Kurmademys, a New Side-Necked Turtle (Pelomedusoides: Bothremydidae) from the Late Cretaceous of India

Home , Araripemydidae, Emydura, Foxemys, Rosasia

PUBLISHED BY THE AMERICAN MUSEUM OF NATURAL HISTORY CENTRAL PARK WEST AT 79TH STREET, NEW YORK, NY 10024 Number3321,16pp.,4®gures February 27,2001

EUGENE S. GAFFNEY,1 SANKAR CHATTERJEE,2 AND DHIRAJ K. RUDRA3

ABSTRACT The Maastrichtian Kallamedu Formation of southern India near the village of Kallamedu, Tamil Nadu, has yielded skulls and postcrania of a new genus of side-necked turtle. Kurma- demys kallamedensis, new genus and species, is based primarily on a single well-preserved skull. Kurmademys is a pelomedusoid pleurodire belonging to the family Bothremydidae Baur, 1891, with these bothremydid characters: (1) exoccipital-quadrate contact, (2) incisura columellae auris closed by bone, and (3) eustachian tube and stapes separated by bone. Kurma- demys is unique among known bothremydids in having extensive temporal emargination, a small postorbital, a large precollumellar fossa, and a foramen posterius canalis carotici interni formed completely by the basisphenoid.

INTRODUCTION from the Neogene of Burma and Pakistan (see Pleurodires, or side-necked turtles, are not Wood, 1970). A skull of a pleurodire, Shwe- part of the recent fauna of the Indian subcon- boemys pisdurensis, has been described by Jain tinent, but they are known as fossils from the (1977, 1986) and an unnamed bothremydid late Cretaceous into the Neogene. Their record has been mentioned in Singh et al. (1998). in India, however, is very poor. Summaries of Shweboemys pisdurensis appears to be a this record are in Wood (1970), de Broin broad-jawed podocnemidid related to Shwe- (1987, 1988), and Jain (1986). Good skulls of boemys and Stereogenys (Wood, 1970). The the podocnemidid Shweboemys are known bothremydid skull (SD S/VPL 1125) is pres-

1 Curator, Division of Paleontology, American Museum of Natural History. 2 Paul Whit®eld Horn Professor of Geology and Curator of Paleontology, Museum of Texas Tech University, Lubbock, Texas. 3 Former Professor and Head, Geology Unit, Indian Statistical Institute, Calcutta, India. 2 AMERICAN MUSEUM NOVITATES NO. 3321 ently being studied by Gaffney, Sahni, Singh, Werner, 1998). Other skulls of pelomedusoids and Schleich. This skull is a bothremydid, but are Araripemys (Meylan, 1996), and an un- distinct from Kurmademys. The shell of pelo- named Santana genus (FR 4922) (Gaffney and medusoids is notoriously conservative and in- Meylan, 1991). A general treatment and de- adequate to distinguish bothremydids from po- scription of pleurodire skulls, turtle skull mor- docnemidids even when completely known. phology and terminology, and a literature re- Thus, the Indian pleurodire record rests on the view are in Gaffney (1979). two skulls indicated above and the new ma- This paper is intended to name and brie¯y terial from Kallamedu. describe this new taxon. More detailed com- Shweboemys pisdurensis consists of a parative descriptions and a systematic anal- skull, nearly complete shell material, and ysis are part of a larger project on pleurodires some postcranial elements from the Maas- (Gaffney, Meylan, and Wood, 1997; Gaffney, trichtian Lameta Formation at Pisdura and Tong, Chatterjee, Moody and Hirayama, Dongargaon, in Maharastra State, central In- 1998). dia. We have not seen this material, but the We use Lapparent de Broin and Werner's skull does appear to be similar to the de- (1998) informal reference to a Bothremys scribed Shweboemys of Wood (1970). The Group and a Nigeremys Group (which in- unnamed bothremydid skull of Singh et al. cludes Taphrosphys). These continue to form (1998) is from the Maastrichtian intertrap- monophyletic taxa in most of our current pean Green Tuff bed of Amboli Quarry, Jo- analyses. Contents of these groups are as fol- geshwari, Bombay. Features of this specimen lows: Bothremys GroupÐBothremys, Rosa- are noted in the comparisons below. sia, Zolhafah, Foxemys; Nigeremys GroupÐ Although the Bothremydidae was named Nigeremys, Arenila, Taphrosphys. as early as 1891 by George Baur, the term fell into disuse for most of this century and Institutional Abbreviations the few included taxa, particularly Bothremys and Taphrosphys, were simply included in ISI Indian Statistical Institute, Calcutta, In- the Pelomedusidae. Antunes and Broin dia (1988) and Broin (1988) revived Bothremy- FR Forschungsinstitut Senckenburg, Frank- didae, provided a new diagnosis, and added furt, Germany AMNH American Museum of Natural History, taxa, such as Rosasia, based on skulls and New York, USA shells. Recent papers on fossil pleurodires, such as Meylan (1996), Broin de Lapparent and Werner (1998), and Tong et al. (1998), Anatomical Abbreviations use the Antunes and Broin (1988) terminol- bo basioccipital ogy, in which Bothremydidae, Podocnemi- bs basisphenoid didae, and Pelomedusidae (restricted to Pe- ex exoccipital lusios and Pelomedusa) are contained in the fpcci foramen posterius canalis carotici interni Pelomedusoides (which equals Pelomedusi- fr frontal dae in the classic sense). Bothremydids are ju jugal now recognized as a more widespread and mx maxilla diverse group than previously considered. na nasal Useful reviews of the literature on bothre- op opisthotic pa parietal mydids can be found in Broin (1988) and An- pal palatine tunes and Broin (1988). Previously described pf prefrontal bothremydid skulls are as follows: Bothremys pm premaxilla (Gaffney and Zangerl, 1968; Gaffney, 1977), po postorbital Taphrosphys (Gaffney, 1975), Rosasia (Antu- pr prootic nes and Broin, 1988), Foxemys (Tong et al., pt pterygoid 1998), Zolhafah (Lapparent de Broin and Wer- qj quadratojugal ner, 1998), Arenila (Lapparent de Broin and qu quadrate Werner, 1998), and Nigeremys (Bergounioux so supraoccipital and Crouzel, 1968; Lapparent de Broin and sq squamosal vo vomer 2001 GAFFNEY ET AL.: KURMADEMYS 3

