Journal of African Earth Sciences, Vol. 29, No. 1, pp.233-241, 1999 g 1999 Elsevier Science Ltd Pergamon Pll:SO899-5382(99)00093-7 All rights reserved. Printed in Great Britain 0899.5362/99 $- see front matter

Middle Triassic vertebrates of India

SASWATI BANDYOPADHYAY * and DHURJATI P. SENGUPTA Geological Studies Unit, Indian Statistical Institute, 203 BT Road, Calcutta 700 035, India

ABSTRACT-Until recently, the Yerrapalli Formation of the Pranhita-Godavari Valley, Deccan, India, was considered as the only Middle Triassic vertebrate-bearing horizon of. India. A new fauna1 assemblage comprising dipnoans, capitosaurids, brachyopids, trematosaurids, dicynodonts, rhynchosaurs and archosaurs has recently been recovered from the Denwa Formation of the Satpura Basin, central India. This fauna1 assemblage gives ample indications to suggest another Middle Triassic vertebrate-bearing horizon from India. The fauna1 elements of the Denwa Formation compare very well with those from the Moenkopi Formation of North America, and appear slightly older than the Yerrapalli fauna. The Bhimaram Formation, occurring immediately above the Yerrapalli and below the Carnian Maleri Formation, has also produced some capitosaurid and dicynodont fragments. An analysis of these three vertebrate-bearing horizons from India reveals that, while the Denwa and Yerrapalli Formations can be dated as Early and Middle Anisian, respectively, the Bhimaram Formation may be anywhere between Late Anisian and Ladinian. o 1999 Elsevier Science Limited. All rights reserved.

RESUME-Jusqu’il y a peu, la Formation Yerrapalli de la vallee de Pranhita-Godavari (Deccan, Inde) dtait considerbe comme le seul niveau $Ivertebres du Trias moyen. Une nouvelle association faunistique comprenant des dipneustes, des capitosaurides, des brachyopides, des tremato- saurides, des dicynidontes, des rhynchosaures et des archosaures a et& recemment decouverte dans la Formation de Denwa, du bassin de Saptura, en lnde centrale. Cette association faunis- tique permet de suggerer I’existence d’un nouvel horizon a vertebres du Trias moyen en Inde. Les elements faunistiques de la Formation de Denwa sont tres proches de ceux de la Formation de Moenkopi d’Amerique du Nord et apparait legerement plus vieille que la faune de Yerrapalli. La Formation de Bhimaram, qui se situe juste au-dessus de la Formation de Yerrapalli et sous la Formation de Maleri d’dge Carnien, a fournit Bgalement quelques fragments de capitosaures et de dicynodontes. Une analyse de ces trois niveaux a vertebres d’lnde revele que les Formations de Denwa et de Yerrapalli peuvent etre datees respectivement de I’Anisien inferieur et moyen, tandis que la Formation de Bhimaram se situerait entre I’Anisien superieur et le Ladinien. Q 1999 Elsevier Science Limited. All rights reserved.

(Received l/7/98: revised version received 20/l O/98: accepted 16/l l/98)

INTRODUCTION In India, Middle Triassic vertebrates were first immediately overlying the Yerrapalli and underlying reported by Jain et a/. (1964) from the Yerrapalli the definite Late Triassic vertebrate-bearing Maleri Formation of the Pranhita-Godavari Valley, Deccan Formation (Kutty and Sengupta, 19891, has yielded (Fig. 1 I. Anderson and Cruickshank (1978) assigned a few fragmentary vertebrate bones. Some fragmen- an Anisian age to the fauna. Apart from the Yerrapalli tary bones are also known from the Middle Triassic Formation, there were sporadic reports of fossil Denwa Formation of the Satpura Gondwana Basin vertebrate fragments of possible Middle Triassic age of central India, situated about 400 km north of the from other places in India. The Bhimaram Formation, Pranhita-Godavari Valley (Fig. 1). These reports never

* Corresponding author

Journal of African Earth Sciences 233 S. BANDYOPADHYA Y and D. P. SENGUPTA

I I 1 780 820 86" ,yt.J IOOkm I ‘1. *i-t:> (,X.~ 24" Rewa Basr \

SatPufa Bas’n

‘:::::I;:...... __ .;.. : dii. . . ..__ CALCUTTA . yJ NAGPUR ??

