Study of Late Triassic Vertebrate Fossils from Tiki Formation, Shahdol District, Madhya Pradesh A Project submitted to Central University of Punjab For the award of Master of Science In Geology By Shabeer Hussain Supervisor Dr. K. Milankumar Sharma Department of Geography and Geology School of Environment and Earth Science Central University of Punjab, Bathinda 2018, May CERTIFICATE I declare that the project entitled "Study of Late Triassic Vertebrate Fossils from Tiki Formation, Shahdol District, Madhya Pradesh" has been prepared by me under the guidance of Dr. K. Milankumar Sharma, Assistant Professor, Department of Geography and Geology, School of Environment and Earth Science, Central University of Punjab. No part of this project has formed the basis for the award of any degree or fellowship previously. Shabeer Hussain Department of Geography and Geology, School of Environment and Earth Science, Central University of Punjab, Bathinda ‐ 151001. Date: ii CERTIFICATE I certify that Shabeer Hussain has prepared his project entitled "Study of Late Triassic Vertebrate Fossils from Tiki Formation, Shahdol District, Madhya Pradesh” for the award of M.Sc. degree of the Central University of Punjab, under my guidance. He has carried out this work at the Department of Geography and Geology, School of Environment and Earth Science, Central University of Punjab. Dr. K. Milankumar Sharma Assistant Professor Department of Geography and Geology School of Environment and Earth Science Central University of Punjab, Bathinda ‐ 151001. Date: iii ABSTRACT Study of Late Triassic Vertebrate Fossils from Tiki Formation Name of student: Shabeer Hussain Registration number: 16mscegs15 Degree for which submitted: Master of Science Name of supervisor: Dr. K. Milankumar Sharma Name of Centre: Department of Geography and Geology Name of school: School of Environment and Earth Science Keywords: Tiki Formation, microvertebrates fossils, Paleo- geography, paleo-environment, Late Triassic period Abstract The objective of the study is to analysis the micro-vertebrate fossil assemblage from Tiki Formation of Late Triassic period and also reveals the significance of paleo-geography and paleo-environment of Tiki Formation, Shahdol District, Madhya Pradesh. Approximately, 1200kg of sample have been collected from Shahdol district, MP. After, washing it properly with water, acetic acid and kerosene, Trinocular Microscope is used for further analysis. Micro-vertebrates which are found in study are Cladodus sp., Actinopterygiian fish, Coelacanthidae fish, Metoposaurian clavicle, Parvodus tikinesis, Archosauriformes indent., Rewacondon tikiensis, Parasuchus hispoli, cynodont, Phytosaur. These faunas are closely resembles with different formation such as Maleri formation in Pranhita-Godavari basin, India, Santa Maria formation of Brazil, Ischiguelasto formation of Argentina, Chinli formation of USA, Dockum fauna of North America with the possibly migration occur during Late Triassic through northern Africa. Thus, it show the global palaeo-biogeographic scenario of Late Triassic time is closely co-relatable Late Triassic taxa in different continents which might have served a relatively rapid intercontinent passage of the vertebrate community extending across the ancient Supercontinent, Pangea. Shabeer Hussain Dr. K. Milankumar Sharma iv ACKNOWLEDGEMENT Every good work starts with the blessings of almighty, so firstly I pay sincere regards to the great God for his blessings. I would like to pay my sincere regards to my parents and family for their love, affection, moral and economical support. Their constant support helped me to complete my M.Sc project work. I would like to thank Dean Academic Affair/COC, Professor V.K.Garg, Central University of Punjab, Bathinda for providing the necessary infrastructure to carry my research work. Further, his valuable suggestions and constant support always encouraged me. I express my deepest feeling of gratitude and appreciation to my Supervisor, Dr. K. Milankumar Sharma, Assistant Professor, Centre for Geography and Geology Central University of Punjab, Bathinda, for his guidance, criticism, encouragement and valuable suggestions which helped me in completing the project work. I would like to thank Dr. Kiran K. Singh (Assistant Professor), Dr. Jitendra Kumar Pattnaik (Assistant Professor) for their help and support in project. Words are inadequate in offering my sincere thanks to my seniors Mr. Nongmaithen Amardas Singh, Mr. Priyananda Singh and my friend Meenakshi Rawat for their valuable support and encouragement. I expand my thanks to Mr. Punit Lab Assistant for their support. SHABEER HUSSAIN v TABLE OF CONTENTS Chapter Contents Page no. No. 1. INTRODUCTION 1-5 1.1 Climate and Rainfall 2 1.2 Vegetation 2 1.3 Drainage Pattern 3 1.4 Physiography 3 1.5 Regional Geology and Stratigraphy 3 2. LITERATURE REVIEW 6-8 3. MATERIALS AND METHODS 9-11 3.1 Field Investigation 9 3.2 Laboratory Investigation 9 3.2.1 Method for separation for microfossils 9 4. RESULT 12-26 5. DISCUSSION 27-29 CONCLUSION 30 REFERENCE 31-35 vi LIST OF FIGURES Figure no. Description Page no. 1.1 Geological Map of Shahdol district 05 and Tiki formation 4.1 Teeth of Cladous sp. 13 4.2 Teeth of Cladous sp. 14 4.3 Teeth of Actinopterygiian fish 16 4.4 Teeth of Coelacanthidae fish 19 4.5 Teeth of Parvodus tikiensis 21 4.6 Teeth of Archosauriformes 23 4.7 Teeth of Cynodont 24 4.8 Tooth of Phytosaur 26 vii LIST OF ABBREVIATIONS S. No Full Form Abbreviation 1. Madhya Pradesh M.P. 2. Geological Survey of India GSI 3. Species sp. 4. Scanning Electron Microscope SCM 5. Hydrochloric acid HCl viii CHAPTER 1 INTRODUCTION In the geological history of Earth, Triassic period is one the most significant time owing to its palaeo-geographic set up in the form of the Supercontinent