Osteology and Diagnosis of Cherninia denwai along (Capitosauridae) from the Denwa Formation, Satpura Gondwana Basin, Central India    

Osteology and Diagnosis of Cherninia denwai along (Capitosauridae) from the Denwa Formation, Satpura Gondwana Basin, Central India | Authorea try { document.documentElement.classList.add('js'); } catch (e) { } var _gaq = _gaq || []; _gaq.push(['_setAccount', 'G-8VDV14Y67G']); _gaq.push(['_trackPageview']); (function() { var ga = document.createElement('script'); ga.type = 'text/javascript'; ga.async = true; ga.src = ('https:' == document.location.protocol ? 'https://ssl' : 'http://www') + '.google-analytics.com/ga.js'; var s = document.getElementsByTagName('script')[0]; s.parentNode.insertBefore(ga, s); })(); Skip to main content Preprints Collections Wiley Open Research IET Open Research Ecological Society of Japan All Collections About About Authorea FAQs Contact Us Quick Search anywhere Search for preprint articles, keywords, etc. Search Search ADVANCED SEARCH SCROLL This is a preprint and has not been peer reviewed. Data may be preliminary. 7 January 2025 V2 Latest version Share on Osteology and Diagnosis of Cherninia denwai along (Capitosauridae) from the Denwa Formation, Satpura Gondwana Basin, Central India Authors : Pummy Roy , Sanjukta Chakravorti 0000-0002-8766-6176 [email protected] , and Dhurjati Prasad Sengupta Authors Info & Affiliations https://doi.org/10.22541/au.172479300.01115366/v2 Published Fossil Record Version of record Peer review timeline 446 views 165 downloads Contents Abstract Abbreviations Institutional Abbreviations: Materials and methods Systemic Palaeontology Emended Diagnosis: Osteology Mandible Post crania Neural arch and neural spine Ribs Pectoral girdle Forelimb Pelvic girdle Hind limbs Conclusion Acknowlegement Information & Authors Metrics & Citations View Options References Figures Tables Media Share Abstract The Middle Triassic Denwa Formation located within the Satpura Gondwana basin of Central India exhibits a significant presence of temnospondyl amphibians classified under the family Capitosauridae. Prior investigations have documented two taxa of the Capitosauridae, namely Cherninia denwai and Paracyclotosaurus crookshanki , from the Denwa Formation. These prior accounts were predominantly predicated upon two holotype skull specimens, thereby neglecting numerous paratype specimens contained within the collection as well as various associated post-cranial materials. Recently, a diverse assortment of novel specimens pertaining to C. denwai has been unearthed from the Denwa Formation. Utilizing both the newly acquired specimens and previously overlooked specimens, this study presents an amended diagnosis of C. denwai . The newly discovered specimens comprise a partial skull, a mandible, clavicles, interclavicles, vertebrae, neural arches and spines, ulnae, an ilium, a femur, and a fibula, all of which are described herein for the first time. An extensive osteological analysis of the skull and mandible has been conducted. It has been determined that C. denwai coexists temporally with C. megarhina , and both taxa exhibit distinct osteological traits; however, they are recognized as separate and unique species. The Middle Triassic Denwa Formation constitutes a significant component of the Satpura Valley basin, which is recognized as one of the four principal Gondwana basins located in India. The Middle Triassic Denwa Formation is predominantly characterized by heterolithic deposits that encompass sequences of sandstone and mudstone. Positioned beneath the Early Triassic Panchmarhi Formation and above the Jurassic Bagra Formation, the Middle Triassic Denwa Formation is notable for its abundance of Triassic vertebrate fossils, which include a variety of fish fossils, temnospondyls, and the renowned horned Shringasaurus indicus , among others \cite{Bandyopadhyay_2020,SENGUPTA_2021} . The prevailing palaeo-environment of the Denwa Formation is characterized as fluvio-lacustrine in nature, contributing to a larger braided river system. Two capitosaur taxa, Cherninia denwai and Paracyclotosaurus crookshanki have already been described \cite{Mukherjee_1998,damiani2001cranial} from the Denwa Formation (Figure 2). The identifications and descriptions were solely based on two skulls designated as holotypes (ISI A 54 and ISI A 55 respectively). Those descriptions were done mainly on the material collected by the late R.N. Mukherjee of the Geological Studies Unit, Indian Statistical Institute, Kolkata. In the existing description, the figures representing the occipital views of those two taxa were inadvertently swapped (see \cite{Mukherjee_1998} ). However, not all the material collected by him have been used in the existing descriptions. Moreover, in subsequent years, many new materials were excavated by the present authors which included several post cranial elements so far that undescribed for Cherninia denwai from the Middle Triassic Denwa Formation which nudges towards the wholesome detailed osteological description of the Indian capitosaurid Cherninia denwai . Capitosaur phylogeny has always remained a fairly debatable issue over the years. The first systematic grouping of the capitosaurids had been done by \cite{ochev1966systematics} . Welles and Cosgriff (1965) attempted grouping of different capitosaurs through the measurements and/ or their ratios of several skull parameters \cite{welles1965revision} . Primary phylogenetic analyses of the capitosaurs have been done by Ingavat & Janvier (1981) and Milner (1990) \cite{Ingavat_1981,milner1993paleozoic} . Later Jupp and Warren (1986) suggested some identifying characters of the mandibles of the capitosaurs and Warren and Snell (1991) worked