Figure 1. The Gondwana basins of India. received proper attention and the Yerrapalli Formation and occasionally in the sandstone. Two new species was considered as the only definite contender for of Parotosuchus (Mukherjee and Sengupta, 19981, the Indian Middle Triassic terrestrial vertebrate- a lonchorhynchine trematosaurid (Welles, 19931, and bearing horizon. some dicynodonts (Bandyopadhyay, 1988) are known Recently, several well-preserved specimens have from the Denwa Formation. Recently, identifiable been collected from the Denwa Formation by wor- fragments of a dipnoan, Ceratodus, a brachyopid, a kers of the Indian Statistical Institute. The new fossil large- and a medium-sized dicynodont, a rhynchosaur assemblage from the Denwa has only a broad simi- and a small archosaur have been recovered from larity with that of the Yerrapalli. Some specific ele- this horizon. ments, however, suggest that a part of the Denwa Capitosaurids are the dominant temnospondyl of assemblage could be older than that of the Yerrapalli. the Denwa fauna. Hughes, in 1877, recovered a The present paper gives an account of the new fauna1 bone fragment which was described as a crocodilian elements and attempts to correlate the Denwa fauna1 scute by Lydekker (1885a). Later, this bone was assemblage with other similar faunas of the world. identified as part of a right clavicle of Mastodono- saurusindicus by Lydekker (1885b, 1890). Chatter- jee and RoyChowdhury (19741, however, doubted FAUNA FROM THE DENWA FORMATION the identification as well as the taxonomic assign- The Denwa Formation of the Satpura Basin, central ment of the specimen. They described it as a left India, conformably overlies the Pachmarhi Formation tabular and assigned it to Parotosuchus. Recently, (Table 1) of probable Permo-Triassic age. It is domin- Mukherjee and Sengupta (1998) reported two spe- ated by red mudstone which is silty in places; cies of Parotosuchus based on several well-preserved variegated coloured mudstone interbedded with cal- skull specimens. careous sandstone and palasosols are also present. The two Parotosuchus species differ from all other The vertebrate remains occur mainly in the mudstone parotosuchids by the presence of a large anterior

234 Journal of African Earth Sciences Middle Triassic vertebrates of India

Table 1. The Upper Gondwana stratigraphy of the Satpura Basin (modified after Crookshank, 19361

Formations Brief Litholoav Lameta Conglomerate, limestone and mudstone ]

Massive white sandstone with Jabalpur conglomerate; white and red mudstone; Ptiloph yllum flora Lower carbonaceous shale and coal lenses Cretaceous

Predominant coarse conglomerate, bands of calcareous sandstone, variegated dinosaurs ? mudstone and calcareous marly lenses Unio, Ceratodus, ti capitosaurids, Predominant red mudstone, variegated brachyopid, Denwa mudstone interbedded with sandstone; trematosaurid, Middle pebbly at places; peloidal calcirudite/ dicynodonts, Triassic calcarenite rhynchosaur, archosaur White coarse-grained sandstone with Pachmarhi lenses of angular to subangular quartz temnospondyl Permo- pebbles, green shales Triassic

Figure 2. Parotosuchus denwai (ISIA 54). (Al Dorsal and (~31ventral views of the skull. Scale bar= 5 cm.