Fig. 1. Type locality of Kurmademys kallamedensis, ISI R152, near the village of Kallamedu, Tamil Nadu, southern India. The turtle material occurs here in a 6 in. thick pocket of ®ne grained sandstone and clay in the upper Maastrichtian Kallamedu Formation.

SYSTEMATICS short in contrast to Foxemys and Bothremys; ORDER TESTUDINES LINNAEUS, 1758 triturating surfaces triangular in contrast to MEGAORDER PLEURODIRA COPE, 1864 (FIDE narrow in Nigeremys Group, but not greatly GAFFNEY AND MEYLAN, 1988) expanded as in Foxemys, Rosasia, Zolhafah, HYPERFAMILY PELOMEDUSOIDES COPE, 1868 and Bothremys; no pit in triturating surface in contrast to Bothremys and Rosasia; max- FAMILY BOTHREMYDIDAE BAUR, 1891 illa-quadrate contact absent; palatine exten- Kurmademys, new genus sively exposed on triturating surface as in all TYPE SPECIES: Kurmademys kallamedensis, Bothremys Group; antrum postoticum large new genus and new species. in contrast to small in Bothremys; eustachian DISTRIBUTION: Maastrichtian of central In- tube separated from stapes as in all other dia. bothremydids; incisura columellae auris ETYMOLOGY: Kurma, ``turtle'' in Sanskrit, closed as in Bothremys and Taphrosphys; in allusion to the second-stage incarnation of precolumellar fossa large as in pelomedusids, Lord Vishnu as a turtle in Hindu mythology. but unique among bothremydids; pterygoideus depression moderate as in Rosasia, not DIAGNOSIS deep as in Foxemys; foramen posterius ca- A genus of bothremydid pleurodire with nalis carotici interni completely enclosed in triangular skull, orbits dorsolaterally placed, basisphenoid unique among pelomedusoids; not dorsally as in Bothremys; extensive tem- supraoccipital-quadrate contact present as in poral and cheek emargination similar to Pe- all Bothremys Group; foramen stapedio-tem- lusios and Pelomedusa, but unique among porale visible in dorsal view in contrast to all bothremydids; jugal enters orbit; postorbital other bothremydids. 4 AMERICAN MUSEUM NOVITATES NO. 3321

Fig. 2. Kurmademys kallamedensis, n. gen. & sp., ISI R152. A, dorsal; B, ventral; C, right lateral; D, posterior; E, left lateral; F, anterior.