on the post cranial elements of that group \cite{Jupp_1986,Warren_1991} . Meanwhile, Shishkin’s (1980) suggested the concept of a diphyletic origin of the otic fenestra in capitosaurs, and Morales & Kamphausen (1984) described a new taxon ( Odenwaldia heidelbergensis ) that appeared to confirm that hypothesis \cite{shishkin1980luzocephalidae,morales1984odenwaldia} . Almost thirty-five years later, Schoch and Milner (2000) and Damiani (2001) reviewed the family and used cladistic analysis on a larger dataset involving computer-aided softwares such as PAUP and MacClade, and came up with contrasting ideas1 \cite{schoch2000stereospondyli,damiani2001cranial} . Damiani, in 2001, proposed the name Mastodonsauridae for the family while Schoch and Milner (2000) retained the family Capitosauride. Steyer (2003) and Schoch (2008 & 2013) also dealt with the characters and phylogeny of capitosaurid \cite{Steyer_2003,schoch2008capitosauria,Schoch_2018} . Schoch (2000, 2008, 2018), like Schoch and Milner (2000) however preferred to retain the family ‘Capitosauridae’. Most of the capitosaurid characters, apart from the diagnostic features of Chernina denwai used below are taken from the above references. Some of the characters of the genus Cherninia are mostly noted from C. megarhina (Chernin 1974) from the Middle Triassic Ntawere Formation of Zambia. It is to be noted that Damiani (2001) put Cherninia in Anisian and Paracyclotosaurus in Ladinian. Abbreviations Anatomical abbreviations : AC: Anterior coronoid; AF: Adductor fossa; AN: Angular; APV: Anterior palatal vacuity; AR: Articular; Cm: Crista muscularis; CT: Cultriform process; D: Dentary; ECT: Ectopterygoid; EO: Exoccipital; F: Frontal; FM: Foramen magnum; GF: Glenoid fossa; IJ : Insulae jugalis; IN: Internal nares; ITPV: Interpteygoid vacuity; J: Jugal; L: Lacrimal; MC: Middle coronoid; MS: Mandibular ; sulcus; MX: Maxilla; N: Narial; NA: Nares; O: Orbit; P: Parietal; PAL: Palatine; PAR: Prearticular; PC: Posterior coronoid; PF: Postfrontal; PMF: Posterior meckelian foramen; PMX: Premaxilla; PNF: Pinneal foramen; PO: Postorbital; POSP: Postsplenial; PP: Postparietal; PQF: Paraquadrate foramen; PRF: Prefrontal; PSP: Parasphenoid; PT: Pterygoid; PTF: Post temporal fenestra; Q: Quadrate; QB: Quadrate boss; QJ: Quadratojugal; SA: Surangular; SMX: Septomaxilla; SP: Splenial; SQ: Squamosal; ST: Supratemporal; STV: Supratemporal vacuity; T: Tabular; V: Vomers. Institutional Abbreviations: ISI: Indian Statistical Institute, Kolkata (Geological Studies Unit) Materials and methods A cumulative total of approximately 30 skeletal elements have now been documented and ascribed to Cherninia denwai (Supplementary Table 1). Drawing from these novel findings, the taxonomic descriptions have been revised to incorporate several new cranial features as well as additional mandibular characteristics, which are presented herein for the first time. Certain post-cranial elements have also been introduced, likewise for the inaugural time. One of the paratypes, a fragmented cranium (ISI A 207), exceeds one meter in length, while a mandible (ISI A 208) approaches 96 cm. Both of these specimens have been unearthed by the senior author DPS from the central stratigraphic section of the Denwa Formation, specifically from violet-hued mudstones, thereby indicating that the central section of Denwa served as a sanctuary for colossal temnospondyls that attained lengths of 4 to 5 meters. Systemic Palaeontology Order Temnospondyli von Zittel 1887-1890 Suborder Stereospondyli von Zittel 1887-1890 Superfamily: Capitosauridea Watson, 1920 [ nom. Trans. \cite{save1935dermal} ex. Capitosauridae \cite{1920} emend \cite{schoch2000stereospondyli} ] Genus: Damiani, 2001 Species: Damiani, 2001(= Mukherjee and Sengupta, 1998); ; Figure 2 - 13 Type horizon Middle Triassic Denwa Formation; Age Anisian. Locality Middle part of Denwa Formation, near Purtala village, Madhya Pradesh, Central India (22°35’38.6” N, 78°32’17.0”E). Holotype ISI A 54, a near complete skull (Figure 2.1). (Figure 2) Paratype A posterior portion of skull found near Kohpani village (ISI A 207; Figure 2.2), nearly complete left mandible near Jhirpa village (ISI A 208; Figure 3- 4). Emended Diagnosis: Synapomorphies with Capitosauridae Pre-orbital part of skull (snout) elongated with orbits placed at the posterior half of the dorsal skull roof; orbital margins raised above the dorsal surface of the skull roof; postero-lateral corners of the cheek anterior to the level of the posterior edge of the tabular horns; well-developed zone of intensive growth on the preorbital and cheek region of the skull; infraorbital sensory canal forms a Z-shaped loop on the lacrimal bone, the lacrimal flexure; deeply incised otic notch; occipital condyles anterior to quadrate condyles; well-developed, lappet -like tabular horns which are partly supported from below by muscular cristae from the paroccipital process; elongated preorbital projection of the jugal, extending to the middle border of the snout; supratemporal bone excluded from the border of the otic notch; postorbital antero-laterally expanded (‘hooked’); crista muscularis of the parasphenoid levels with the posterior border of the pterygoid-parasphenoid suture; unbroken chain of teeth on the vomer-palatine-ectopterygoid series; occipital face of the pterygoid bears a tall, narrow, crest-like flange of bone, the oblique ridge of the pterygoid; posttemporal fenestra triangular in shape; mandible with an antero-posteriorly expanded symphysis; a short accessory sensory sulcus, dorsal to the oral sulcus present. Synapomorphies with Pterygoid-parasphenoid suture short, less than the width of the corpus of the parasphenoid; occiput moderately shallow; skull