Journal of African Ewth Sciences S. BANDYOPADHYAYand D.P. SENGUPTA

L Figure 3. Parotosuchus crookshanki IISIA 55). (A) Dorsal and (l3) ventral views of the skull. Scale bar = 5 cm. palatal vacuity, which is single and reniform but and Estes, 1969). This is the second Hadrokkosaums- longitudinally divided by a narrow and low strip of like skull found from anywhere in the world. There bone appearing inside the vacuity. One of the spe- are, however, certain important differences that cies, Parotosuchus denwai, (Fig. 2) is a large form require the Denwa brachyopid to be distinguished with a broad snout and an almost open otic notch as a separate taxon. It is also important to note that defined by posteriorly-pointed tabular horns curved the mandible and the skull of Hadrokkosaurus from at tips. The other species, Parotosuchus crookshanki the Moenkopi Formation were collected from two (Fig. 31, has a slender snout, anteromedially flattened different localities and questions have been raised tabular horns, which define a semiclosed otic notch on their being part of the same animal (Morales, (closure is more marked than that of the other spe- 19871. Jupp and Warren (19861, while working on cies), and an extremely broad body of the pterygoid temnospondyl mandibles, commented that the Ha- behind the interpterygoid vacuities. drokkosaurus mandible has many characteristics of The other temnospondyl is a brachyopid represen- an archosaur which were not apparent in any temno- ted by a well-preserved skull along with both the spondyl. It should be noted that the skull and the mandibles. The skull is typically parabolic in outline mandibles of the new brachyopid were found arti- with large anterolateral eyes, widely-spaced nares culated. Moreover, the mandible is very similar to and a deeply vaulted palate. Tabular horns and the that of Hadrokkosaurs, and the so-called archosaur otic embayment are absent. The mandible has a characters as mentioned by Jupp and Warren (1986) very long retroarticular process. Only the anterior are all present in the mandible of this new brachyopid third of the total mandible length is dentigerous. specimen. The skull roof morphology, and particularly the large A trematosaurid rostrum, represented by the tip extension of the post glenoid area of the mandible, and another fragment immediately behind that, has are comparable with Hadrokkosaurus bradyi from been described by Welles (I 9931 from the Denwa the Moenkopi Formation of North America (Welles Formation. The rostrum has been assigned by Welles

236 Journal of African Earth Sciences Middle Triassic vertebrates of India

Figure 4. (A) Dorsal and (B) ventral views of the left humerus of the Denwa rhynchosaur. Scale bar = 4 cm.

(1993) to Lonchorhynchinae: a subfamily which inclu- proximal end to the entepicondylar foramen (broken) des all the extremely slender snouted trematosaurs. and 28.5 cm wide near the head. The femur (Fig. Apart from the temnospondyls, the Denwa Forma- 6) is stout, thick and is 43.5 cm long, 17.2 cm tion contains a dicynodont, a rhynchosaur and a wide proximally, 16 cm wide distally and 8 cm wide small archosaur. The rhynchosaur is represented by at the shaft. a complete humerus and some fragmentary limb It should be noted that the Denwa reptilian material bones. The humerus (Fig. 4) is 21 cm long, 9 cm allows identification only to the family level. The wide proximally and 11.8 cm wide distally. A long large size and massiveness of one of the dicynodonts cylindrical centrum, limb fragments and some teeth are comparable only with Stahleckeria potens of indicate the presence of a probable archosaur in the Santa Maria Formation of Brazil. the Denwa fauna. Dicynodont fragments are represented by several cranial and post cranial bones of two distinctly AGE OF THE DENWA FAUNA different sizes. Two fragmentary squamosals and The Denwa Formation consists of three successive an articular, several centra, neural arches, ribs, an lithological units. The basal sand dlominated unit ilium and a femur of a medium-sized dicynodont occurring just above the Pachmarhi Formation is show typical kannemeyeriid features. The femur (Fig. poorly fossiliferous. The middle unit, consisting of 51 is remarkably slender; it is 29.3 cm long, 10.5 violet mudstone alternating with white to greenish, cm wide proximally and 10 cm wide distally. calcareous, fine to medium sandstone, has yielded Besides the above material, very large-sized P. denwai, dicynodonts (both medium- and large- vertebrae, humerus, femur, ilium and ischium of sized) and Unio shells. The upper part comprises another dicynodont have also been collected. The brick red mudstone, subordinate white sandstone dorsal vertebrae range in length from 18 to 20 cm and peloidal calcirudite/calcarenite and has produced from the base of the centra to the tip of the Ceratodus, P. crookshanki, a rhynchosaur, the Ha- transverse process (the neural spines are broken). drokkosaurus-like brachyopid and fossil wood. The Part of a very large, stout and thick humerus has dicynodonts and P. denwai continue to occur in the also been found. The bone is 35 cm long from the upper unit, as do the Unio shells.

Journal of African Earth Sciences 237 S. BAND YOPA DH YA Y and D. P. SENGUPTA

Figure 5. (A) Dorsal and /B) ventral views of the left femur of the medium-sized Denwa dicynodont. Scale bar = 4 cm.