Kurmademys kallamedensis, new species praoccipitalis, and part of the left quadratojugal. TYPE SPECIMEN: ISI R152, a nearly com- TYPE LOCALITY: Near the village of Kal- plete skull lacking the dorsal part of the pre- lamedu, Tamil Nadu, southern India. Map of frontals, the posterior part of the crista su- locality is in Sastry et al. (1972). 2001 GAFFNEY ET AL.: KURMADEMYS 5

Fig. 3. Key for ®g. 2.

HORIZON: Kallamedu Formation of the described from other exposures of the Kal- Ariyalur Group. Formation named and de- lamedu (Matley, 1929; Yadagiri and Ayyas- scribed by Sastry et al. (1972), who corre- ami, 1987). The Kurmademys locality is a lated it with the uppermost Maastrichtian; the small pocket of ®ne-grained sandstone and Cretaceous-Tertiary boundary is its upper clay, about 6 in. thick. It also contained croc- limit (Sastry et al.: 6). Dinosaurs have been odiles, gar scales, and freshwater gastropods 6 AMERICAN MUSEUM NOVITATES NO. 3321 and bivalves and is interpreted as a freshwater pond deposit. DIAGNOSIS: As for genus. ETYMOLOGY: For the Kallamedu Forma- tion. REFERRED MATERIAL: ISI R155A, partial skull; ISI R155B, partial skull; ISI R155C, partial skull; ISI R158, partial skull; ISI R159, partial skull.

DESCRIPTION The Kurmademys kallamedensis type skull, ISI R152 has a premaxilla-condyle median length of 47.2 mm, a maximum width of 49.4 mm, and a height from condylus mandibularis of the quadrate to the top of the skull roof of 28.2 mm. The order of elements and the order of topics within elements follows Gaffney (1979). More general features of the pleurodire skull and explanations of terminology can also be found in Gaffney (1979) and are not repeated here. Although at present six skulls of Kurma- demys are known, the type specimen, ISI R152, forms the basis of this description, with a few contributions from the other specimens. The type skull is the best preserved and nearly complete. The other ®ve skulls are not as well preserved and are not yet ful- ly prepared.