margins moderately concave lateral to the orbits so that the cheek region is flared; prefenestral division of the palate (the region of the palate anterior to the interpterygoid vacuities) markedly elongated; anterior palatal vacuity paired but set in an oval depression so that the vacuities lie below the plane of the palate; hyper-elongated and exceptionally broad snouts so that parts of the anterior lateral margin of the skull on both sides are almost parallel to each other; skull roof with conspicuous, paired, forked ridges anterior to the orbits; anterior rim of the otic notch angular; temporal sensory canal tending posteriorly to the tips of the tabular horns; internal nares (choanae) are placed extremely far forward than the interpterygoid vacuities. Autapomorphies of Cherninia denwai can be distinguished from Cherninia megarhina the second known species of Cherninia by short and narrow postparietals; the presence of septomaxilla, interpterygoid vacuities pointed both anteriorly and posteriorly; lack of parasphenoid groove; lack of occipital sensory canal; tabular horns posteriorly directed and recurved distally; the occipital margin of the skull is deeply concave; otic notch is broad posteriorly. Skull margins moderately concave lateral to the orbits so that the cheek region is flared, hyper-elongated and moderately broad snouts, lateral margins of the middle part of the skull tapering. The skull roof has conspicuous, paired and forked ridges anterior to the orbits on both sides of the skull. Anterior rim of the otic notch angular (40-60 degrees), tabular horns posteriorly directed and recurved distally, postparietals antero-posteriorly compressed and narrow, deeply concave occipital margin of skull, well expressed lateral line sensory canals, septomaxilla present \cite{Mukherjee_1998} . The pterygoid-parasphenoid suture is short and the interpterygoid vacuities pointed both anteriorly and posteriorly. The skull table is shallow, the region of the palate anterior to the interpterygoid vacuities is elongated, anterior palatal vacuity paired but set in an oval depression so that the vacuities lie below the plane of the palate, cultriform process merges with the vomers and the occiput is moderately shallow. The mandible is slender and long (the largest measuring almost 96 cm in length), distinct post glenoid area (PGA) is present, hamate process of prearticular is well developed but not as pronounced as in Tatrasuchus , labial wall of adductor chamber is dorsally horizontal, prearticular is separated from splenial by dentary and coronoid series, glenoid facet lies below the dorsal surface of dentary. The PGA is a combination of type 1 and type 2 sensu Jupp and Warren (1986) \cite{Jupp_1986} . Osteology Skull roof The detailed osteological description of the dorsal part of the skull roof are as follows: Premaxillae The anterior-most rim of the snout is formed by the premaxillae. The premaxillae suture with the maxillae laterally, with the nasals posteriorly, and forms a part of nares. In ventral view, they contact the vomers posteriorly, the maxillae laterally, and form the anterior rim of the anterior palatal vacuity. The ornamentation is made up of pits and grooves. Maxillae The anterior one-third of the lateral skull margin is formed by the maxillae, commencing from the posterior rim of the nares. Dorsally, they also form the lateral rim of the nares. Septomaxillae were figured by Mukherjee and Sengupta (1998), however, they are ill preserved. Anteriorly, the maxillae are sutured with the premaxillae, medially by the nasals, the lacrimals, and posteriorly by the jugals. In ventral view, the maxillae bordered the premaxillae anteromedially, and medially with the vomers and the palatines. The anterolateral rim of the choanae was formed by the maxillae. Very faint pit and groove ornamentation is present in the anterior part. Nasals : The nasals are quite broad, and form a major part of the anterior skull roof medially. Nasals are paired medially, contacting the premaxillae anteriorly. The nasals form a triple junction posteriorly, contacting the frontals posteromedially and the prefrontals posterolaterally. The nasals are bordered by the maxillae behind the nares and contact the lacrimals laterally. Nasals form posterolateral rims of the narial. Pit and groove ornamentation are present in the anterior-most part and ridge and furrow in the posterior part. Lacrimals They are tear-drop shaped, suturing with maxillae anterolaterally, nasal medially, prefrontal posteriorly, and jugal posterolaterally. Pit and groove in the anterior part and ridge and furrow ornamentation in the posterior part are present. The lacrimals do not enter the orbit margin. Jugals The jugals form the majority of the cheek regions are. Anteriorly, they contact the maxilla and anterolaterally the lacrimals. Laterally, the jugals are sutured anteriorly by the prefrontals and posteriorly by the postorbital. The jugals form a narrow tip and enter the orbit, thereby separating the prefrontals and the postorbital. Posteriorly, the jugals sutures with the squamosals medially and the quadratojugals laterally. Pit and groove ornamentation in the posterior part and ridge and furrow are present in the anterior part. Prefrontals The elongate prefrontals suture with the nasals and lacrimals anteromedially. Laterally, towards the midline, they suture with the frontals and towards the lateral margin with the jugals and form the anteriormost rim of the orbits. Ridge and furrow like ornamentation is present in them. Frontals The frontals are conspicuous, elongate bones, anteriorly subtriangular in shape that contact along the midline. They run from the middle part of the snout region to the middle