It should be noted that P. denwai is more primitive Moenkopi Formation of ‘Early’ (not the earliest) than P. crookshanki as far as the otic notch closure Anisian age (Morales, 1993) has the characteristic and the width of the pterygoid and parasphenoid brachyopid Hadrokkosaurus and Eocyclotosaurus are concerned. P. denwai has some similarity with (Welles, 1967) and dicynodont ichnospecies. The P. megarhinus from the N’tawere Formation of the brachyopid is similar to that of the upper assemblage Luangwa Valley of Zambia, which has been con- of the Denwa, but Eocyclotosaurus has not been sidered to be older than the P. pronus-bearing Manda found in the Denwa Formation. No brachyopid is Formation of Tanzania (Chernin and Cosgriff, 1975). noted from the Yerrapalli Formation. In fact, in the The temnospondyl members also show a close entire Pranhita-Godavari Valley sequence, no bra- similarity with those of the Moenkopi Formation of chyopid material has so far been collected. Two western North America. The otic notch closure of chigutisaurids and a metopsaurid from the Late Tri- the parotosuchid of the Torrey fauna1 zone of the assic Maleri Formation are the only representatives Moenkopi Formation, which has been dated as Early of short-faced temnospondyls of the Pranhita- Spathian (Welles, 1967; Morales, 1987, 19931, is Godavari Valley; the latter two are again not found similar to P, denwai. The Wupatki Member of the anywhere in the Satpura Valley. Moenkopi Formation of Late Spathian age contains The other Denwa fauna1 elements consist of P. peabodyi (Welles and Cosgriff, 1965) and Steno- Ceratodus, at least two dicynodonts, a rhynchosaur cephalus birdi, Cosgriffius campi (a lonchorhynchine and probably an archosaur. These broad groups are trematosaurid) and various ichnospecies. The otic also present in the Yerrapalli Formation. The di- notch closure of P. peabodyi is similar to that of P. cynodonts are represented by a medium-sized and crookshanki of the upper unit of the Denwa For- a large-sized animal. The stout and massive limb mation. The uppermost Holbrook Member of the bones of the latter resemble Stahkeckeria potens

238 Journal of African Earth Sciences Middle Triassic vertebrates of India

A

Figure 6. (Al Dorsal and IB) ventral views of the left femur of the large-sized Denwa dicynodont. Scale bar = 4 cm.

and indicate the probable presence of a stahleckeriid. DISCUSSION However, the Denwa dicynodonts, at present, cannot As has already been mentioned, the Middle Triassic be properly compared with the Yerrapalli dicynodonts vertebrates also occur in the Yerrapalli and Bhimaram until identifiable skull material is collected. The two Formations of the Pranhita-Godavari Valley. The parotosuchids and the new brachyopid from the Yerrapalli Formation occurs above thle Kamthi For- Denwa Formation are therefore the only fauna1 mation, which is of Permo-Triassic age, containing elements which can be used for correlation at Glossopteris flora and Brach yops laticeps. Among present. the Yerrapalli fauna (Table 21, Parotosuchus raja- The parotosuchids of the Denwa Formation do reddyi, with its distinctly expanded tabular horns not show the entire trend of advancement, as do and constricted otic notch, is considered to be more their counterparts from the Moenkopi Formation. advanced than the typical Early Triassic capitosau- It is possible to assign an Early Anisian age to the rids. RoyChowdhury (1970a) considered P. rajareddyi upper fauna of the Denwa on the basis of the rare to be closer to P. prows. The rhynchosaur, Meso- brachyopid, which is noted only from the upper dapedon kuttyi, is characterised by maxillary tooth part of the Denwa and in the Holbrook Member rows, dissected by two anteroposteriorly-directed of the Moenkopi Formation. If this holds good, longitudinal grooves (Chatterjee, 1980) and is the rest of the lower part of Denwa would be comparable to Stenaulorhynchus stockleyi of the older and could be earliest Anisian or even Late Manda Formation of Tanzania. The most common Spathian, like the Wupatki Member of the same member of the Yerrapalli Formation is the kanne- formation. meyeriid, Wadiasaurus indicus (RoyChowdhury,

