75 cms, Epifaunal tiering has not been exhibited by Marwar fossil which is probably the largest Ediacaran disc reported so far. assemblage, which supports the true representation of Ediacaran Fauna. The assemblage also shows biomat Most of the fossils from the present assemblage inclined dependant palaeoecological as well as palaeoenvironmental towards animal affinity, except discs with well marked folds setting of Neoproterozoic. The assemblage is dominated by and wrinkles. Specimens are well comparable in morphology disc shaped organisms assignable to medusoid affinities and with carbonaceous megafossils Chuaria, established taxa, of some considered being resting impressions of hold fast like plant affinity. Size and presence of carbonaceous matter are the structures. A fairly good population of small sized cluster of features differentiating these taxa from the Ediacaran fossils. mounds, assignable to Beltanelli form is has also been reported The unique biology, preservational mode and ecology of the from the present assemblage. Ediacaran biota marks the end of Proterozoic Eon and may

Ediacaran fossils are generally restricted to 635-541 Ma old herald the beginning of the Phanerozoic Era (Narbonne, 1998). sediments all over world. Global dominance of Ediacaran discs These fossils occupy a pivotal position in the evolution of life can be seen and a distinct group of complex macroscopic on Earth, between largely microbial communities organisms that flourished on the eve of Cambrian radiation has characterizing Precambrian and shelly biota of Cambrian and taken place. The Ediacaran organisms have been considered the younger Phanerozoic system (Sepkoski, 1981). precursors to Cambrian animals, including cnidarians, annelids, Evolutionary History arthropods and echinoderms (Gehling, 1991). Despite high taxonomic diversity and wide geographic distribution, the Prior to the appearance of Ediacaran biota, benthic phylogenetic interpretation of many Ediacaran fossils is communities had been dominated by prokaryotic microbes controversial. Since Ediacaran fossils occur as casts and molds along with some algae (Knoll, 1992). Evidences from of organisms, the real body of the original creature is not molecular phylogeny suggest that microscopic animals could preserved, which has preluded the study of actual nature of have evolved much earlier but the exact timing remains these organisms and created difficulty in their biologic debatable (Wray et al., 1996, Morris and Collins, 1996). The classification. However, the Ediacaran fossils represent many oldest Ediacaran fossils occur in Twitya Formation, unique taxa which are found to be extinct at the Ediacaran- northwestern Canada, immediately below tillites, correlated Cambrian transition and follow initial morphological expansion with Marinoan- Varangerian glaciation (Hofmann, et al., 1990). and subsequent taxonomic saturation within an established A global rise in atmospheric oxygen in the Late Proterozoic morphospace (Shuhai Xiao, 2010). may have been the trigger that permit animals to achieve megascopic size (Canfield and Teske, 1996). Very peculiar and most probably the characteristic feature of Jodhpur Ediacaran assemblage is the association of macro-algal Affinities Virtually most of the workers emphasized similarities structures and microbial mats (Fig.10), with medusoid discs. in two dimensional structures between Ediacaran fossils and Palaeoecological distribution of such morphologies, indicate living groups of jelly fish, soft corals, annelids and arthropods the ecosystem of that time, when algal bloom flourished with and suggested that the Ediacaran biota represent the direct the Ediacaran organisms (mostly of animal affinity) and ancestors of these modern groups (Narbonne, 1998). Seilacher, probably served as their food. The presence of macro algae not 1992, suggested that Ediacaran organisms be referred to as a new phylum “Vendozoa’, which consist of quilted organisms only improved the environment at that time (enhanced the O2 level) but also played an essential support for the growth, that lacked mouths and guts and received energy by absorbing evolution and propagation of metazoans which directly related dissolved organic molecules or by harboring photosynthetic or to the oxygen level. chemosynthetic symbionts. Now Seilacher term this phylum ‘Vendobionta’ and consider them as true animals. Narbonne The modern cohesive microbial mats developing in peritidal (1998) placed these fossils into a single taxonomic group, be it siliciclastic systems, mostly settle on clear fine grained sands of jellyfish, protists, lichens or vendozoans, which is unacceptable the intertidal to lower supratidal zones (Gerdes and Krumbein, by scientific community. These fossils exhibit a wide size 1987; Gerdes et al., 2003; Noffke et al., 2001, 2003).In this range, variable symmetry and their resistance towards particular environment, the mats are constructed by decomposition. It is now believed that different taxa lived cyanobacteria, because these zones are favourable sites for together under conditions ranging from shallow to deep sea oxygenic photosynthesis in which cyanobacteria provide floors. In present assemblage the fossils have been categorized abundant energy (from solar radiation), Co2 (from the under three groups. These are: 39186 | P a g e Purnima Srivastava., Soft–Bodied Ediacaran Metazoans and Ichnofossils from the Marwar Supergroup, Western Rajasthan: Emergence of Earliest Multicellular, Complex Megascopic Animal Life