PREFRONTAL Almost all of both prefrontals are missing in the type skull, ISI R152. The only frag- ments of prefrontal remaining are the ven- tralmost tips of right and left prefrontals ly- Fig. 4. Kurmademys kallamedensis, n. gen. & ing on the medial side of the maxillae in the sp. Partially restored ventral view. See ®g. 3 for anterior orbital walls. There is more of the bone identi®cations. right prefrontal than the left. The dorsal area of the prefrontals has been restored arbitrari- breaks rather than sutures, so the contact ly on ISI R152. However, ISI R158 has pre- with the prefrontals is not determinable. frontals preserved. However, the extent preserved is comparable The shape and extent of prefrontal in Kur- to the frontal as seen in Pelomedusa and Pe- mademys is consistent with that seen in most lusios, so it is not likely that much frontal is other pelomedusoids, such as FR 4922. They missing. meet for their entire length on the midline. As in other pelomedusoids, the frontal of There are no nasals. Kurmademys contacts the postorbital laterally and the parietal posteriorly. Its contacts FRONTAL and relative size are very similar to Pelusios Both frontals are nearly complete in ISI and Pelomedusa. On the ventral surface the R152, but their anteriormost margins are frontal of Kurmademys has a parasagittal 2001 GAFFNEY ET AL.: KURMADEMYS 7 ridge separating fossa orbitalis from the sul- otic as in most other pelomedusoids. Poste- cus olfactorius. The sulcus is slightly narrow- riorly the parietal contacts the supraoccipital er anteriorly than in Pelusios, but widens and prootic as in other pleurodires. The pa- posteriorly as in Pelusios and most pelome- rietal of Kurmademys also has a short ventral dusoids. The processus inferior parietalis process below its lateral margin that contacts meets the posterior edge of the frontal ridge the dorsal process of the palatine in the lat- dorsally as in FR 4922, without a ventral eral wall of the sulcus palatinopterygoideus process as in Pelusios. (Antunes and Broin, 1988). In other pelomedusoids, such as Baurue- mys, FR 4922, and Pelusios, the frontal is JUGAL thickened lateral to the sulcus olfactorius The jugal in ISI R152 is preserved on both ridge and forms a wall for the fossa orbitalis sides, but surrounding contacts are seen only anteriorly and a dorsal margin of the ptery- on the right side. go-palatine channel posteriorly. This wall is The jugal is relatively small in Kurmade- usually continuous with the posterior orbital mys, similar to Pelusios and Pelomedusa, wall laterally and ventrally. However, in Kur- and much smaller than in podocnemidids. mademys the frontal is relatively thin and The jugal is exposed in the posterior margin there is no distinct connection between the of the orbit, but its exposure is reduced by a sulcus olfactorius ridge and the posterior or- posterodorsal process of the maxilla. Ven- bital wall. The pterygo-palatine channel is trally the jugal contacts the maxilla, and pos- relatively open dorsally, at the anterior end, teriorly the quadratojugal. The jugal is not in contrast to most other pelomedusoids, exposed on the dorsal edge of the cheek which have a more restricted ridge of bone emargination due to a contact of the quad- here. ratojugal and maxilla. This is unusual because Kurmademys has a cheek emargination PARIETAL at least as extensive as in Pelomedusa and Both right and left parietals are complete Pelusios, and they have a large exposure of in ISI R152, except for small cracks and the jugal along this emargination. breaks. The medial process of the jugal in ISI The dorsal plate of the parietal is not ex- R152 is preserved and visible on both sides. tensive in Kurmademys. Anteriorly there is a It is basically similar to that area described transverse contact with the frontal, and lat- by Gaffney (1979: ®gs. 53, 130) for Pelusios. erally a contact with the postorbital. There is The medial process of Kurmademys contacts no contact between the parietal and quadra- the palatine medially and the maxilla ante- tojugal; the postorbital is widely exposed riorly. The jugal forms part of the anterior along the edge of the temporal emargination. wall of the adductor muscle chamber as in The temporal part of the parietal is about as most pelomedusoids. The jugal contacts the extensive as in the living Pelomedusa, which postorbital medially and the pterygoid ven- is slightly more emarginate than most Pelusios. tromedially. Ventrally the jugal contacts the The ventral wall of the parietal, the pro- maxilla. All of these contacts are quite sim- cessus inferior parietalis, can be seen in both ilar to Pelusios. sides of ISI R152. The wall is similar to that in other pelomedusoids. The anterior margin QUADRATOJUGAL of the wall is formed completely by the pa- The quadratojugal in ISI R152 is nearly rietal and curves ventrally without the pos- complete on the right side. Some small parts terior indentation seen in some Pelusios. of the dorsal and ventral margin are probably Ventrally the parietal contacts the pterygoid, missing, but based on the tapering of the which is low until it reaches the area anterior bone and on surrounding bones, very little of to the foramen nervi trigemini, where the the quadratojugal is missing. On the left side, pterygoid/parietal contact is more dorsal. The however, only the anterior part and a small foramen nervi trigemini in Kurmademys is fragment of the posterodorsal contact with formed by the parietal, pterygoid, and pro- the quadrate are preserved. 8 AMERICAN MUSEUM NOVITATES NO. 3321