of the orbits. Anteriorly, they are bounded by the nasals, laterally by the prefrontals, posteriomedially by the parietals, posterolaterally by the postfrontals and forming a part of the rim of the orbit projecting between the prefrontals and the postorbitals. Ornamentation in the anterior part is feeble but in the posterior part pit and groove ornamentation are present. Postfrontals The postfrontals are pentagonal in outline and form the posteromedial margin of the orbits. Anteriorly, they suture with the frontals, posteromedially by the parietals, posteriorly by the supratemporals, and laterally by the postorbitals. Pit and groove ornamentations are present throughout. Postorbitals The postorbitals are wing-like projections, contributing to the posterolateral rim of the orbits. The postorbital contacts the postfrontals medially, and supratemporals posteromedially, projecting into the jugals anterolaterally, thus suturing with them. The postorbitals contact the squamosals posterolaterally. Like the postorbitals, pit and groove ornamentation throughout. Quadratojugals The quadratojugals form the cheeks and the lateral-most part of the skull. Dorsally, they have a sub-rectangular outline, contacting the jugals anteriorly and the squamosals medially. On the ventral side, they contact the quadrates medially. They form a larger part of the lateral and posterior rims of the supratemporal vacuities. In the occipital view, the quadratojugals sutures with the quadrates medially and the squamosals dorsally. Quadratojugals have ridge and furrow ornamentation throughout. Parietals The parietals contact medially, and resembles a trapezium in outline. They enclose the parietal (pineal) foramen along the medial suture towards the posterior half of the bone. The parietals suture with the frontals anteriorly, the postfrontals and the supratemporals laterally, and the postparietals posteriorly. Pit and groove ornamentations are present throughout. Supratemporals The supratemporals are polygonal in outline and contact the postorbitals, postfrontals anteriorly, the parietals medioanteriorly, and the postparietals medioposteriorly. The supratemporal sutures with the squamosals laterally and posteriorly with the tabulars. The supratemporal does not contribute in the rim of the otic notch. Pit and groove ornamentations are present throughout. Squamosals The squamosals form the postero-lateral rim of the skull, almost half of the margin of the otic notch, projecting in between the quadratojugals and the tabulars. The squamosals are bordered by the postorbitals and the jugals anteriorly, the supratemporals medially, and the quadratojugals laterally. Squamosals suture with the tabulars posteromedially. Pit and groove ornamentation present near the otic notch rim which gradually grades into ridge and furrow radially at the potero-lateral corner. Postparietals The postparietals form the posterior concavity of the dorsal rim of the skull. They are polygonal and contact along the midline. The postparietal sutures with the parietal anteriorly, supratemporals and the tabulars laterally. They are much shorter than the parietals. Pit and groove ornamentations are present throughout. Tabulars The elongated polygonal shape of the tabulars broadens anteriorly and directed posteriorly, forming a portion of the margin of the otic notch. They contact the supratemporals anteriorly, the postparietals medially, and the squamosals anterolaterally encircling the otic notch. The tabulars do not suture with the squamosal laterally thereby forming a small embayment between the tabular and the squamosal. In dorsal view, the tabular horns project posteriorly and have a concavity anteriorly. Pit and groove ornamentations are there throughout. Lateral Line Sulci The lateral line sulci are present as pairs of supraorbital, infraorbital, and postorbital sulci. They are more evident laterally, especially in the posterior part of the skull, and weakly impressed medially, and again pronounced across the snout. However, all the lateral line of sulci is discernible. The supraorbital sulci arise from the anterior region of the snout and traversing medially to the nares. They meander anteroposteriorly from the nasals to the prefrontals, running along the prefrontal-lacrimal suture reaching the posterior half of the frontals before weakly terminating on the postfrontals near the orbital midline. The infraorbital sulci begin lateral to the nares and meander anteroposteriorly on the maxillae before bending acutely onto the lacrimals forming a Z- flexure and terminating atop or parallel to the jugal-maxilla sutures. The postorbital sulci originate from the postorbital rim of the orbits. They run in a sigmoid form onto the jugals and quadratojugals first, then along the squamosal-quadratojugal sutures, terminating at the posterior border of the skull. Palate Two series of tooth present on the palate. Vomers Vomers are elongated bones tapering posteriorly. Anteriorly, the vomers sutures with the premaxillae, forming the posterior margin of the anterior palatal vacuity. Immediately behind this opening, the vomers bear teeth row arranged transversely. The vomers also bear palatal tusks, circular in cross-section. Anterolaterally, the vomers contact the maxillae and laterally contact the palatines. The vomers contribute a part of the choanae and to most of the medial rim of the palatal vacuities. At their contact along the interpterygoid vacuities, the vomers form a ‘V-shaped ridge that projects ventrally. Palatines The palatines are crescent in shape and bear the second pair of palatal tusks. The palatines form the posterior rim of the choanae and contact the maxillae anteriorly. The palatine-maxilla sutures run