occur frequently in fossil bearing horizon. It is therefore 1) Bottom dwelling (benthic) non resistant discs, considered to inferred that most fossiliferous strata were deposited between have been permanently attached to the sea bottom (Seilacher wave base and storm wave base, and were probably within the 1992; Narbonne and Aitken 1990; Gehling 1991, Fedonkin and euphotic zone. It is observed that Ediacaran ecosystem was Waggoner, 1997). Beltanelliformis; a centimeter scale discs are different from the modern systems. Microbial mats were the most common Ediacaran fossils world-wide, which are abundant and covered maximum marine surfaces, which has comparable in morphology to the base of some anemones and affected sedimentation pattern of that time. Burrowing also to some extent with Palaeozoic burrows attributed to organisms were rare and were probably ineffective anemones ( Narbonne and Hofmann, 1987, Fedonkin and bioturbators. It is also inferred that animal preying on soft Runnegar, 1992). Larger discs with hair like projections bodied immobile organisms were not evolved by that time. (Heimalora), have been regarded as the tentacles of a polyp or the root like structures of a hold fast (Runnegar, 1995). Correlation 2) This category comprises quilted structures (very rare in Microbial mat structures like Arumberia and ediacaran present assemblage) considered to be true ‘vendobionts’. These metazoans like Beltanella, Aspidella, Heimalora and organisms were regarded as representing the ancestors of Charniodiscus are typical Ediacaran fossil forms reported from several modern groups ranging from soft corals to annelid the Bonavista Peninsula of the Newfoundland, Canada, Mcllroy worms. et al., 2005 and Hofmann et al., 2008. Almost similar 3) This category comprises trace fossils, representing simple assemblage (excluding Charniodiscus) has also been reported subhorizontal burrows. Presence of these fossils is indicative of from the Maihar Sandstone of the Vindhyan Supergroup, India, mobility and implies concentration of sensory organs at a head assigned Ediacaran age (Kumar and Pandey, 2009). This region (Narbonne, 1998). Previously these fossils were correlation has a palaeogeographic implication. During considered as the work of bilaterian animals (Seilacher 1989, Ediacaran period both Marwar and Vindhyan Basins existed Fedonkin and Runnegar, 1992). It was concluded that and might have linked with each other through a corridor of Vendobionta were distinctive clade of Neoproterozoic Lesser Himalaya, represented by the Krol deposits. During that organisms, but its taxonomic position is uncertain; different period, sedimentation in the Vindhyan Basin ended as there is views ranging from an extinct group within the phylum no evidence of Cambrian deposits, but it continued in the Cnidaria (which also includes modern corals and anemones) to Marwar basin upto the Lower Cambrian. Regression started an extinct phylum unrelated to subsequent Phanerozoic from the west via present day Himalayan zone and reached evolution of animals (Buss and Seilacher, 1994). western Rajasthan during the Lower Cambrian. It continued further in NW in the Salt Range in Pakistan (Kumar and Preservation Siliciclastic sediments are not considered to be Pandey, 2009). appropriate for the fossils preservation with minor details. In contrast despite soft bodied animals of Ediacara affinity, these CONCLUSION organisms are well preserved. Generally these fossils are found in event beds, but in Jodhpur Sandstone, these beds have not There is a possibility that Jodhpur macrofossil assemblage of been observed. It is believed organisms lived on the mud body fossils, macro algal forms, microbial mats and trace bottom and their impressions were preserved, when the fossils, collectively represents one of the best repository for organisms catastrophically covered by a bed of sand-sized evolutionary palaeobiology. The assemblage is also debris (Narbonne, 1998). Preservation of Ediacarans in warty, representing a transitional phase of megascopic life proceeding putular texture that was referred as ‘old elephant skin’ has from water to land. It seems that megascopic plant life, was provided the search image for Ediacaran fossils. These are probably heading a step towards terrestrialization at the time of rarely found in beds that do not contain Ediacaran fossils. Jodhpur Sandstone sedimentation. Coexistence of terrestrial Gehling (1987) first recognized the identical features in modern plant like structures (with well developed Hold fast like microbial mats and concluded that these mats were structures, stolon like horizontal rhizome- like structures) with instrumental in stabilizing the mud surface and providing a aquatic communities (Ediacaran metazoans, microbial mats and ‘death mask’ on the organism itself. Bioturbations, cropping acritarchs) also reflect an evolutionary phase of aquatic life, and competitions seem to be the reason of abruptly declined from coastal marine to land environment. Since the present mat occurrences in Cambrian and negligible preservation of paper is based on preliminary study, more details and extensive soft –bodied organisms in sandstone rocks. It was inferred that study is required to draw substantial conclusion and more clear disappearance of Ediacarans was not solely preservational as picture of evolution and ecosystem at the time of sedimentation these fossils are almost absent in Cambrian Lägerstäten such as of Jodhpur Sandstone.