The quadratojugal of Kurmademys is un- lusios, Pelomedusa, and FR 4992 have a usually small for pelomedusoids, because the large antrum postoticum, but the antrum of temporal and cheek emarginations are both Kurmademys is even larger. The size in FR extensive. Kurmademys is unique among 4992, Pelusios, and Pelomedusa is interpret- bothremydids in having such a small quad- ed as the primitive condition for Pelomedu- ratojugal and narrow temporal arch. The soides, because this is the condition in chel- quadratojugal contacts the quadrate posteri- ids. The Kurmademys condition is tentatively orly, the postorbital anterodorsally, the jugal interpreted as a unique autapomorphy of this anteriorly, and the maxilla anteroventrally. genus, because our current analyses show The dorsal margin of the quadratojugal forms Kurmademys deep within the Bothremydidae. the lateral edge of the temporal emargination and the ventral margin forms the dorsal edge POSTORBITAL of the cheek emargination. The postorbital is preserved on both sides of ISI R152. The posterior edge of the left SQUAMOSAL postorbital is missing bone when compared with the more complete right postorbital. Both right and left squamosals are present The size and relations of the postorbital in in ISI R152 and both have some damage to Kurmademys are very similar to the postor- their posterolateral margins, but are other- bital in Pelusios and Pelomedusa. The post- wise complete. orbital of Kurmademys lies between the orbit As in most turtles, the squamosal of Kur- anteriorly and the temporal margin posteri- mademys is cone shaped, forming the pos- orly and forms part of the margins of those terolateral portion of the antrum postoticum. openings. Medially the postorbital contacts As in other turtles, the squamosal of Kur- the frontal anteriorly and the parietal poste- mademys ®ts onto the circular posterior end riorly. Laterally the postorbital contacts the of the quadrate and contacts the opisthotic jugal anteriorly and the quadratojugal pos- medially. Kurmademys has an extensive tem- teriorly. All of these contacts are as in Pe- poral emargination and the squamosal has no lusios and Pelomedusa. FR 4922 differs in contact with parietal or postorbital. The right having a broad contact of the parietal and side of ISI R152 also shows that, although quadratojugal posterior to the postorbital. the squamosal has a narrow anterior process The ventral process of the postorbital is lying on the quadrate, the process does not also similar to that in Pelusios and Pelome- reach the quadratojugal. In Pelusios and Pe- dusa. As exposed in the anterior wall of the lomedusa there is usually a contact between temporal fossa, the postorbital contacts the the quadratojugal and squamosal, although parietal medially, the pterygoid ventrally, there is some individual variation with some and the jugal laterally. On the right side, specimens having a very slight contact or no which is better preserved than the left side, contact. However, none of the specimens of there is a small contact between the parietal Pelusios or Pelomedusa available to us show and pterygoid, preventing exposure of the the degree of reduction of quadratojugal and postorbital in the pterygo-palatine channel at squamosal nor the extent of dorsal exposure this point. In the posterior wall of the fossa of the quadrate seen in Kurmademys. This orbitalis, the ventral process of the postor- feature is unique to Kurmademys. The dor- bital contacts the dorsal process of the pala- solateral surface of the squamosal in Kur- tine medially in a sloping suture. Laterally mademys is rounded, with no parasagittal the postorbital contacts the jugal. The medial ridge or wall seen in other Pelomedusoides. surface of the ventral process of the postor- This ridge is a function of the degree of tem- bital forms the lateral wall and the lateral part poral emargination and is present to a vary- of the roof of a relatively short pterygo-pal- ing extent in all other Pelomedusoides. atine channel. The antrum postoticum is preserved on both sides of ISI R152, and its internal extent PREMAXILLA is visible. The antrum is larger in Kurma- Both premaxillae are present in ISI R152 demys than in any other Pelomedusoides. Pe- and are nearly complete. 2001 GAFFNEY ET AL.: KURMADEMYS 9