Pangea, its fossil content especially the composition of tetrapod communities comprising of the terrestrial ecosystems in the past which underwent significant reorganizations of its faunas (Behrensmeyer et al., 1992; Sues et al., 1994; Bandyopadhyay, 1999; Chatterjee and Scotese, 1999; Heckart and Lucas, 2000). This period is also well known for the evolutionary event marked by a dramatic transition from more primitive faunas dominated by “mammal-like reptiles” (synapsids-dicynodonts and cynodonts), rhynchosaurs, temnospondyl amphibians, and primitive archosauromorphs to a more advanced fauna dominated by derived archosaurs (including dinosaurs and crocodiles), mammals, lissamphibians (frogs and salamanders), and lepidosaurs (lizards, sphenodonts, and snakes) (Hackert, 2004). The Tiki Formation exploration in the Late Triassic continental sedimentary rocks exposed in the South Rewa Gondwana basin of Madhya Pradesh was taken up by the Geological Survey of India (GSI) during the early nineties. It represents an important lithostratigraphic as well as a biostratigraphic unit. In view of this, a systematic palaeontological investigation carried out by Das and Datta during the field seasons 1993-94 to 1995-96 and later by Datta in 2000-01 to 2003-04 and Abir Gupta during the field season 2011-13. The Tiki Formation is well known for its vertebrate fossils assemblage of Late Triassic age (Chatterjee, 1978; Chatterjee and Majumdar, 1987; Datta and Das, 1996; Datta et al., 2004). The earlier records of certain well known mega tetrapod fossil remains from the Tiki Formation are the parasuchids like Parasuchus (Chatterjee, 1978), rhynchosaurids like Hyperodapedon (Benton, 1983), rauisuchuid such as Tikisuchus romeri (Chatterjee and Majumdar, 1987) and metoposaurid amphibians Buettneria (Sengupta, 2002), and dipnoan lung fish Ceratodus (Datta and Das, 1996). However, micro-tetrapod assemblages are comprised of an advanced Cynodont Rewaconodon (Datta et al., 2004), a morganucododontid mammal Gondwanadon (Datta and Das, 1996) and Tikitherium (Datta, 2005). Acrodont of Iguanian lizard Tikiguania (Datta and Roy, 2006) is also 1 found. Isolated dental remains of ornithischian dinosaurs like Galtonia sp., Pikinosaurus sp., and the Revoeltosaurus sp. (Das and Datta, 1997, 1999; Datta, 2003) are obtained. The Piscean remains of the Tiki Formation comprise of elasmobrannch selachian fauna Xenacanthus sp., Hybodus sp. and dipnonian fish Ceratodus sp. (Das and Datta, 1997, 1999; Datta and Das 2001, and Datta, 2003; Abir Gupta, 2013). Due to its global significance, a project under the title ‘Study of late triassic vertebrate fossils of Tiki Formation’ has taken into consideration. The faunal correlation with other parts of the world also studied in Tiki Formation exposed in parts of Shahdol district (Survey of India Toposheet no. 64E/5 and 63H/8) of Madhya Pradesh during the field season 2017-18. This study involves the field work as well as the laboratory investigations. Nearly 30 sq. Km area around Tiki village (23°55.345’N; 81°21.439’E), Jora village (23°53.884’N; 81°21.019’E), Simaur village (23°54.707’N; 81°19.802’E), Ghorsa village (23°57.456’N; 81°21.932’E), Tenduar village (23°59.009’N; 81°23.95’E), in Shahdol district. Laboratory investigation involves analysis of mega vertebrate fossils as well as microvertebrate fossils. Later on, palaeo-environmental and palaeo-geographical significance also discussed. 1.1 Climate and Rainfall – Shahdol district has a characteristic climate and rainfall, hot summer, mild winter and experiences south-west monsoon. The climate is humid according to Tharnthwaite due to the presence of forest type vegetation. The average temperature (46.50 to 270) and humidity is maximum in August (exceed 85%) during the South-West monsoon. In April relative humidity is less then 40%. As per the rainfall statistics, the area receives average rainfall of 1130.5mm. normal and mild drought is 25% respectively, where as the severe drought is only 5%. So there is a normal and mild drought in every 7 year and severe drought is ones every 20 year. 1.2 Vegetation – The vegetation is moist tropical deciduous forest which dominates the swamp habitat in the region. The forest floor is usually occupied by green grassed. The flow of the 2 stream is very slow so that meager aquatic vegetation is found i.e. stationary water bodies. the agriculture is not very well developed. The yearly yield products are only enough for their own needs. Mustard oil, groundnut, till, sunflower and soybeans are the main production for oilseeds. 1.3 Drainage Pattern – As the study area fall under the Ganga Basin, the Son river and its tributaries drained from the area. Kunak Nadi and Chowadi Nadi are the two important tributaries of Son river which passes through the area. In North-West direction the other important tributaries like Tipon, Chadas and Bakan flows with a dendritic pattern, drained the district plains. Another most important tributaries of river is Banas river flowing along the eastern boundary of the district, which marks the boundary of Shahdol district with Sindhi District. The Banas river meets with the Son river at the Northern area of Shahdol district. 