Burgess Shale. References Ecology Albani, A.E., Bengtson, S., Canfield, D.E., Bekker, A., Now, it is believed that Ediacaran biota mainly consist of Macchiarelli, R.,Mazurier, A., Hammarlund, E.U., benthic organisms, which are mostly preserved in their original Boulvais,P., Dupuy, J.J.,Fontaine, C., Fursich, F.T., position. Most of the Ediacaran fossil sites are shallow- water Lafaye, deposits (Gehling, 1998, Narbonne et al., 1997; Grazhdankin F.G.,Janvier,P.,Javaux.E.,Ossa.F.O.,Wickmann,A.C.,Rib and Ivantsov, 1996). In Marwar Supergroup also, fossil bearing oulleau,A.,Sardini,P.,Vachard, D.,Whitehouse, M and Jodhpur Sandstone exhibits wave ripples, cross bedding and Meunier, A. 2010, Large colonial organisms with occasionally occurring desiccation cracks. Microbial mats 39187 | P a g e International Journal of Recent Scientific Research Vol. 11, Issue, 07 (B), pp. 39177-39191, July, 2020

coordinated growth in oxygenated environments 2.1 Gyr Crimes, T. P., 1970. Trilobite tracks and other fossils from ago., Nature 466, 100-104 the upper Cambrian of North Wales, Geological Journal, Anderson, M. M. and Conway Morris, 1982. A review with 7(1): 47-68. descriptions of four unusual forms of the soft bodied Crimes, T. P. and Mcllroy, D. 1999, “A biota of Ediacaran fauna of the Conception and St. John’s Groups (Late aspect from lower Cambrian strata on the Digermul Precambrian), Avalon Peninsula Newfoundland. Peninsula, Arctic Norway”,Geological Magazine 136, Proceedings of the Third North American pp.633-642. Palaeontological Convention, 1:1-8. De, C. and Prasad, S. 2012. Discovery of Ediacaran Fossil Battison, L. and Brasier, M. D. 2009. Exceptional assemblage from the Marwar Supergroup of Rajasthan., Preservation of Early Terrestrial Communities in Geol. Surv. India, western region. Lacustrine Phosphate one Billion Years Ago, in Smith, Erwin, D. H., Laflamme, M,. Tweedt, S.M., Sperling, E.A., M. R., O’Brien L. J., Caron, J-B International Pisani, D. & Peterson, K.J., 2011. The Cambrian Conference on the Cambrian Explosion (Walcott 2009) Conundrum: Early divergence and later ecological3 Abstract Volume. Toronto, Ontario, Canada: The success in the early history of animals. Science, 334 Burgess Shale Consortium. (6059), 1091- 1097. Billings, E., 1872. On some fossils from the primordial rocks Farmer, J., Vidal, G., Moczydlowska, M., Strauss, P., of Newfoundland:Canada. Naturalist, n.ser. 6:465-479 Ahlberg, P., and Siedlecka, A., 1992. Ediacaran fossils pp. from the Innerly Member (Late Proterozoic) of the Boynton, H.E., and Ford, T. D., 1995. Ediacaran fossils from Tanafjorden area, northern Finnmark, Geol. Mag.129 the Precambrian (Charnian Supergroup) of Charnwood (2): 181-195. Forest. Leicestershire, England, Mercian Geologist, Fedonkin, M. A. 1980. New Precambrian Coelentrata in the 13:165-182. T. P.Crimes and D. Mcllroy, “A biota of north of the Russian Platform . Palaeontological Journal, Ediacaran aspect from lower Cambrian strata on the 2:1-10 Digermul Peninsula, Arctic Norway”,Geological Fedonkin, M. A. 1981. Belomorskaya biota venda Magazine 136, 1999633-642. (dokembriskaya berskeletnaya fauna severa Russkoi Bradley, J., 1973. Zoophycus and Umbellula: their synthesis Platform (The White Sea Biota of the and identity, Palaeogeograpgy, palaeoclimatology and Vendian:Precambrian soft bodied fauna of the northern palaeoecology, 13: 103-128. Russian Platform). Moscow, Nauka. Brigg, D.E.G, Erwin, D. H. and Collier, F. J., 1994. The Fedonkin, M. A. 1985. Systematic description of Vendian- fossils of the Burgess Shale, Smithsonian Institution Metazoa, in B. S. Sokolov and A, B.Ivanovsky (eds) Press, Washington and London. Vendskaya sistema, Istoricko-geologicheskoe; Buss, L. W. and Seilacher, A.,1994. The phylum paleontologicheskoe obosnovanie. T.I. Paleontologia, Vendobionta: A sister group of the Eumetazoa?, Nauka, Moscow. 