Laterally the premaxilla contacts the max- soids. The apertura is not produced anteriorly illa in a parasagittal suture, and it contacts as in Pelusios and Podocnemis. The dorsal the other premaxilla medially. The posterior process of the maxilla in Kurmademys is margin of the premaxilla forms at least part thinner than in Pelusios and Pelomedusa and of the apertura narium interna, but has a bro- is similar to FR 4922. The ventral margin of ken edge medially. The broken edge does not the orbit, is formed by the maxilla, which has show a sutural surface anywhere and there is a dorsal process along the posterior margin no fragment of a vomer, but it is possible that of the orbit reducing the contribution made one was present. The premaxilla in FR 4922 by the jugal to the orbit. Forms such as FR has a process of the maxilla lying behind it, 4922 and Podocnemis have a posterodorsal but this is absent in ISI R152. The dorsal process, but it is separated from the orbit by surface of the premaxilla forms part of the the jugal. Pelusios and Pelomedusa do not ¯oor of the fossa nasalis. In Kurmademys the have this process, but its expression is vari- premaxillae curve dorsally to form a sharply able in pelomedusoids. The posterior edge of rising median ridge in the fossa. This median the maxilla forms the margin of the cheek ridge is present in other pelomedusoids, but emargination. Between the cheek emargina- it is lower and smaller than in Kurmademys. tion and the jugal, the maxilla contacts the The ventral surface of the premaxilla bears quadratojugal. the continuation of the labial ridge. The la- The horizontal plate of the maxilla is ex- bial ridge of Kurmademys is narrower than posed in the orbital ¯oor where the maxilla in Pelusios and lacks the anterior projection contacts the palatine medially and forms a of the margin of the apertura narium externa small part of the border of the large foramen in that form. This area between the apertura orbito-nasale. Posteriorly the maxilla con- narium externa and labial ridge is very thin tacts the jugal. in Kurmademys, similar to FR 4922. Kur- The triturating surfaces of Kurmademys mademys has a shallow median notch similar are narrow anteriorly and widely expanded to Pelusios and wider than in FR 4922. In posteriorly. The maxilla itself, however, ta- contrast to Pelusios and Pelomedusa, Kur- pers posteriorly, so that the palatine forms mademys has a posterior extension to the pre- the posterior and medial portion of the trit- maxilla that bears a distinct, ventrally facing urating surface. The triturating morphology concavity on the midline that is the ventral is similar to that in Foxemys, which is also surface of the dorsal ridge in the fossa na- narrow anteriorly and expanded posteriorly, salis. Lower jaws of Kurmademys show a with a signi®cant contribution from the pal- marked symphyseal hook. The ¯at part of the atine. In both skulls the narrow, anterior part triturating surface narrows considerably from has a raised medial edge along the lingual the maxilla to form a narrow shelf between ridge. They differ in that Foxemys has a the concavity and the labial ridge. This mor- slight pinching of the snout anteriorly, as in phology is also seen in Bothremys and Ro- Rosasia; this is absent in Kurmademys, sasia as well as Neochelys and some gener- where the snout is straight. Foxemys has two alized cryptodires like baenids. There is no accessory ridges, absent in Kurmademys. sign of a foramen praepalatinum. The triturating surface in Kurmademys is raised anteriorly along the margin of the ap- MAXILLA ertura narium interna. The posterior expanded area is slightly concave. There are no ac- Both maxillae of ISI R152 are complete cessory ridges on the triturating surface, except for the distal ends of the dorsal pro- which ends posteriorly in a V-shaped margin cesses, which are missing. completely formed by the palatine. The vertical or alveolar plate of the maxilla is deep and fairly massive, not narrow as VOMER in FR 4922, but similar to Podocnemis. The maxilla forms the apertura narium externa There is no vomer present nor are there anteriorly and it is wider at its base than in sutural surfaces remaining for a vomer. How- FR 4922, similar to most other pelomedu- ever, the bone edges in this area are not en- 10 AMERICAN MUSEUM NOVITATES NO. 3321 tirely complete, and the morphology sur- the postorbital and a short ventral process of rounding the apertura narium interna is very the parietal, all visible on the posterior sur- close to that in Foxemys, which has a well- face of the postorbital wall. developed vomer. Thus it is quite possible that one was present in Kurmademys. QUADRATE