1.4 Physiography – Shahdol district is located in the north-eastern part of the Madhya Pradesh of India. Physiographically, a major part of the area is an undulating plain with small mounds, dense forest and cultivated lands and badland topography interspersed by small seasonal streams and gullies. In the northern part, a prominent hill range trends ENE- WSW with an average altitude of 400 m. 1.5 Regional Geology and Stratigraphy – The Rewa Basin is dipping towards the North direction and the strike direction of ENE-WSW Chakraborty et al. (2003) considered the Rewa basin as a fault-controlled subsidence basin. The basal part of Rewa basin of Gondwana succession shows stability in the lithological characteristics with other basins of Gondwana. It follows the glacio-fluvial Talchir Formation, and then successively followed by the coal-bearing Barakar Formation, Barren Measures and Raniganj Formation. Based on the lithology and the physical attribute of upper gondwana of Rewa basins is differ from the other 3 basins. The characterised ratio of sand: mud is 4:1 in the basal part of Pali Formation where the pebbly to medium grained micaceous and quartzose sandstones formed the dominant lithotype with subordinate amounts of carbonaceous and red colored mudstone units. Large trough and planar cross beds, and synsedimentary deformation structures such as convolute bedding are common. Study of the Gondwana succession of the Rewa basin was initiated by Hughes (1881). The Upper Gondwana succession was classified into three different stratigraphic horizons, namely the Pali, Parsora and Tiki Formation in succeeding order based on lithology and fossil content (Cotter, 1917; Fox, 1931, 1934). In the studied area, the Middle Triassic Pali Formation is overlain by the Late Triassic Tiki Formation, which is unconformably overlain by Middle Jurassic Parsora Formation. Lithostratigraphically, the Tiki Formation is divisible into Lower Tiki and Upper Tiki. The Lower Tiki Formation is characterized by predominantly reddish mudstone/claystone with calcareous sandstone; the Upper Tiki Formation contains lime-pellet conglomerate intercalated with alternate beds of sandstones and mudstones. The unionid bivalve bed acts as marker bed for the fossiliferous horizons of the Lower Tiki Formation. The Tiki Formation is considered to be deposited in the channel and overbank facies where the river channel was confined within the extensive floodplain, based on the sedimentological characteristics of mud-dominating lithology intercalated with subordinate siltstone and sandstone beds (Mukherjee et al., 2012). However, there is no record of vertebrates fossil from the Pali and Parsora Formation. The objective of the study is to analysis the micro-vertebrate fossil assemblage of Late Triassic period from Tiki Formation. Also, reveals the paleo-environment and paleo-geographic significance of Tiki Formation. Figure 1.1C shows the geological map of my study area. 4 Fig-1.1: The Geological map showing: A. Major Gondwana basins of the peninsular India; B. Position of the study area within the Rewa basin (Source: Mukherjee et al., 2012:); C. Geological Map of the study area, Tiki area in parts of Shahdol district of M.P. India (Source: Datta and Das, 2001; Datta et al., 2004; Mukherjee et al., 2012); D. Lithostratigraphical sequence in Tiki area, south Rewa Gondwana basin, Shahdol, M.P. India, (Source: Datta and Das, 2001; Datta et al., 2004). 5 CHAPTER 2 LITERATURE REVIEW The Vertebrate microfossil collections are a source of faunal abundance material for studying of ancient ecosystems. The Tiki Formation of Gondwana basin in Shodhol district M.P is well-known for Late Triassic macro and micro-vertebrate fossil assemblages. These collections of disarticulated bones and teeth belonging to a diverse group of vertebrate taxa. These are the source of information on ecologically and phylogenetically significant taxa with body size less than about 5 kg (Fisher, 1981). There is a distinct angle towards collection, exploration and study of large vertebrate fossil, that are relatively easy to excavate something conformation source of information (Bandyopadhyay, 1999, 2011; Brusatte et al., 2010; Novas et al., 2011; Nesbitt et al., 2013). In the past many researchers have studied vertebrate microfossils (e.g., Heckert, 2004) which may be due to the difficulty in recognizing their fossils with naked eyes, along with laborious and time-consuming procedures of extraction. The Permo-Triassic deposits of India are rich storehouses of mega- vertebrate fossils collection of different Gondwana basins of (Ray, 2000; Bandyopadhyay, 2011), which are mostly collected from the ubiquitous red mud-rock dominated fluvial sediments. Sedimentary rocks such as limestone, sandstone, mudstone which are deposited in ancient lakes, rivers, seas and desserts are the well preserver of Vertebrate micro-fossils (Behrensmeyer et al., 1992; Benton, 2005). Vertebrate micro-fossils were commonly found in channel bars and logs (Heckert, 2004), the carbonate nodules of pedogenic and ash falls (Cifelli et al., 1996; Schiebout et al., 1998) also act as possible sources for the preservation of vertebrate microfossils. Chatterjee 1967, reported that complete skeleton of phytosaur had been collected from Maleri Formation and another specimen from Tiki Formation of the Son- Mahanadi valley. They referred specimens to Parasuchus hislopi but they followed by Ballew, (1989) and called them Paleorhinus hislopi. They concluded that Paleorhinus was the least derived phytosaur (Ballew 1989) because it had external nares anterior to fenestrae and a posterior temporal arcade at the level of skull roof so they 6 considered the fossil may be young. The mud-dominated succession of Tiki Formation had yielded mega-vertebrates. These were followed by metoposaurid temnospondyl Buettneria (Sengupta, 2002), a phytosaurid Parasuchus (Chatterjee, 1978), a rauisuchid Tikisuchus (Chatterjee and Mazumdar, 1987), and a rhynchosaurid Hyperodapedon (Benton, 1983). In addition, micro-vertebrates, such as a dromatheriid non-mammalian cynodont, Rewaconodon (Datta et al., 2004), a morganucodontid mammal Gondwanodon (Datta and Das, 1996), and the earliest mammal with transversely expanded upper molar Tikitherium (Datta, 2005) were also known from the Tiki Formation. Based on its faunal content, the Tiki Formation had been correlated with the lower