posterolaterally from the choanae to the ectopterygoid and form the anterolateral rim of the interpterygoid vacuity. The palatine has a discontinuous row of teeth more or less becoming larger anteriorly. Pterygoids The pterygoids have a bow-like structure extending between the interpterygoid vacuities and the supratemporal vacuity and forming part of their respective rims. Anterolaterally, the pterygoids contact the ectopterygoids and point contact with the insulae jugalis . Posteriorly, the pterygoids contact the quadrates and form part of the posterior margin of the skull. The pterygoids contact the parasphenoid medially. The pterygoids do not contact the palatines. The pterygoids suture with the exoccipitals ventrally. The palatine ramus of pterygoid bears feeble ornamentation. Parasphenoid The parasphenoid is an unpaired bone present in the middle. The cultriform process of the parasphenoid projects beyond the anterior margin of the interpterygoid vacuities. The parasphenoid-exoccipital suture has a flat and long ‘M’ shaped structure. The crista muscularis of the parasphenoid runs transversely. Laterally, the parasphenoid sutures with the pterygoids. The parasphenoid-pterygoid sutures are as long as the width of the parasphenoid basal plate. The parasphenoid forms the posteromedial rim of the interpterygoid vacuities. The basal plate of parasphenoid is short and parasphenoid pockets are present entirely on the ventral surface of the palate. The cultriform process of the parasphenoid process is placed on a crest posteriorly which gradually merges with the vomers anteriorly. Ectopterygoids The ectopterygoids are longitudinal bones. Posteromedially, the ectopterygoid sutures with the pterygoid, posteriorly with the insula jugalis , and anteriorly with the palatines. The ectopterygoid bears a number of laterally compressed teeth. Ectopterygoids contribute a small portion to the rim of the interpterygoid vacuities. Insulae Jugalis The insulae jugalis are the triangular bones forming a point contact with the pterygoids posteriorly. Anteromedially, insulae jugalis is bound by the ectopterygoid and posteriorly forms a part of the anterior rim of the supratemporal vacuity. Quadrates The quadrates are polygonal-shaped that contact the pterygoids medially and the quadratojugals laterally. Anteriorly, the quadrates contribute to the posterior rim of the supratemporal vacuity, and posteriorly form the posterior margin of the skull. A prominent quadrate boss is present on the occipital face. Exoccipitals The exoccipitals project posteriorly to create the atlas-exoccipital articulation and diverge dorsally to encompass the ventral portion of the foramen magnum. They suture with the parasphenoid anteriorly. The posterior projection of the exoccipitals lies anterior to the projection of the quadrates. The posterior contact surfaces of the exoccipitals with the atlas are sub-oval in shape and are flat and narrow. Skull openings Skull openings Skull openings from the holotype ISI A 54 are listed below: Nares are the two tear-drop-shaped dorsal openings with a maximum length of 4.33 cm and maximum width 2.52 cm for the left and maximum length 4.76 cm and maximum width of 2.63 cm for the right. A small bone, septomaxilla is present at the inside of the anterolateral margin of the nares. The orbits are positioned at the posterior half of the skull with a maximum length of 5.70 cm and 5.85 cm and a maximum width of 3.76 cm and 3.30 cm respectively for left and right and they are elliptical in outline. The orbits are slightly elevated than the plane of the skull roof. The pinneal foramen is subcircular its long axis is 1.68 cm and short axis is 1.03 cm, and is completely enclosed by the parietals. The otic notch is wide open posteriorly and and are bounded by the squamosals and the tabulars. The anterior rim of the otic notch is angular, the angle between the squamosals and the tabulars are 41 and 62 degrees in the left and right respectively. The anterior palatal vacuities are subdivided laterally stretched with a maximum length of 2.5 cm and width of 7 cm. The left one with dimensions 2.08 X 3.22 cm and the right one with 1.46 X 3.49 cm. It is enclosed by the premaxillae and the vomers. The anterior palatal vacuity is bean shaped with a constriction in between. The choanae are two slit-like cavities in the anterior part of the palate. The choanae are quite deformed (taphonmically) with dimension 3.49 X 1.28 cm and 5.07 X 1.21 cm for left and right respectively. The interpterygoid vacuities are the largest openings of the palate, with a ‘D’-shaped outline, with lengths are 28.93 X 8.08 cm and 28.91 X 9.60 cm for left and right respectively. The lateral rims have a parabolic curvature with the posterior curvature more acute than the anterior one. The marginal rims are almost straight. The interpterygoid vacuities are enclosed by the vomers, the palatines, the pterygoids, the parasphenoid and a small portion of the ectopterygoids. The cultriform process of the parasphenoid forms most of the medial margin. The subtemporal vacuities are the posterior most vacuities in the palate. The vacuities are arch shape with a gentle lateral convexity and a more acute medial concavity. The maximum length measured parallel to the midline is 17.30 cm, and the maximum width is 7.90 cm for the left one and 17.47 cm and 9.01cm respectively for the right one. The supratemporal vacuities are enclosed by the pterygoids, the quadrates, the quadratojugal and the insulae jugalis. The tip of the anterior projection of the subtemporal vacuity does not reach half of the interpterygoid vacuities. Occiput The occipital process to postparietal and tabular is extremely short and stout dorso-ventrally in occipital