70-112, Palaeobiology, v. 20, 1-4. Fedonkin, M. A., Gehling, J.G., Grey, K., Narbonne, G.M. & Butterfield, N. J. 1990. Organic preservation of Vicker Rich, P. 2007. The Rise of animals: Evolution nonmineralising organisms and the taphonomy of the and diversification of the Kingdom Animalia, Baltimore: Burgess Shale, Palaeobiology, 16: 272-286. John Hopkins, Univ. Press. Bykova, N. V., 2011. Ediacaran soft- bodied organisms and Fedonkin, M. A. and Runnegar, B. 1992. Proterozoic macrophytes: Two sides of one coin? In Neoproterozoic metazoan trace fossil in Schopf, J.W Klein, C. (eds.), Sedimentary Basins: Stratigr., Geodynamics and The Proterozoic Biosphere: A multidisciplinary study, Petroleum Potential, Proc. Internat. Conf. (Novosirbirsk: Cambridge, UK, Cambridge University Press. 389-396. Trofimork. Inst. Petrol. Gas . Geolo. Geophys. Sib. Fedonkin, M. A. and Waggoner, B. M. 1997. The Late Branch Russia. Acad. Sci. 14-15 Precambrian fossil Kimberella is a mollusk-like Canfield, D. E. and Teske, A., 1996. Late Proterozoic rise in bilaterian organism, Nature, v. 388, 868-871. atmospheric oxygen concentration inferred from Ford, T. D. 1958. Precambrian fossils from Charnwood phylogenetic and sulphur isotope studies. Nature, v. 382, Forest. Proc. Yorks. Geol. Soc. Vol. 31, 211-217 127-132. Gehling, J.G., 1987. Earliest known echinoderm- A new Chauhan, D.S., Ram, B, and Ram,N., 2004. Jodhpur Ediacaran fossil from the Pound Subgroup of South Sandstone: A gift of ancient beaches to western Australia, Alcheringa, v.11, 337-345. Rajasthan. Jour. Geol. Soc. Ind. 64: 265-276. Gehling, J.G., 1991. The case for Ediacaran fossil roots to the Cloud, P. E., Wright, J., and Glover , L., 1976. Traces of metazoan tree. In Radhakrishna, B. P. (ed): The world of animal life from 620 million year old rocks in North Martin F. Glaessner, 181-224. Geological Society of Carolina, American Scientist, 64, 396-406. India, Memoir 20. Geol. Soc. of India, Bangalore. Conway Morris, S. 1976. Molecular Clocks: Defusing the Gehling, J.G., 1999. Microbial mats in terminal Proterozoic Cambrian Explosion; Current Biology, v, 7, R71-R74. siliciclastics: Ediacaran death masks. Palaios,14, 40-57 Conway Morris, S., 1993. Ediacaran –like fossils in Gehling, J.G., Narbonne, G.M. and Anderson, M.M., 2000. Cambrian Burgess Shale type faunas of North America, The First named Ediacaran body fossil, Aspidella Palaeontology, 36:593-635. terranovica. Palaeontology, 43(3): 427-456. Conway Morris, S. and Collins, D. H., 1996. Middle Gehling, J.G., Jensen, S., Droser, M. L.Myrow, P. M., and Cambrian ctenophores from the Stephen Formation, Narbonne, G. M., 2001. Burrowing below the Basal British Columbia, Canada. Phil.Trans. Roya soc. Cambrian GSSP, Fortune Head, Newfoundland, London, B.351: 27308. Geological Magazine, 138: 213-218. 39188 | P a g e Purnima Srivastava., Soft–Bodied Ediacaran Metazoans and Ichnofossils from the Marwar Supergroup, Western Rajasthan: Emergence of Earliest Multicellular, Complex Megascopic Animal Life

Gerdes, A. and Krumbein, W.E., 1987. Biolaminated Jenkins, R. J. F. and Gehling, J.G., 1978. A review of the deposits.In: Bhattacharya, G.M., Friedmann, G.M., frond like fossils of the Ediacara assemblage. Records of Neugebauer, H.J. and Seilacher, A.