member of the Maleri Formation of the Pranhita- Godavari basin in southern India (Lucas, 1998; Datta, 2004) and the Camp Springs member of the Dockum Formation, USA (Datta et al., 2004), and was assigned a Late Triassic (Carnian) age (Benton, 1994; Datta, 2004; Datta et al., 2004). The earlier record of well-known fossil remains of small tetrapod assemblage are comprised of an advanced cynodont Rewaconodon (Datta et al., 2004), an unnamed rank mammal Tikitherium (Datta, 2005) and an acrodont Iguanian lizard Tikiguania (Datta and Ray, 2006). Isolated dental remains of orinitheischian dinosaurs like Galtonia species Pikinosaurus species and the Revueltosaurus species are also reported (Das and Datta, 1997, 1999; Datta, 2003). The Piscean remains of the Tiki Formation belong to elasmobranch selachian fauna Xenacanthus species Hybodus species and dipnonian fish Ceratodus species (Das and Datta, 1997, 1999; Datta and Das 2001, and Datta, 2003; Gupta, 2013). In late Triassic clade of extinct archosaurs which are predominantly found in marine environment had characteristics similar to a modern crocodile. Sharma and Kumar, 2015 found phytosaurid osteoderm remain from Tiki Formation. They concluded that osteoderms were found together with phytosaurian teeth, jaws and vertebrae. Mukherjee and Ray, 2012 discovered graveyard of rhynchosaur, an archosauromorph reptile in the upper Triassic Tiki Formation of South Rewa Gondwana basin. They interpreted mass mortality event during sudden flooding. They continued on the same site identified rhynchosaur bones as belonging to a new species Hyperodapedon 7 tikiensis that represent the most advanced species among rhynchosaurs (Mukherjee and Ray, 2014). Bandyopadhyay, 1999 and Chatterjee, 1974 reported that rhynchosaurs are found in the middle Triassic Denwa Formation of the Satpura basin and in the middle Triassic Yerrapalli Formation and the Upper Triassic Maleri Formation of the Pranhita– Godavari (PG) basin. Based on its faunal content, Lucas, 1998 and Dutta, 2004 correlates the Tiki Formation with the lower part of the Maleri Formation of the Pranhita–Godavari basin. The Camp Springs Member of the Dockum Group, USA and Benton, 1994 correlates Tiki Formation with Upper Triassic (Carnian) age. Rogers, 1990 found that Tiki Formation was susceptible to drought, so close association of the rhynchosaur bone-bed is suggestive of drought-related mortality, similar to that seen in the dinosaur bone-beds of Cretaceous Two Medicine Formation, Montana and Fiorillo et al., (2000) also seen that association in Placerias Quarry of the Upper Triassic Chinle Formation. Late Triassic freshwater fishes show resemblences to that found in the Chinle Group of USA (Heckert, 2004). The Tiki Formation is correlated with the lower part of the Maleri Formation of the Pranhita-Godavari Basin, the Ischigualasto Formation of Argentina, and the Hyperodapedon assemblage zone of the upper part of the Santa Maria Formation of Brazil (Ray, 2015). Rewaconodon tikiensis is rat like cynodont related to the early ancestors of mammals is similar to Microconodon from the Newark Supergroup of North America reported by Datta et al., (2004). Tikisuchus is similar to Rauisuchus of Santa Maria Formation of Brazil, Postosuchus of the Tecovas Formation of North America and Teratosauras of Stubensandstein of Germany as revealed by Chatterjee and Majumdar, 1987. 8 CHAPTER 3 MATERIALS AND METHODS The methodology involved in the present study can be grouped into two parts including the field work and laboratory work. 3.1 FIELD INVESTIGATION Detailed geological field work and mapping of the area will be carried out selecting the suitable section for paleontological and studies. Through fieldwork will also be done at the previously well-known fossil sites and its adjoining lithostratigraphic units. This survey will create the spatial-temporal context of fossil-bearing localities (including the collection of GPS coordinates for all localities) taking into consideration with the exposures of most complete and less tectonically undeformed section for documenting fossils, a collection of samples for stable isotopic and sedimentary geochemical study. For the palaeontology bulk sampling from each horizon will be carried out during the section measurement. Approximately, 1200kg of the sample will be taken for the analysis of micro-vertebrate including pisces, micro-mammal, reptile etc. besides the in-situ collection of mega- fossils from the field. The location and horizon from the collected samples will be properly recorded in the dairy and field photograph will be taken. Each collected sample will be properly labeled and each fauna collected will be properly catalogued with (PBL/CUP/MKH-T) which will be store in palaeontology and biostratigraphy lab of Department of Geography and Geology. For clay mineralogical analysis, sampling will be done from the measured section taking into consideration with the lithology exposed in the section with approximately 6-6 cm interval from bed by bed. At least 1kg of the sample from each litho-unit will be collected and proper procedures for the preparation and analysis of the samples in the laboratory will be followed. 