view. The pterygoid oblique ridge is exceptionally short and dorsomedially oriented, located just anterior to the otic notch and approaching the crista falciformis of the squamosal. The descending ramus of the squamosal partially covers the lateral sections of the pterygoid and quadrate and extends dorsolaterally to the paraquadrate foramen at the quadrate. From its posterior corner, the quadratojugal produces a narrow but thicker flange that extends medially for a short distance along the rear surface of the articular. The quadrate bears prominent quadrate boss. The post temporal fenestra is triangular in outline. The torsion in the quadrate is visible in the occipital view. Mandible A well-preserved right mandible of Cherninia denwai (ISI A 208) was recovered from the Baki-Nala section near the village of Jhirpa, measuring almost 96 cm in length (Figure 4). The curvature of the mandible exactly fits with the skull outline of C. denwai . Moreover, skull fragments of C. denwai are found from the same locality. Dentary The dentary forms the dorsal and some of the labial and lingual surfaces of the anterior part of the ramus. The dentary is bound ventrally by the splenials and angular in the labial view and posteriorly by the surangular tapering to a point. In lingual view, it is bound by the splenial and the coronoids ventrally, while it forms the anterior margin of the adductor fossa posteriorly. The dentary and the splenial is present in the anterior symphysis. The main marginal teeth are located on the dorsal surface. The symphysial region of the dentary has one large tusk. Splenial and postsplenial In labial view, the splenial and postsplenial lies ventral to the dentary with the splenial located anteriorly. The region of the posterior meckelian foramen is broken. The ventral surface is not fully preserved. Angular Angular forms the remaining bone in the ventrolabial and some part of ventrolingual surface. It is bound by the dentary anterodorsally, by the postsplenial, anteroventrally, and by the surangular dorsally in labial view. In lingual view, it is bound by prearticular dorsally and articular posteriorly. Surangular The surangular forms most of the posterodorsal part of the labial surface, including that of the postglenoid area. In labial view, the surangular sutures with angular ventrally and with dentary anteriorly. In lingual view, it sutures with articular posteriorly; with angular ventrally and anteriorly with prearticular. In dorsal view, the surangular sutures with dentary anteriorly; lingually with articular and forms lingual margin of the adductor fossa. Coronoid series Coronoid series of bones comprising of anterior coronoid, middle coronoid and posterior coronoid on the lingual surface. They are placed ventral to the dentary and extend from a little behind the symphysis to the leading edge of the adductor fossa. The anterior coronoid sutures with the splenial ventrally, the middle coronoid sutures with the postplenial and some part of the splenial and the posterior coronoid with the prearticular. No dentition is present in the coronoid series nor is there any process present in the coronoids. Prearticular In lingual view, the prearticular forms the posterior half of the ramus. It sutures with postsplenial anteriorly and dorsally with middle and posterior coronoid. The prearticular forms the lingual margin of the adductor fossa and the lingual edge of the glenoid fossa. Prearticular does not form part of post glenoid area. The prearticular does not suture the splenial because it is separated by the coronoid. Articular Articular is the endoskeleton bone and forms all of the glenoid fossa. Anterodorsally, the articular forms the posterior border of the adductor fossa. The articular is enclosed labially by the surangular and lingually by the prearticular. Ventrally, the articular is overlain by the angular. Posteriorly the articular enters the postglenoid area. The posterior extremity of the articular is not completely ossified. Other important features of mandible The hemi-mandible is low except for its posterior area, which is dorsally straight along the posterior two-thirds and slightly curved in the anterior part. In the labial view, the angular shows well−defined ornamentation that becomes less evident in the postsplenial. A highly developed mandibular sulcus is found in the back of the hemi-mandible, and the marked oral sulcus extends all the way to the dentary. There is also a tiny accessory sulcus that runs from the mandibular sulcus to the surangular. The posterior meckelian foramen is not preserved and anterior meckelian foramen is absent. The adductor fossa is quite big, with an oval shape and dimensions of 21.2 cm along the long axis and 2.3 cm along the short axis. The articular lingual border, the surangular labial border, the antero-labial border of the dentary, the antero-lingual border of the posterior coronoid, and the posterior border of the glenoid fossa all encircle the adductor fossa. One anterior tusk and multiple teeth with relatively depressed bases can be seen in the dentary. As is characteristic in temnospondyls, both the dentary and the splenial are a part of the symphyseal area. The articular and surangular, but not the prearticular, are located in the postglenoid region, which is considerably extended. To the ventral side of the articular is the angular. The foramina chorda tympani is situated more on the labial side of the post-glenoid region (dorsal). Although Jupp and Warren (1986) discussed the lower jaw anatomy of temnospondyls and stated that capitosaurs mostly have Type I post glenoid area, the morphology of the post− glenoid area of C. denwai cannot be clearly placed in any of the two types. It shares