(Eds), Lecture notes the south Australian Museum, 17(23): 347-359. in Earth Sciences, 9. Springer-Verlag, Berlin, pp 1-183. Khan, E.,A., 1971.Geological mapping in parts of Jodhpur Gerdes, A., Friedt, G.,Parrish, R. R. and Finger, F., 2003. and Nagaur district , Rajasthan, In Seminar on Oil, Gas High resolution geochronology of Variscan granite and Lignite scenario with special reference to Rajasthan, emplacement, the south Bohemian Batholith., Jour. of pp. 69-72. Czech Geological Society, 48:53-54. Kumar, S. and Pandey, S. K., 2008-a. Discovery of trilobite Glaessner, M. F., 1962. Precambrian fossils, Biol. Rev. vol. trace fossils from the Nagaur Sandstone, the Marwar 37, 467-494. Supergroup, Dulmera area, Bikaner district, Rajasthan, Glaessner, M. F., 1969 .Trace fossils fr om the Cambrian and Curr. Sci, 94(8): 1081-1085. basal Cambrian. Lethaia 2, 369-393. Kumar, S. and Pandey, S. K., 2008-b. Arumberia and Glaessner, M. F., 1979. Precambrian, In: Robinson, R.A., associated fossils from the Neoproterozoic Maihar Teichert, C. (Eds), Treatise on Invertebrate sandstone, Vindhyan Supergroup, Central India. Jour. Palaeontology. Part A. Geological Society of America, Pal. Soc. India, 53(1):83-97 pp79-118. Kumar, S. & Pandey, S. K. 2009. Note on the occurrence of Glaessner, M. F., 1984. The dawn of animal life: A Arumberia and associated fossils from the Jodhpur biohistorical study. Cambridge , UK, Cambridge Sandstone, Marwar Supergroup, Western Rajasthan, University Press, 244pp. Jour. Pal. Soc. Ind. 54(2), 171- 178. Glaessner, M. F., and Daily, B. (1959) "The Geology and Kumar, S., Srivastava, D.K. and Ahmad, S.2012. Five armed Late Precambrian Fauna of the Ediacara Fossil body fossil from the Ediacara Jodhpur Sandstone, Reserve". Records of the South Australian Museum 13: Marwar Supergroup, western Rajasthan, India: A 369-407 retrieved 26 January 2008 possible precursor of phylum Echinodermata. Curr. Sci. Glaessner, M. F. and Wade, M., 1966. The late Precambrian 10291):24-26. fossils from Ediacara, South Australia, Palaeontology, 9: Kumar, V., 1999. Evolution and Geological set up of the 599- 628. Nagaur-Ganganagar Basin, North Western Rajasthan, In: Grazhdankin, D. V. and Ivantsov, A.Y., 1996 Reconstruction Paliwal, B, S.(Ed.) Geological Evolution of of Biotopes of ancient Metazoa of the late Vendian Northwestern India, Scientific Publishers, India, pp.34- White Sea biota. Palaeontological Jour. 30:674-678. 60. Grotzinger, J. P., Bowering, S. A., Saylor, B. Z. and Knoll, A. H., 1992. Biological and biochemical preludes to Kaufman, A. J., 1995. Biostratigraphic and the Ediacaran radiation. In: Lipps, J. H. and Signor, P. Geochronologic constraints on early animal evolution. W.(eds) Origin and early evolution of metazoan. New Science, vol. 270, 598-604. York, Plenum Press, 53- 86. Gureev, Y. A., 1986. On the perspectives of Kumar, S. and Pandey, S. K., 2009. Note on the occurrence palaeoichnological methods in stratigraphy. Tectonica of Arumberia banksii and associated fossils from the Stratig.27: 42-47 (in Russian). Jodhpur Sandstone, Marwar Supergroup, Western Gureev, Y. A.,1987. Morphological analyses and systematic Rajasthan., Jour. Palaeontol. Soc. India, 54(2): 171- 178. of the Vendiata. Akademiya Nauk Ukrainskoy,SSSR, Kumar, S., Misra, P.K. and Pandey, S. K., 2009. Ediacran InstitutGeologicheskikh Nauk,87-15:1-54. megaplant fossil with Vaucherian affinity from the Hofmann, H. J., Narbonne, G. M. and Aitken, J. D. 1990. Jodhpur Sandstone , Marwar Supergroup, Western Ediacaran remains from inter tillite beds in Rajasthan., Curr. Sci. 97(5):701-705. northwestern Canada, Geology, v.18,1199-1202. Laflamme, M.,2010.Wringing out the oldest sponges. Hofmann, H. J., Mountjoy, E.W., and Teitz, M.W., 1991. Nature Geoscience, Advance online Publication Ediacaran fossils and dubiofossils, Miette Group of Laflamme, M., Narbonne, G. M. and Anderson, M. M., 2004. Mount Fitzwilliam Area, British Columbia, Canadian Morphometric analyses of the Ediacaran frond Jour. of Earth Sciences, 28:1541-1552. Charniodiscus from the Mistaken Point Formation, Hofmann, H. J., O’Brien, S. J. and King, A. F., 2008. Newfoundland. Jour. of Palaeontology, 78, 827-837. Ediacaran biota on Bonavista Peninsula, Newfoundland, Liu, A, G., Kenchinton, C. G. and Mitchell, E. G., 2014. Canada. Jour. of Palaeontol. 