3.2 LABORATORY INVESTIGATION 3.2.1 Method for separation of microfossils. A. Maceration 9 i. Maceration with water: In this process, the bulk sample brought from field investigation will be first soaked in water for 24 hours and then wet sieving in the three different messes unit and drying the collected sample from the wet sieving. ii. Maceration with water and acetic acid: In this method, the dried samples will first be treated with acetic acid and then water is poured into it thereafter. This type of maceration will be undertaken on harder samples such as hard clays, sandstone, shale, etc. The strength of the acid and the time of dissolving the sample depends on the type of the samples. iii. Maceration with kerosene and water: In this process, the sample will purely dry at first before putting it in kerosene for 3-4 hours. If the sample is not dried well then the kerosene will not enter into the sample. After the sample is fully soaked with kerosene, kerosene is drained out of the sample which is followed by pouring water into the tubs containing the samples. Due to the pressure created by the oil coming out of the interstice and the water forcing into it, the sample starts disintegrating by itself. This type of maceration technique is applied to samples such as hard mudstone. The loose macerates of the above samples will be further subjected to wet sieving process in which the macerates will be passed through the different size of sieves. The residues collected from the sieves were made dry at first and then they were seen under the binocular microscope for picking the fossils contained on it. The important fossils will be separated by picking up from the macerated through the Leica APO binocular microscope. B. Scanning Electron Microscope (SEM) Proper preparation for microfossil samples will be carried out before SEM-EDX analysis. Firstly, the collected fossil sample is clean-up by using ultrasonic bath by 5- 10 minute. After cleaning, the sample is put on the SEM plate by using double side tap and clean again with acetone. Thereafter, the fossil will be subjected to gold coating. For microstructure analysis of mammalian, enamel and dentine, the cleaned samples will be the first cut both in the longitudinal and transverse section and the surface will be polished. Etching of the polished surface will be done using 5% HCL 10 before gold coating. SEM-EDX analysis of the gold coated samples will be done including proper photography at the Carl Zeiss Merlin Compact 6073 Scanning Electron Microscope housed at Central Instrumentation Laboratory, Central University of Punjab, Bathinda. C. Photography under Trinocular Stereozoom Microscope and taxonomic study: For further study of the morphology of the fossil collected will be photographed and studied under Leica M205C Trinocular Microscope. The mega-vertebrates fossils which are collected from the field will be thoroughly washed with water and air dried. Then the dried fossil sample will be photographed using a digital camera (NIKON D7200) and proper mould and cast of the sample will be prepared for precious samples before carrying out any further study such as the thin slide study, SEM study for microstructural analysis, etc. Proper taxonomic identifications of the fossils will be done by comparing the morphology and structure of the fossil with their holotypes, syntypes, etc. and also by comparing with the identical fossils reported in the certain literature by the previous workers. Biostratigraphic study of samples collected from the measured section will be done taken into consideration of the followings points:  Quantifications of assemblage zones;  Quantification of index fossils concept.  Calculation of sequence of events. 11 CHAPTER 4 RESULT Order: Ctenacanthiformes Family: Ctenacanthidae Genus: Cladodus (Egerton, 1841) Cladodus sp. (Fig-4.1: A-E) (Fig-4.2: A-H) Locality and horizon: Nala section near Tiki, Simaur and Jora village sections; red mudstone bed above the unio bed. Material: Teeth of a Cladodus species sample number (PBL/CUP/MKS-T/01) and (PBL/CUP/MKS-T/02) Remarks: The present specimens have conical in shape with, concave cusp. The specimen has triangular root or base in which the two cusps are preserved (fig-4.1: A,A’) in some well-preserved specimens there is three cusps one is in the middle which is small as compare to the other two cusps (fig-4.2: B,B’). Striation is developed in the whole crown but the maximum specimens striations are present only in upper part of the cusp (fig-4.1: A,C,E) and (fig-4.2: C,D,E,F,G) while in few, these are continued in whole cusps. (fig-4.1: B,C) In most of the specimens have only one cusp preserved (fig- 4.1: A-E) (fig-4.2: C-H) but in few specimens both median and lateral cusps are present (fig-4.2: A,B) but lateral cusps are broken in some specimens while maximum specimen have cusp well preserved. Roots are well preserved only in the few specimens; (fig-4.1: E) (fig-4.2: A, B, D, G) presents. The above characters are similar to Cladodus shark teeth. These are earlier reported literature of Duffin J. Christopher and Ginter, M. (1843) and Ginter, et al., (2005) and. The above specimens are fragmented so there is a problem in the classification up to species level. 12 Fig-4.1: Teeth of “Cladodus sp” (PBL/CUP/MKH-T/01). A, B, C, D, F) In labial and lingual view E) Labial view and E’) In sterio-occlusal view. All (A-F are in 1mm of scale). 13 Fig-4.2: Teeth of a Cladodus sp. (PBL/CUP/MKS-T/02) A) Labial view. B) In Basal view and A’ and B’ in distal. C, E, H) In Labial-lingual view. D) In occlusal view D’) In oblique-liable view G) In labial view and G’) In distal view. 14 Subclass: Actinopterygii (Klein, 1885) Actinopterygiian indet. (Fig-4.3: A-H) Locality and horizon: Nala section near Tiki, Jora and Sirmaur village section; red Mudstone bed above the unio bed Material: Teeth of Actinopterygiian fish. (PBL/CUP/MKH-T/03) Remarks: Complete or near-complete teeth as pertaining to actinopterygian (bony) fishes are also picked up from the macerated samples of Jora and Tiki village. These teeth can be distinguished into three distinct morphotypes and the remaining teeth are miscellaneous and probably may represent into genera. Some of these teeth are well preserved as evidenced by the presence of well preserve crystal tips, however, some specimen have broken tip. The root portion is not preserved well which are ken to break off. Fish