with Type I the following features: the prearticular does not extend into the postglenoid area, the articular is the major component of the postglenoid area, the angular lies ventral to the articular. On the other hand, as in Type II ( sensu Jupp and Warren 1986), the angular lies labial to the articular and the postglenoid area is slightly elongated. The prearticular hamate process is highly developed, huge, and spatulate. In comparison to the hamate process, the quadrate trochlea is shorter. There were two separate processes called the crista articularis and crista medialis . Underneath the dentary's dorsal surface is the glenoid fossa. The elliptical, sizable adductor fossa has a horizontal and straight labial wall. Torus arcuatus (muscle scar near hamate process) is prominent and the muscle insertion area is comparatively large (Figure 4). Each tooth's base is distinct and elliptical. Nearly all of the sutures are distinct and noticeable. The labial wall of the mandible has noticeable ornamentation. There is one tusk on the symphysis. The shape of the glenoid fossa is rectangular. The adductor fossa's anterior process is elevated and noticeable. Post crania All post-cranial elements have been retrieved from the Middle part of the Denwa Formation, thereby associating them with with Cherninia denwai , as they were discovered in close proximity to the cranial fragments of of Cherninia denwai . Furthermore, the occurrence of the other trematosaurid temnospondyl is exceedingly rare. Vertebrae and Ribs Vertebral column is the most significant component after the skull and mandible. Vertebral elements mainly the intercentra, neural arches, and neural spines were preserved (Figure 5). The post-atlantal and presacral neural arches are two different forms of neural arches found in Permian and Triassic temnospondyls \cite{sulej2007osteology} . The presacral has less developed posterior zygapophyses with a short neural spine and transverse process than the post-atlantal. The neural spines of the post-atlantal have erect spines and transverse processes. Most of the neural arches and neural spines were incomplete and broken in C. denwai. Atlas and axis were not present in the collection. The intercentra were wedge-shaped in the lateral view and crescentic in anterior and posterior views. Distinctly, two types of intercentra were present, type 1 in the anterior part and type 2 in the posterior part. The anterior intercentra were massive, quite thick, and robust. The dorsal most part is blunt. The ventrolateral surface of the intercentrum is concave anteroposteriorly and is finished. The parapophysis, is well developed. Five numbers of anterior intercentra are present (ISI A 209-213). The posterior intercentra are flattened ventrally and the lateral surfaces are more distinct from the ventral face. They are shallower and less ossified than the anterior ones. The parapohyses were directed more ventrally, thus their lower edges project below the ventral line with a triangular outline. There are four posterior intercentra present (ISI A 214-217) (Figure 5). Neural arch and neural spine Neural arch and neural spines (Figure 6) were broken in the specimens present in the middle part of the Denwa Formation. The body of the neural arch extends laterally. A rectangular facet, the area of attachment of the pleurocentrum (Watson, 1958) is present. At the end a squarish facet is present which is the diapophysis. Neural spines were detached from the body of the neural arch. They are short and stout. Eight single broken neural arches (ISI A 218-225); and one pair of neural arches (ISI A 226a & b) are present. Eight neural spines were present (ISI A 227-234). No split neural spines are present. No haemal arches are present. Ribs A total of 30 numbers of rib fragments and one almost complete rib were present (Figure 7). The ribs were distinct and morphologically varied. The position of the ribs has been decided by the existing literature of temnospondyls ribs \cite{dutuit1976introduction,sengupta2002indian,sulej2007osteology} DuTuit 1976, Sengupta 2002, Sulej 2007). The cervical ribs have two separate rib facets. The anterior thoracic rib ( ISI A 235; ISI A 238-250) heads were elliptical in cross-section with a narrow extension for attachment of capitulum and tuberculum. The distal end of the anterior is expanded and plate-like. The posterior thoracic ribs (ISI A 251-266) are cylindrical and have elliptical proximal heads and rounded distal ends (Figure 8). Sacral ribs are expanded both proximally and distally but short and stout (ISI A 236-237). No caudal ribs were preserved. Pectoral girdle Two almost complete and three fragmented interclavicles (ISI A 267 & 268) and one complete left and one fragmented left and one fragmented right clavicle (ISI A 272-274) and six fragmented cleithra (ISI A 275-280) are present. The interclavicles are quite big and robust and its thickness varies from 3 to 5 cm. The clavicles have a long dorsal process originating at the posterolateral corner of the clavicle plate. The dorsal process is slender and twisted. The clavicles are slender and spatulate (Figure 9). Cleithra are slender processes having costae on their surfaces. Scapular crest is present at its head. The shaft's external surface is convex. The internal facet has a concave attachment area for the clavicle and a convex attachment area for the scapula (Sulej 2007). However, all the cleithra are broken (Figure 8). Forelimb The forelimb comprises of radius and ulna. Humerus and radius is not present in the present collection. Only the proximal and distal heads of ulnae are preserved (ISI A 182/3 & 4; ISI A 281-282, Figure 10). The specimen ISI A 182/4 is stated as the distal shaft of ulna