82(1): 1-36. Remarkable insight into the palaeoecology of the James, J. F., 1885.The fucoids of the Cincinnati group: Avalonian Ediacaran macrobiota. Gond. Res. Cincinnzti Soc, Nat.Hist. Jour.vol.7,no.4, p151-166 http://dx.doi.org/10.1016/J.gr 2014.11.002 James, U.P., 1879. Description of new species of fossils and Mazumdar, A. and Strauss, H., 2006. Sulfur and Strontium remarks on some others from the Lower and Upper isotopic compositions of carbonate and evaporate rocks Slurian rocks of Ohio. The Paleontologist, 3:17-24. from the Late Neoproterozoic –Early Cambrian Bilara Jenkins, R. J. F. 1984. Interpreting the oldest fossil Group (Nagaur- Ganganagar Basin, India): Constraints cnidarians. Palaeontogr. Am. , vol. 54, 95-104. on intrabasinal correlation and global sulfur cycle. Jenkins, R. J. F. 1989. The “supposed terminal Precambrian Precambrian Research 149, 217-230. extinction event” in relation to the Cnidaria, Mem. Ass. Mcllroy, D. and Walter, M. R., 1997. A reconsideration of the Australas Palaeontols 8: 307-317. biogenicity of the Arumberia banksi Glaessner and Walter. Alcheringa: an Australasian Journal of

39189 | P a g e International Journal of Recent Scientific Research Vol. 11, Issue, 07 (B), pp. 39177-39191, July, 2020

Palaeontology, 21: 79, doi: Paroda, H. and Bouougri, E. 2008. Neoproterozoic trace 10.1080/03115519708619187. fossils vs microbial mat structures: Example from the Mcllroy, D., Crimes, T.P. and Pauley, J. C., 2005. Fossils and Tandilia Belt of Argentina. Gond. Res., 13: 480-487. matgrounds from the Neoproterozoic Longmyndian Patricia, V.R., Komarower, P. (Eds).2007. The rise and fall of Supergroup, Shropshire, UK, Geological Magazine, 142: the Ediacara Biota. Geol. Soc. London, Special 441-455. Publication, 286, pp 440. McMenamin, M. A. S., 1998. The Garden of Ediacara: Peterson, K. J., Waggoner, B, and Hagadorn, J. W., 2003. A Discovering the first complex Life. New York, Columbia fungal analog for Newfoundland Ediacaran fossils? University Press. Integr. Comp Biol. 43(1), 127-136. Narbonne, G. M., 1998. The Ediacara Biota: a Terminal Pflug, H. D. 1974. Vor-und Fruhgeschicht der Metazoa, N. Neoproterozoic experiment in the evolution of life. GSA Jb. Geol. Palaeontol., Abh. Vol. 145, 328-374. Today 8 (2), 1-6. Poire, D. G., 1984. Trazas fosiles en euarcitas de la formacion Narbonne, G. M., 2005. The Ediacara Biota: Neoproterozoic sieras Bayas (Precambrico) y su comparicion con las de origin of animals and their ecosystem. Ann. Rev, Earth la Formacion Balcarce (Cambro-Ordov), Sieras Planet. Sci. 33, 421-442. Septentrionales de la provincial de Buenos Aires. Narbonne, G. M. and Aitken, J. D. 1990. Ediacaran fossils Congreso Geologico Argentino, San Carlos de from the Sekwi Brook area, Mackenzie Mountains, Bariloche, Actas, iv, 9, 249-266. northwestern Canada: Palaeontology, v. 33, 945-980. Raghav, K. S., De, C. & Jain, R. L., 2005. The first record of Narbonne, G. M. and Hofmann, H. J., 1987. Ediacaran biota Vendian medusoids and trace fossil bearing algal mat of the Werneck Mountains, Yukon, Canada, ground from the basal part of the Marwar Supergroup of Palaeontology, v. 30, 647-676. Rajasthan, India. Ind. Min., 59 (1-2), 23- 30. Narbonne, G. M., Dalrymple, R. W. and MacNaughton, R.B., Rathore, S. S. Venkatesan, T. R. and Srivastava, R. K., 1999. 1997. Deep-water microbialites and Ediacara-type fossils Rb/ Sr isotope dating of Neoproterozoic (Malani Group) from North Western Canada, Anstr. Prog. Geol. Soc. Magmatism from southwest Rajasthan, India: Evidence Am. 29, A-193. of younger Pan- African Thermal Event by 40Ar- 39Ar Noffke, N., Gerdes, G., Klenke, T. and Krumbein, W.E., studioes. Gondwana Research 2(2), 271- 181. 