Journal of African Earth Sciences 239 S. BAND YOPADHYA Y and D. P. SENGUPTA

Table 2. Vertebrate fossils from the Yerrapalli Tanzania; the latter has been referred to the Middle Formation of the Pranhita-Godavari Valley Anisian by Anderson and Cruickshank (1978). Hence the Yerrapalli Formation can also be assigned to the Fauna1 List Stratigraphical Affinities Middle Anisian. Fish The Bhimaram Formation, occurring immediately Dipnoan Ceratodus sp. Mesozoic above the Yerrapalli Formation, is a thick sandstone- Actinopterygian Saurichthys sp. Mesozoic dominated horizon previously believed to be unfossili- Amphibians Capitosaurid Parotosuchus Middle Triassic ferous (Jain eta/., 1964; Sengupta, 1970). However, rajareddyi Kutty et a/. (1987) noted the presence of at least a Reptiles temnospondyl and a dicynodont from this horizon. Dicynodonts Wadiasaurus indicus Middle Triassic The temnospondyl fragments indicate the presence Rechnisaurus Middle Triassic cristarhynchus of a large capitosaurid in this horizon. Only a snout Cynodont Trirachodontid teeth, Lower or of a dicynodont has been collected; Kutty et a/. genus and species Middle Triassic (1987) suggested that this dicynodont appears to indeterminate be different from those of the Yerrapalli dicynodonts Rhynchosaurid Mesodapedon kuttyi Middle Triassic Prolacertid unnamed genus and Middle Triassic as it has a very pointed snout. Of the Yerrapalli species dicynodonts, Wadiasaurus has a tapering snout not Rauisuchid unnamed genus and Middle Triassic ending in a point, which is typical of Kannemeyeria, species Placerias or Rabidosaurus, whereas Rechnisaurus has a wide and blunt snout. However, nothing con- 1970b; Bandyopadhyay, 1988). The other dicyno- clusive can be said about the Bhimaram dicynodont dont is Rechnisaums cristarhynchus (RoyChowdhury, until more material is found. 1970b; Bandyopadhyay, 19891, which is also found The Bhimaram Formation occurs between the in the Manda Formation of Tanzania (Cox, 1991) Middle Anisian Yerrapalli Formation and the Carnian and the Puesto Viejo Formation of Argentina (De- Maleri Formation (Kutty and Sengupta, 19891, hence Fauw, 1993). Of the two Yerrapalli diapsids, the it can be tentatively deduced that the Bhimaram prolacertid has a small skull and thecodont dentition, Formation has an age ranging from Late Anisian to but shows advancement over the South African Ladinian. Prolacerta (Dasgupta, 1993). The other is a rauisu- From the above discussion it can be concluded chid with a large skull and blade-like recurved serra- that, in India, three Middle Triassic horizons of three ted teeth (Dasgupta, 19931, which closely resembles slightly different ages occur. While the Denwa For- Ticinosuchus and perhaps also Mandasuchus. mation, with a fauna resembling that of the Moen- While discussing the Middle Triassic vertebrates kopi, is assigned to the Early Anisian, the Yerrapalli of the world, Anderson and Cruickshank (1978) Formation, considered as equivalent to the Manda precisely delineated the Yerrapalli fauna as Anisian, Formation, appears to be younger than the Denwa which was later corroborated by Cox (1991 I. The and is assigned to the Middle Anisian. The Bhimaram fauna1 assemblage of the Yerrapalli Formation shows Formation, occurring above the Middle Anisian Yerra- a close similarity to that of the Manda Formation of palli Formation and below the Late Triassic (Carnian)

Table 3. Correlation of the Middle Triassic vertebrate-bearing horizons of India, northwestern USA and Tanzania

Age Pranhita- Satpura Basin Northwestern Tanzania I(Ma) I Godavari Valley INDIA USA AFRICA