morphotype 1: This type of tooth is conical, bent/curved at the middle of the crown, with a thicker base and the apex is tapering (fig-4.3: A-F). The tip is not directly above the centre of the base, but more to one side. This group comprises certain teeth which are mostly conical, with a broader base and significantly smaller towards the tip, thickness, length and degree of curvature each specimen may vary. It is difficult to identify the specimen into genera and species. Fish morphotype 2: Like the tooth of morphotype 1, this type of tooth is also conical, slightly curved to straight, the crown with pointed apex; it differs from the first one in having an abruptly tapering crystal tip at the upper portion of the crown (fig-4.3: A,A’, B,B’,E,E’,F,F’) but in (fig-4.3: C,C’) the upper tapering crystal tip is broken off. The base of the teeth are broader, roots are not preserved. 15 Fig-4.3: Specimen number (PBL/CUP/MKH-T/03) A-F) Teeth of Actinopterygiian fish in labial and lingual view. (All are in 1mm of scale) 16 Class: Sarcopterygii (Romer, 1955) Order: Coelacanthiformes (Berg, 1937) Family: Coelacanthidae (Agassiz, 1843) Coelacanthidae indet. (Fig-4.4: A-C) Locality and horizon: Tiki, Jora, Semour, village section; red mudstone/clay bed just below and above the unio bed. Material: Incomplete fragments of tooth plates of Coelacanthidae fish (PBL/CUP/MKH-T/04) Remarks: Several tooth plates oriented in a regular fashion along the plate (fig-4.4: A,A’, C, C’). These teeth small size sometimes forms a tooth row, several teeth appearing as an elongate block. Wear facets can be seen on some tooth surfaces as the striations have been worn down. The present tooth plate are having similar looks with those of Coelacanthidae tooth plates from Triassic Snyder Quarry North-Central New Mexico (Heckert and Jenkin, 2005) and also Chinle group have previously been referred to as Colobodontidae or Perleididae, extinct groups of actinopterygians similar to the palaeoniscids (Murry, 1982; Huber et al., 1993; Heckert, 2001). This report is first of this kind from the Late Triassic deposit of India. Pectoral Girdle of Metoposaur sp. Indet (Fig-4.4: B, B’ and D, D’) Locality and horizon: Nala section from Tiki, Jora, Semour, village; red mudstone/clay bed below the unio bed. Material: Incomplete fragments of a metoposaurid clavicle. (PBL/CUP/MKH-T/04) Remarks: The specimens are incomplete fragments of pectoral girdle of Metaposaurian origin. As the specimen is very small fragments identification of the specimen and its position in the pectoral girdle as a part of clavicle and interclavicles portions is quite difficult. The specimens are highly sculptured, ornamented with a very small area of rounded or horizontal cavities or depression. In fig, the specimen has small holes like porous structure (fig-4.4: B and D). 17 Subclass: Actinopterygii (Klein, 1885) Actinopterygiian indet. Specimen number. (PBL/CUP/MKH-T/04) Actinopterygian Fish scale: The majorities of the collected fish scales are rhomboid in shape, smooth external surface, and have a slightly concave external surface at the middle with convex leading edge that articulates with a shallow concavity in the next most anterior scale (fig-4.4: E, F). Some of the present specimen show well-preserved scale (fig-4.4: E, F) with their original iridescence of the enameloid are still present. The specimens represent an array of morphotypes ranging from simply smooth rhombodhedral shape to nearly robust external ornamentation (fig-4.4: E, F). The majority of the osteichthyan scales, teeth and bone fragments from Tiki Formation, even though they seem quite diverse, are difficult to identify below to genus and species level. 18 Fig-4.4: Tooth plates of Coelacanthidae fish. (PBL/CUP/MKH-T/04). A) Labial view and A’) Occlusal view. C, C’). In Stereo-occlusal view. (PBL/CUP/MKH-T/04) B, D) In labio-occlusal view; Actinopterigiian fish scale (PBL/CUP/MKH-T/04) E and F in external view. (All figs are in 1mm scale) 19 Order: Euselachia (Hay, 1902) Superfamily: Hybodontoidae (Owen, 1846) Family: Lonchididae (Herman, 1977) Genus: Parvodus (Rees and Wonderwood, 2002) Parvodus tikiensis Prasad et al., 2004. (Fig-4.5: A-E) Locality and horizon: Nala section near Jora village section; red mudstone bed below the unio bed. Material: Teeth of a fresh-water shark. (PBL/CUP/MKH-T/05) Remarks: The present specimens are similar with the Parvodus tikiensis morpho- type 2 of Prasad et al., 2014. Earlier the first report of Parvodus tikiensis has been made by Prasad et al., 2004. The species is signified by the presence of labio- lingually compressed tooth bearing two pairs of lateral cups let on either side of the main cusp. The main cusp is the highest of all which followed by the first lateral cusp which nearly half of the height of the principal cusp (fig-4.5: A, A’, B, B’ and C, C’and D, D’). The root of the teeth has elongated depression in ventral view. The principal cusp is high, diamond shaped. Both the labial and lingual surface of the tooth absent in vertical striation. The occlusal crest is sharp, except fig-4.5: E crenulated developed strongly and lingo-laterally curved. The specimen E, E’ has an arc shape in labial view has a smooth base, the crown is not well developed as compared to other specimens. 