PALATINE Most of both quadrates are complete and free of matrix. Part of the medial area of the Both palatine bones are present in ISI left quadrate is broken and partially restored R152, but are missing some of the anterior with something awful. edges that form the margin of the apertura The squamosal lies at the posterolateral narium interna and the possible vomerine corner of the quadrate and its relations and contact. contacts with the quadrate in Kurmademys The anterolateral part of the palatine in are similar to those in Pelusios and Pelo- Kurmademys contacts the maxilla and forms medusa. The quadrate exposure along the lat- the posteromedial part of the triturating sur- eral edge of the temporal emargination pre- face. The triturating surface is a low platform vents contact of squamosal and postorbital. that ends in a V-shaped margin completely Anteriorly the quadrate contacts the quadra- formed by the palatine. Foxemys is similar to tojugal, but the contact is relatively small be- Kurmademys in this area. The palatine forms cause of the extensive temporal and cheek the posterior margin of the apertura narium emargination. The quadratojugal contact is interna, but most of this margin is missing in smaller in Kurmademys than in Pelusios, Pe- ISI R152. The choanal grooves are barely lomedusa, or FR 4922. discernible in Kurmademys; they are better Most of the quadrate is involved in the for- de®ned in Foxemys. The foramen palatinum mation of the cavum tympani and its two posterius is formed in the palatine-pterygoid spaces, the antrum postoticum and the pre- suture by both bones (®g. 4), as in Foxemys collumellar fossa. The antrum postoticum of and in contrast to FR 4922, Pelusios, Pelo- Kurmademys is unusually large for pelome- medusa, and Podocnemis, in which most of dusoids; it is as large as the antrum in Emy- the foramen is in the palatine. As in other dura, the presumed primitive condition for pleurodires, there is a median contact with pelomedusoids. The antrum of Kurmademys the other palatine and a transverse, posterior is swollen to completely ®ll the area inside contact with the pterygoid. On the dorsal sur- the space formed by squamosal and quadrate. face both right and left palatines are visible The precollumellar fossa is also deep and and free of matrix. The palatine forms the very large in Kurmademys, and also com- medial part of the orbital ¯oor and the lateral parable in size to primitive chelids. However, margin of the large foramen orbito-nasale. other pelomedusoids, such as Pelusios and Posteriorly the palatine of Kurmademys Pelomedusa, also have a large precollumellar has a large dorsal process forming the lateral fossa. In many features, the cavum tympani wall of the pterygo-palatine channel. This of Kurmademys is more primitive than in process contacts the jugal laterally, the post- other bothremydids. orbital dorsolaterally, and the parietal dor- The other feature of interest in the cavum somedially. The process tapers dorsally, so tympani is the incisura columellae auris, that its medial edge is higher than its lateral which still has the stapes present in the right edge. The medial edge forms that lateral quadrate of ISI R152. The incisura is reduced margin of the adductor channel. This dorsal to a completely closed, small foramen con- process of the palatine reaches the parietal in taining only the stapes in Kurmademys, in the postorbital wall, which is quite unusual contrast to the open incisura of Foxemys. Ar- and has not been found so far in other pe- enila, Zolhafah, Bothremys, and Taphros- lomedusoids. The posterior wall of the orbit phys have the closed incisura as in Kurma- is complex in pelomedusoids and particularly demys but in Araripemys, FR 4922, and chel- so in Kurmademys. Behind the dorsal process ids, it is open. The combination of a com- of the palatine is the ventromedial process of pletely closed incisura columellae auris with 2001 GAFFNEY ET AL.: KURMADEMYS 11 a gigantic antrum postoticum is a combina- the exoccipital. The basioccipital contact of tion unknown so far in pleurodires. The the quadrate characterizes the Bothremydi- quadrate of Kurmademys has a kidney- dae plus Podocnemididae, but the exoccipital shaped cavum tympani as in Bothremys. Al- contact is more restricted, found so far only though the incisura columellae auris is com- in the Bothremydidae. pletely closed by bone, behind it is a groove The medial contacts of the quadrate in- formed in the quadrate for the eustachian clude the opisthotic and prootic as in other tube. Dorsally the groove slopes up to a hor- turtles. The roof of the fenestra postotica (ad- izontal, straight-edged ridge that separates itus canalis stapedio-temporalis) has a low the eustachian tube surface from the opening parasagittal ridge showing the passage of the for the stapedial artery in the fenestra pos- more lateral stapedial artery from the more totica. medial (and ventral) lateral head vein. This The medial contact of the quadrate is with ridge is largely formed by the quadrate, with the prootic anteriorly. As in nearly all other the opisthotic contributing medially. The fe- turtles, the prootic and quadrate form the fo- nestra postotica of Kurmademys is subdivid- ramen stapedio-temporale. The foramen sta- ed by bony partitions, so that the lateral head pedio- temporale is certainly placed more an- vein and stapedial artery are separated from teriorly in Kurmademys than in Emydura, but the more medial parts of the fenestra posto- not any more anteriorly than in FR 4922 or tica. The opisthotic forms this wall dorsally Pelusios and Pelomedusa. The canalis sta- and the quadrate forms it ventrally. On both pedio-temporalis can be followed posteriorly sides of ISI R152 the contact of these two on the right side to the aditus canalis stape- bones is broken, and a small amount of com- dio-temporalis, which is partially divided pression is visible on the left side. The quad- from the rest of the fenestra postotica, show- rate also forms the ventral margin of a small ing the entry of the stapedial artery into the opening medial to the one just described, but skull. A separate canal for the stapes itself is lateral to the foramen jugulare posterius, present from the incisura columellae auris to which seems to be a remnant of a more open the aditus canalis stapedio-temporalis as in fenestra postotica. In the Bothremydidae in Bothremys and Taphrosphys. general the fenestra postotica is strongly sub- Behind the prootic there is a contact with divided and separated by bony partitions. In the supraoccipital in ISI R152 (®g. 3A), that Kurmademys the bony partitions are thinner intervenes between the usual opisthotic-pro- and some are probably represented by carti- otic contact. This contact also occurs in all lage or thin bone, allowing the collapse and other Bothremys Group taxa. The opisthotic breakage of foramen edges during fossiliza- contact appears to be reduced in favor of ex- tion. There is a well-developed quadrate-ex- pansion of the supraoccipital contact when occipital contact medially as in Bothremys; compared with a more generalized pleurodire this is considerably more extensive than the like Pelusios. The quadrate-opisthotic con- small contact in FR 4922. tact is anteromedial to posterolateral in Kur- The presumed foramen chorda tympani in- mademys, between the supraoccipital and ferius is present on the posterior surface of squamosal. the processus articularis, roughly similar in On the ventral surface the quadrate in Kur- position to Podocnemis. Because of glue and mademys contacts the pterygoid anterome- crud on the right quadrate, this is only visible dially from the base of the condylus mandi- on the left. bularis along the anterolateral edge of the processus articularis, as in most pleurodires, PTERYGOID such as Emydura and Pelusios. Kurmademys has a medial process of the quadrate as in Both pterygoids are preserved in Kurma- other pleurodires, that contacts a narrowly demys and both are nearly complete. Most of exposed prootic, and broadly contacts the ba- the thin pterygoid ¯ange extending ventrally sisphenoid. Behind the basisphenoid, the from the quadrate process is missing from quadrate has a broad contact with the basi- both pterygoids; the right one is more pre- occipital. Dorsal to that the quadrate contacts served than the left. 12 AMERICAN MUSEUM NOVITATES NO. 3321