240 Journal of African Earth Sciences Middle Triassic vertebrates of India

Maleri Formation, should therefore be considered Hughes, T.H.W., 1877. Wardha valley coalfields. Memoir as Late Anisian to Ladinian (Table 3). Further Geological Survey India 13 (11, 71. Jain, S.L., Robinson, P.L., RoyChowdhury, T.K., 1964. A new discovery of better material from the Denwa and vertebrate from the Triassic of Deccan, India. Quarterly Bhimaram Formations will strengthen the above Journal Geological Society London 120, 1 15-l 24. conclusions. Jupp, R., Warren, A.A., 1986. The mandibles of the Triassic temnospondyl amphibians. Alcheringa 10, 99-l 24. Kutty, T.S., Jain, S.L., RoyChowdhury, T.K., 1987. Gondwana sequence of the northern Pranhita-Godavari valley: its ACKNOWLEDGEMENTS stratigraphy and vertebrate fauna. Palssobotanist 36, 263- The authors express their deep gratitude to Prof. 282. T.K. RoyChowdhury of the Indian Statistical Institute Kutty, T.S., Sengupta, D.P., 1989. The Late Triassic formations of the Pranhita Godavari valley and their vertebrate fauna1 for his constant encouragement and suggestions succession - A reappraisal. Indian Journal Earth Sciences throughout the work. Most of the temnospondyl 16 (3-4), 189-206. material from the Denwa Formation was collected Lydekker, R., 1885a. Fossil Reptilia and Batrachia. by the late R.N. Mukherjee. The line drawings were Palaeontologia lndica Series 4 1 (31, l-20. Lydekker, R., 1885b. The Reptilia and Amphibia of the Maleri done by Mr Ajoy K. Das. Finally, the authors are and Denwa Groups. Palasontologia lndica Series 4 1 (5), l- grateful to the Indian Statistical Institute for financial 20. support. Lyddekker, R., 1890. Catalogue of the fossil Reptilia and Amphibia. British Museum (Natural History), London. pp. 145-146. Morales, M., 1987. Terrestrial fauna and flora from the Triassic REFERENCES Moenkopi Formation of the Northwestern United States. Anderson, J.M., Cruickshank, A.R.I., 1978. The biostratigraphy Journal Arizona - Nevada Academy Sciences 22, l-l 9. of the Permian and the Triassic. Pala3ontologia Africana 21, Morales, M., 1993. Tetrapod biostratography of the Lower - 15-44. Middle Triassic Moenkopi Formation. In: Lucas, S.G., Morales. Bandyopadhyay, S., 1988. A kannemeyeriid dicynodont from M. (Eds.), The Nonmarine Triassic. New Mexico Museum of the Middle Triassic Yerrapalli Formation. Philosophical Natural History and Sciences Bulletin 3, pp. 355-356 Transactions Royal Society London 8320, 185-233. Mukherjee, R.N., Sengupta, D.P., 1998. New capitosaurid Bandyopadhyay, S., 1989. The mammal-like reptile amphibians from the Triassic Denwa Formation of the Satpura Rechnisaurus from the Triassic of India. Palaeontology 34, Basin, Central India. Alcheringa 22, 317-327. 305-312. RoyChowdhury, T.K., 1970a. A new capitosaurid amphibian Chatterjee, S., 1980. The evolution of rhynchosaurs. from the Triassic Yerrapalli Formation of the Pranhita-Godavari Ecosystems Continentaux Masozoique. Memoir Society valley. Journal Geological Society India 19 (2), 155- 162. Geology France 139, 75-65. RoyChowdhury, T.K., 1970b. Two new dicynodonts from the Chatterjee, S., RoyChowdhury, T.K., 1974. Triassic Gondwana Triassic Yerrapalli Formation of Central India. Pa&ontology vertebrates from India. Indian Journal Earth Sciences 1 (I), 13 (11, 132-144. 96-l 12. Sengupta, S., 1970. Gondwana sedimentation around Chernin, S., Cosgriff, J.W., 1975. Further consideration of the Bheemaram (Bhimaram) in the Pranhita - Godavari Valley, Capitosaurids from the Upper Luangwa Valley, Zambia. India. Journal Sedimentary Petrology 40, 140-l 70. Palaaontolgia Africana 18, 143-l 48. Welles, S.P., 1967. Arizona’s giant amphibians. Pacific Crookshank, H., 1936. Geology of the northern slopes of the Discovery (California Academy Sciences, San Francisco) 20 Satpuras betwen the Morand and Sher rivers. Memoir (41, 10-15. Geological Survey India 66, 173-381. Welles, S.P., 1993. A review of the Lonchorhynchinae Cox, C.B., 1991. The Pangaea dicynodont Rechnisaurus and trematosaurs (Labyrinthodontia), and a description of a new the comparative biostratigraphy of the Triassic dicynodont genus and species from the Lower Moenkopi Formation of faunas. Pa&ontology 34, 767-784. Arizona. Palaeobios 14 (3), l-24. Dasgupta, K., 1993. New reptiles (Diapsids: Archosauro- Welles, S.P., Cosgriff, J.W., 1965. A revision of the morph) from the Triassic Yerrapalli Formation, Deccan, labyrinthodont family Capitosauridae, and a description of India. Ph.D. dissertation, University of Calcutta, Calcutta, Parotosuchuspeabodyi, n. sp. from the Wupatki Member of India, 125~. the Moenkopi Formation of northern Arizona. University DeFauw, S., 1993. 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