20 Fig-4.5: Parvodus tikiensis (PBL/CUP/MKH-T/05) All fig-4.5: A-E) In lingual and labio- occlusal view. (Scale of 1mm) 21 Archosauriformes (Gauthier, 1984) Archosauriformes indet. (Fig-4.6: A-F) Locality and horizon: Nala section from Tiki, Jora, Semour, village; red mudstone/clay bed below the unio bed. Material: Teeth of “Archosauriformes” (PBL/CUP/MKH-T/06) Remark: Indeterminate teeth that probably pertain to Archosauriformes occur as diverse morphotypes at the tiki locality. This is a somewhat more refined listing than “Reptiliaindet.” and follows the criteria of Godefroit and Cuny (1997) to recognize a tooth as “archosauriform.” The given morphotype below describes in detail. (Archosauriformes) There are about 700 reptilian teeth preserved, of which nearly 300 teeth have slightly recurved, subtriangular crowns with expanded bases, asymmetrical in basal view and distinct denticles both on the posterior or anterior carinae (Fig-4.6: A–F). Morphotype A In (fig.6: A,A’ and C,C’) is a moderately low, conical, slightly laterally compressed tooth bearing fine anterior and posterior serrations. This tooth is too tall to be a phytosaur type B tooth (Hunt, 1994). Most of the specimen crown part is preserved slightly broken and represent the shedded teeth. Roots are not preserved in any specimen. This morphology suggests that the teeth are similarly belonging to archosauriforms and are similar to Revueltosaurus, tecovasaurus, and other (Irmis et al., 2007; Heckert, 2004; Godefroit and Cuny, 1997). The (fig.6: E and F) is a moderately conical tooth fragment. Only the posterior margin is serrated. The tooth is laterally compressed and was probably moderately tall to tall when complete. 22 Fig-4.6: Teeth of Archosauriformes (PBL/CUP/MKH-T/06) A) Labial view A’) Lingual view B, B’) In labial and lingual view C) Labial view C’) In stereo-labial view E) In lingual E’) In lateral view F) Labial and F’) In lingual view. (All are in scale 1mm) 23 Class: Cynodontia (Owen, 1861) Family: Dromatheriidae (Gill, 1872) Genus: Rewaconodon Datta et al., 2004 Rewaconodon tikiensis Datta et al., 2004. (Fig-4.7: A-B) Locality and horizon: Nala section near Tiki, Jora and Semour village extension section. Red clay bed above the unio bed Material: Teeth of a cynodont. (PBL/CUP/MKH-T/07) Remark: The present specimens are identical with the Rewaconodon tikiensis teeth described by Datta et al. (2004). In (fig-4.7: A- B) are bicuspid teeth post canines which closely resemble the micro teeth of small mammal-like a reptile. In the given (fig-4.7: B) both root and cusp and are preserved characterized by bicuspid. In these specimens, the two crown is developed in conical or diamond shape from the base in lateral view. In fig-4.7: A (PBL/CUP/MKH-T/07) there is also two cusps (bicuspid) and root is not preserved. In all specimens, the cusps are pointed. In both, the specimen the medial cusps are tall as compare to lateral cusps. The above characters indicate these are an advanced non-mammalian cynodont, possibly from the Dromotheriidae family. Fig-4.7: Teeth of a cynodont (PBL/CUP/MKH-T/07) A) In Labial B) In Lingual view. C) In lateral view. (Scale 500μm) 24 Order: Phytosauria (von Meyer, 1861) Family: Phytosauridae (Jaeger, 1828) Genus: Parasuchus (Lydekker, 1885) Parasuchus hislopi (Lynekker, 1885) (Fig-4.8: A-C) Locality and horizon: Nala section near Jora village; red clay bed below the unio bed Material: the Isolated tooth of Phytosaur (PBL/CUP/MKH-T/08) Remarks: fig-4.8: (PBL/CUP/MKH-T/08) morphologically the specimen is well comparable with the phytosaurian reptile Parasuchus hislopi describe from the Tiki Formation (Chatterjee et al., 1974). Of the three phytosaurs teeth, one (fig-4.8: B) is almost complete and the remaining two are incomplete (fig-4.8: A and C) with broken apex. There is a depression in the broken apex (fig-4.8: A). The teeth are conical and cylindrical and little-curved crown. The striation is well preserved and they bear fine longitudinally on the whole crown part. The roots are not preserved. In (fig-4.8: C) there is a linear depression through the middle of teeth in ventral view. It is considered osteoderms are commonly found in many groups of extant and extinct reptiles and amphibians including phytosaurs, crocodylomorphs, various groups of dinosaurs, aetosaurs, etc (Rogers et al., 2011) and are widespread amongst the phytosaur and aetosaur (Romer, 1956; Scheyer, et al., 2013). Earlier, Chatterjee (1978) reported osteoderms, certain bones and a well-preserved skull of a basal phytosaur, Parasuchus hislopi, from the coeval deposits of Late Triassic deposits of Tiki Formation. 25 Fig-4.8: Isolated tooth of Phytosaur (PBL/CUP/MKH-T/08) specimens A-C) In labial and lingual view (Scale of 1mm) 26 CHAPTER 5 DISCUSSION During the Late Triassic period continents were grouped together as a supercontinent Pangaea and these certain continental processes were probably very active. Hence, the terrestrial or non-marine Triassic deposits are extensively found in India, South Africa, South America, Western and Eastern North America, Antarctica, Australia, USSR, Western Europe, etc. The presence of closely related Late Triassic taxa in different continents may be one of the major attribute of a relatively rapid inter- continent passage which in turn is the result of manifestation of the integral part of the vertebrate community extending across the ancient supercontinent, Pangea (Bandhopadhyay, 1999). In Indian Gondwana deposit, the faunal remains of Tiki Formation are closely resembles with it equivalent deposits of Maleri Formation in Pranhita-Godavari basin. Tiki Formation has been correlated with the lower member of the Maleri Formation of Prahnita-Godavari basin, Santa Maria Formation of Brazil, Ischiguelasto Formation of Argentina, Chinli Formation of USA, Dockum fauna of North America with the possible migration occur during Late Triassic in northern Africa. The records of Late Triassic Dromatherid Cynodont, Rewaconodon tikeinsis (Datta et al., 2004) broaden the perspective in the study of paleo-environment, evolutionary aspects of cynodont reptile and their palaeo-geographic correlations with the similar faunas of the rest of the Late Triassic Gondwana deposits of Pangaea. Datta et al., 2004 described the extend of palaeobiogeographic distribution of