2001. Microbially induced sedimentary structures- A Retallack, G. J. 1994. Were the Ediacaran fossils Lichens? new category within the classification of primary Palaeobiology, 20(4), 523-544. sedimentary structures. Jour. of Sedimentary Research, Runnegar, B., 1995. Vendobionta or Metazoa? Developments 71: 649-656. in understanding the Ediacara fauna., Neus Jahrbuch fur Noffke, N., Gerdes, G. and Klenke, T. 2003. Benthic Geologie und Palaeontologie Abhandlunden, v 195, 303- cyanobacteria and their influence on the sedimentary 318. dynamics of peritidal depositional systems (siliciclastic Sarkar, S., Bose, P. K., Samantha, P., Sengupta, P. & salty and evaporitic carbonatic). Earth Science Reviews Eriksson, P., 2008. Microbial mat mediated structures in 62: 163-176. the Ediacaran Sonia Sandstone, Rajasthan, India and O’Brien, S. J. and King, A. F., 2004. Ediacaran fossils from their implications for Proterozoic sedimentation, the Bonavista Peninsula (Avalon zone), Newfoundland: Precamb. Res. 162(1-2), 248- 263. Preliminary descriptions and implications for regional Shanker, R., Bhattacharya, D. D., Pandey, A. C. and Mathur, correlation. Current Research (2004), Newfoundland, V. K., 2004. Ediacaran biota from the Jarashi (Middle Department of Mines and Energy, Geol. Survey report, Krol) and Mahi (Lower Krol) Formations, Krol Group, vol.1, 203-212. Lesser Himalaya, India. Geol. Soc. India, 63 (6), 649- Pandey, D. K. & Tej Bahadur, 2009. A Review of the 654. stratigraphy of Marwar Supergroup of west-Central Seilacher, A., 1953.Studien zur Palichnologie. Z. Die Rajasthan. Journal Geological Society of India 73, 747- fossilien Ruhespuren (Cubichnia), Neues Jb. Geolo. 758. Palaontol. Abh. 98, 87-124. Palij, V. M., 1976. Remains of soft- bodied and trace fossils Seilacher, A., 1989. Vendozoa: Organismic construction in from deposits of Upper Precambrian and Lower the Proterozoic Biosphere, Lethaia, 22, 229-239. Cambrian. In: Ryabenko (ed).Paleontology and Seilacher, A., 1992: Vendobionta and Psammocorallia Lost stratigraphy of upp. Precamb. And Lr. Paleoz.of SWof construction of Precambrian evolution, Jour. Geol. Soc. Eastern-European Platform. Naukova dumka. 63-76. London, 149, 607-613. Pandit, M. K., Sial, A. N., Jamrani, S. S. & Ferreira, V. P., Seilacher, A., Grazhdankin, D. and Legouta, A. 2001. Carbon isotope profile across the Bilara Group 2003.Ediacaran Biota: the Dawn of animal life in the rocks of Trans Aravalli Marwar Supergroup in Western shadow of giant protists. Palaeontol. Res. 7(1), 43-54. Rajasthan, India: Implications for Neoproterozoic- Sepkoski, J. J., 1981. A factor analytic description of the Cambrian transition. Gond. Res. 4: 387- 394. Phanerozoic marine fossil record., Palaeobiology, vol. 7, Paliwal, B.S., 1998. Felsic volcanic interlayered with 36-53. sediments of the Marwar Supergroup at Choti Khatu, Serezhnikova, E. A, Ragozina, A. L., Dorjnamjaa, D., District Nagaur, Rajasthan. Jour. Geol. Soc. India., andZaitseva, L. V. 2014. Fossil microbial communities 52:81-86. in Neoproterozoic interglacial rocks of western Pareek, H. S. 1984. Pre-Quaternary Geology and mineral Mongolia, Precamb. Res. 245: 66-79. resources of northwestern Rajasthan. Memoir of Sharma M., Kumar, S., Tewari M., Shukla Y, Pandey, S. K,, Geological Survey of India. 115: 1-95. Srivastava, P. and Banerjee, S., 2012. 39190 | P a g e Purnima Srivastava., Soft–Bodied Ediacaran Metazoans and Ichnofossils from the Marwar Supergroup, Western Rajasthan: Emergence of Earliest Multicellular, Complex Megascopic Animal Life