by Mukherjee et al, (2019), but the olecranon process is distinct in the specimen, thus the specimen is the proximal shaft of the right ulna \cite{Mukherjee_2019} . The proximal and distal ends of the head are expanded towards the radius. The proximal head is rhombohedral in outline and the posterolateral corners of the head are raised with the olecranon process. On the posterior surface, a low crest is present. The distal heads are laterally flattened articulation surface for ulnae and intermedium. Pelvic girdle The pelvic girdle comprises only the left illum as no other elements are preserved (Figure 11). Overall, the illum is cylindrical with a convex ventral margin that tends to meet the puboischiadic blade (ISI A 283). A depression at the posterior part of the illum is the area of attachment of the acetabulum head which is mostly cartilaginous. The acetabular buttress is thicker anteriorly and laterally projected. The acetabulum is bounded by two notches. The anterior supracetabular notch is slenderer and dorsally placed while the posterior acetabular notch is ventrally placed. The iliac blade is robust and cylindrical and bears some striations for attachment of muscle. The shaft is oval in cross-section. Hind limbs The hind limbs comprise of femur, tibia and fibula. The proximal shaft of a right femur and proximal shaft of left fibula is preserved. Tibia is not present in the collection. The proximal shaft of the right femur is preserved (Figure 12). The femur shows concave anterior and posterior margins. The bone is strong and stout. It bears a medial ridge, adductor crest, is not present in the specimen, but the adductor blade is present housing the fourth trochanter. The proximal articular surface is oval in outline (ISI A 284). The fibula is dorsoventrally flattened, with trapezoidal outline of the proximal head (Figure 13). The anterior margin is concave. The well-developed posterior fibular ridge is visible. This is bordered by a furrow in the front and a raised surface made by two short parallel ridges in the back (ISI A 285). No digit bones were so far present. Conclusion Supported by the new material, an emended diagnosis of C. denwai has been presented here. Paratypes including a partial skull, a mandible, clavicles, interclavicles, vertebrae, neural arches and spines, ulnae, an ilium, a femur and a fibula are also described for the first time. A detailed osteological description of the skull and the mandible has been done. It has been noted that the skull roof has conspicuous, paired and forked ridges anterior to the orbits and the otic notch is angular, tabular horns posteriorly directed and recurved distally. The mandible is slender, distinct post glenoid area (PGA) is present, hamate process of prearticular is well developed and the prearticular is separated from splenial by dentary and coronoid series. The PGA is a combination of type 1 and type 2. The hemi-mandible is low. In the labial view, the angular shows well−defined ornamentation that becomes feeble in the postsplenial. A highly developed mandibular sulcus is found in the back of the hemi-mandible, and the oral sulcus extends all the way to the dentary. The specimen ISI A 182/4 is noted as the distal shaft of ulna by Mukherjee et al, (2019) is found to be the proximal shaft of the right ulna \cite{Mukherjee_2019} . The proximal head of the ulna is rhombohedral and the posterolateral corners of the head are raised with the olecranon process. The acetabulum is bounded by two notches. The anterior supracetabular notch is slenderer and dorsally placed while the posterior acetabular notch is ventrally placed. The detailed osteology and the emended diagnosis of C. denwai is the first step to systematically study the huge array of temnospondyls that are present in Anisian Denwa Formation of Central India. Acknowlegement The present work is a part of the Ph.D. dissertation on the temnospondyls of the Denwa Formation by PR carried out with the financial support from the Indian Statistical Institute (ISI). The authors are grateful to Geological Studies Unit, Indian Statistical Institute for the laboratory and Financial support. We acknowledge Mr. Pushpal Kundu of Photography and Reprography Unit, ISI, Kolkata for all the photography assistance provided and preparator Mr. Lakshman Mahankur for his assistance in preparation of the specimens. We also acknowledge Mr. Swapan Aich, Mr. Ashok Saha and Mr. Sasanka Sekhar Sahu of the Transport unit of ISI, Kolkatafor transport and driving assistance during field- work in the Satpura basin. The authors sincerely thank the unknown peer - reviewers whose comments have significantly helped to develop the manuscript further. Information & Authors Information Version history V1 Version 1 27 August 2024 V2 Version 2 07 January 2025 Peer review timeline Published Fossil Record Version of Record 30 Dec 2024 Published Copyright This work is licensed under a Non Exclusive No Reuse License. Keywords capitosauridae gondwana india temnospondyl the early-middle triassic Authors Affiliations Pummy Roy View all articles by this author Sanjukta Chakravorti 0000-0002-8766-6176 [email protected] View all articles by this author Dhurjati Prasad Sengupta View all articles by this author Funding Information Indian Statistical Institute Prof. Dhurjati Prasad Sengupta Metrics & Citations Metrics Article Usage 446 views 165 downloads .FvxKWukQNSOunydq8rnd { width: 100px; } Citations Download citation Pummy Roy, Sanjukta Chakravorti, Dhurjati Prasad Sengupta. Osteology and Diagnosis of Cherninia denwai along (Capitosauridae) from the Denwa Formation, Satpura Gondwana Basin, Central India . Authorea . 07 January 2025. 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