Dromatherid described earlier from the Carnian sediments of the Newark Supergroup and Dockum Group of North America (Sues et al., 1994; Lucas and Oakes, 1988), Norian- Rhaetian sediments of Europe (Hahn et al., 1994; Godeifroit and Battail 1997) and closely correlatable therioherpetid from the Carnian stage of Santa Maria Formation of Brazil (Hahn et al., 1987; Battail, 1991; Bornaparte and Barberena, 1975; Abdala and Ribeiro, 2000). In India, records of fossil osteoderms are very poor. Earlier, Parasuchus hislopi, a phytosaurian reptile associated with dermal armour was recorded from the Late Triassic Maleri Formation of Pranhita-Godavari Valley (Chatterjee, 1978). Milankumar 2 7 and Jitendra Kumar (2015) reported osteoderms of phytosaurian remains from Tiki Formation. However, osteoderms of archosaurian reptile are known from certain upper Triassic strata of all the continents except Antarctica and Australia (Heckart and Lucas, 2000). The occurrence of phytosaurian remains from the Late Triassic Gondwana deposits have been reported from western United State (Lower Dockum Group, Camp Spring member of the Trecovas Formation, Lower Chinle Formation, Popo Agie Formation), Germany (Blasensandstein), Morocco (Argana Formation), Austria (Opponitzer Beds), Scotland (Lossiemouth Sandstone Formation), South America (Santa Maria Formation and Ischigualasto Formation) (Hunt and Lucas, 1991). The piscean remains of Xenacanthus/Cladodus sp., Hybodus sp. and dipnonian fish Ceratodus sp. show close affinities with those of Santa Maria Formation of Brazil, Ischiguelasto Formation of Argentina, Keuper deposits of German, Late Triassic of Morocco, Late Triassic of Madagascar, etc. Triassic hybodont sharks have been reported earlier from North America, South America and Europe. The late Triassic hybodont sharks from the Gondwana of India (Tiki and Maleri Formations) comprises of five species viz. Lonchidion estesi, L. incumbens, Lissodus duffini, Parvodus tikeinsis and Polyacrodus (Prasad et al., 2008). Coelacanthidae tooth plates is first of this kind from Late Triassic deposit in India is similar to Triassic Snyder Quary North-Central New Mexico (Heckert and Jenkin, 2005) and also Chinle group have previously been referred to as Colobodontidae or Perleididae. Extinct groups of actinopterygians similar to the palaeoniscids (Murry, 1982; Huber et al., 1993; Heckert, 2001). Based on the sedimentological characteristics, Tiki Formation predominantly comprising of mud dominating lithology intercalated with subordinate siltstone and sandstone beds is considered to be deposited in the channel and overbank facies where the river channel was confined within the extensive floodplain (Mukherjee et al., 2012). Majority of the fossil fauna reported here were collected from the reddish mudstone beds which are exposed just above and below the Unionid bivalve beds which is considered as marker for the fossiliferous horizons of Lower Tiki Formation indicating the presence of either swamps in this extensive floodplain deposits or 28 lacustrine type of environment (Milankumar and Jitendra, 2015). The palaeontological and sedimentological datas from the study area suggest the prevailing of semiarid climate with the seasonal rainfall of monsoon type (Chatterjee, 1978; Mukherjee et al., 2012). The faunal records comprising of mullusc, fresh water selachian, rhynchosaur, metoposaurid, rauisuchid, phytosaur and cynodonts, Dinosuarid (Chatterjee and Majumdar, 1987; Das and Datta, 1997, 1999; Datta, 2003; Datta et al., 2004) suggest the presence of higher trophic which were directly or indirectly dependent on each other. 29 CONCLUSION The present palaeontological investigation from report of tooth remains of Coelacanthidae indet. Cladodus sp. from Tiki Formation is one of a kind from the Late Triassic deposit of India. The faunal records of the Tiki Formation comprising of phytosaur osteoderms, teeth, vertebrate, jaw associated with other fossil remains of fresh water metoposaurid, cynodonts, mammals and other microvertebrate assemblages of microvertebrates assemblages fossil fishes remains including hybodonts, Ceratodus, Actinopterigiians, etc. strongly suggest the presence of higher level ecosystem which was directly or indirectly dependent on each other. Both the sedimentological and fauna data from the Tiki Formation especially the lower parts suggested an extensive floodplain to lacustrine environment of deposits. The current faunal remains from the Late Triassic Tiki Formation conforms to the earlier evidences witnessing a more biologically diverse series of origin and diversification events in the Triassic period including very early or first appearances of mammals, dinosaurs, lepidosaurs, pterosaurs, turtles, crocodiles, and lissamphibians, etc. The mega and microvertebrate remains of the Late Triassic of India also show close biogeography affinities with those of similar deposits of Gondwanaland and Laurasia comprising of Santa Maria Formation of Brazil, Ischiguelasto Formation of Argentina, Chinli Formation of USA, Dockum fauna of North America; Late Triassic continental assemblage of German Keuper; Late Triassic of Morocco, Late Triassic of Madagascar; Carnian sediments of the Newark Supergroup; Camp Spring member of the Trecovas Formation, Lower Chinle Formation, Popo Agie Formation of United State; Norian- Rhaetian sediments of Europe; Morocco (Argana Formation), Austria (Opponitzer Beds), Scotland (Lossiemouth Sandstone Formation) etc. Thus, the global palaeo-geographic scenario of Late Triassic is rapidly correlated with Late Triassic taxa in different continents which might served a relatively promptly inter- continent passage of the vertebrate community extending across the ancient Supercontinent, Pangea. 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