Palaeobiological Constraints and the Precambrian Tarhan, L. G., Droser, M. L., Gehling, J. G., Dzaugis, M. P., Biosphere: India Evidence, Proc Indian Natn Sci. Acad. 2015. Taphonomy and morphology of the Ediacara form 78 (3), pp.1-17. genus Aspidella . Precambrian Research, 257, 124- 136. Sprigg, R.C. 1947. Early Cambrian (?)Jelly fish from the Wade, M., 1968. Preservation of soft bodied animals in Flinders Ranges, S. Australia, Trans. R. Soc. South Precambrian sandstone at Ediacara , South Australia. Australia, no. 71, 221-224. Lethaia, v. 1, 238-267., B., 2003. The Ediacaran bita in Sprigg, R.C. 1949. Early Camb. Jellyfishes of Ediacara; space and time. Integrative and comparative Biology, South Australia and Mount John, Kimberley district, 32:104-113. western Australia, Trans. R. Soc. South Australia, no. 73, Waggoner, B. 2003. The Ediacaran biota in space and time, 72-79. Integrative and Comparative Biology, 32:104-113. Srivastava, P. 2006. Possible Ediacaran fossils from the Walcott, C. D., 1911. Report of the Director, Smithsonian Bundi Hill Sandstone, Upper Vindhyans, Rajasthan. Institute Ann. Report. 1910, 1-39. Diamond Jubilee International Conference on Changing Wood, D. A., Dalrymple, R. W., Narbonne, G. M., Gehling, Scenario in Palaeobotany and allied subjects, held at J. G. and Clapham, M. E., 2003. Palaeoenvironmental BSIP, Lucknow, Nov. 2006, p-188. analyses of the Late Neoproterozoic Mistaken Point and Srivastava, P. 2012-a. Treptichnus pedum: An ichnofossil Trepassey Formation, southeastern Newfoundland, representing Ediacaran-Cambrian boundary in the Canadian Jour. Of Earth Sciences, 40, 1375- 1391. Nagaur Group, the Marwar Supergroup, Rajasthan, Wray, G. A., Levinton, J. S. and Shapira, L. H. 1996. India. Proceedings Indian National Science Academy Molecular evidence for deep Precambrian divergences 78(2), 161-169. among metazoan phyla, Science, v. 274, 568-573. Srivastava, P. 2012-b. Problematic Worms and Priapulid-like Xiao, Shuhai, 2010. Evolutionary patterns of Ediacara Biota, Fossils from the Nagaur Group, the Marwar Supergroup, p 129-133, In Weil and Blumel (eds), McGraw-Hill Year Western Rajasthan, India, Ichnos, 19: 156-164. Book of Science and Technology, The McGraw-Hill Srivastava, P. 2012-d. Ediacaran Discs from the Jodhpur Companies, New York. Sandstone, Marwar Supergroup, India: A biological Xiao, Shuhai, and Laflamme M., 2009. On the eve of animal diversification or taphonomic interplay? International radiation: phylogeny, ecology and evolution of the Jour. Geosciences, 3:1120-1126. Ediacara biota. In Trends Ecol. Evolution, 24(1), 31- 40. Srivastava, P. 2013. Largest Ediacaran discs from the Jodhpur Young, F. G., 1972. Early Cambrian and older trace fossils Sandstone, Marwar Supergroup, India: Their from the Southern Cordillera of Canada. Canadian Jour. palaeobiological significance, Geoscience Frontiers, 30: of Earth Sciences, 9: 1-17. 1-9. Zhang, L., 1986. A discovery and preliminary study of the Srivastava, P. 2015. Giant-sized algal fossils (sea-weeds) late stage of the Late Gaojiashan biota from Sinian in from the Marwar Supergroup, Western Rajasthan, India: Nanjiqiang county, Shaanxi, Bull. Xian. Inst. Geol. A step towards terrestrialization, International Jour. of Mineral Resources, 13: 67- 88 Natural and Applied Sciences, 2(1), 49-74. Zhuralev, a. Yu, 1993. Were Ediacaran vendobionta Steiner, M., and Reitner, J., 2001. Evidence of organic multicellulars?. Neus. Jharb. Geol. Palaeontol. Abh. structure in Ediacara type fossils and associated 190, 299-314. microbial mats. Geology, 29(12), 1119-1122.

How to cite this article:

Purnima Srivastava.2020, Soft–Bodied Ediacaran Metazoans and Ichnofossils from the Marwar Supergroup, Western Rajasthan: Emergence of Earliest Multicellular, Complex Megascopic Animal Life. Int J Recent Sci Res. 11(07), pp. 39177-39191. DOI: http://dx.doi.org/10.24327/ijrsr.2020.1107.5457

*******

39191 | P a g e