On the ventral surface the pterygoid con- gion inside the cavum cranii is variably ob- tacts the palatine in a roughly transverse su- scured by pieces of matrix. The crista pter- ture that trends slightly anterolaterally. The ygoidea is relatively low. The pterygoid foramen palatinum posterius is formed in the forms the ventral margin of the foramen ner- palatine-pterygoid suture as in nearly all vi trigemini as in other bothremydids, but the bothremydids. Medially the pterygoids meet foramen is not placed very close to the fo- on the midline for a bit less than half their ramen stapedio-temporale as it is in many length. They are separated posteriorly by the other bothremydids, such as Bothremys and triangular basisphenoid. Foxemys. The pterygoid forms the ¯oor of As in all pleurodires, there is a laterally the sulcus palatinopterygoideus, which lies projecting processus trochlearis pterygoidei. between the side wall of the cavum cranii In Kurmademys the processus does not ex- and the processus trochlearis pterygoidei. tend at a sharp right angle as in the Santana The anterior contacts of the pterygoid at bothremydids (Gaffney et al., in press), but the base of the processus trochlearis ptery- is only slightly less than a right angle, much goidei are visible on both sides of ISI R152. as in Foxemys. It is not as acute as in chelids The pterygoid plus the palatine and jugal and Araripemys. The ¯ange or web that ex- form the postorbital wall, as exposed poste- tends ventrally from the base of the proces- riorly in the adductor muscle chamber. The sus trochlearis pterygoidei along the quadrate pterygoid has a very narrow contact with the process in all pleurodires is mostly missing parietal medially, broader contacts with the in Kurmademys. The portion preserved is postorbital more laterally, and with the jugal consistent with that seen in other bothremy- most laterally. dids. The posterolaterally extended quadrate SUPRAOCCIPITAL processus in Kurmademys is narrower and longer than in FR 4922, Araripemys, pelo- The supraoccipital in ISI R152 is complete medusids, and chelids. In these groups the ventrally and anteriorly, but is missing the process is relatively ¯at and more horizontal, posterior part of the crista supraoccipitalis. while in Kurmademys and some other both- The supraoccipital in turtles is Y-shaped, remydids, such as Foxemys, it is narrower with paired lateral projections forming the and more vertical. This condition seems to medial part of the cavum labyrinthicum on be related to the presence of a ventrally con- each side. In cryptodires and most pleurodi- cave depression in the posterolateral part of res the supraoccipital has a tripartite suture, the pterygoid in these forms (®g. 4). This with the prootic and opisthotic visible on the depression is presumably the pterygoideus dorsal surface of the otic chamber. It is un- muscle attachment site. In Kurmademys the usual to ®nd that in a group of bothremy- depression is shallower than in Nigeremys didsÐthe Bothremys Group of Lapparent de and Foxemys, but it covers a larger area. Its Broin and Werner (1998)Ðthe supraoccipital margins are not as well de®ned in Kurma- contacts the quadrate and separates the pro- demys as it is in those taxa. There is no de- otic from the opisthotic. In Kurmademys this velopment of an overhang of this depression unusual condition is present. The supraoccip- by the pterygoid as in the Podocnemididae. ital on the right side has a broad contact with The foramen posterius canalis carotici in- the quadrate laterally and separates the pro- terni in Kurmademys lies entirely within the otic from the opisthotic. On the left side the basisphenoid (®g. 3B); the pterygoid does supraoccipital is complete and separates the not participate in its formation as it does in prootic and opisthotic, but the quadrate is many other bothremydids. The Kurmademys damaged. This degree of quadrate contact by condition is unique within pelomedusoids. the supraoccipital is similar in Kurmademys, The posterior margin of the pterygoid con- Bothremys, Foxemys, and Rosasia. The contacts the narrowly exposed prootic between tact is absent in Taphrosphys and indeter- the basisphenoid and quadrate contacts. minate in Zolhafah, Nigeremys, and Arenila. Most of the dorsal surface of the pterygoid The crista supraoccipitalis is usually rela- is visible in Kurmademys, although the re- tively short in bothremydids. In Kurmademys 2001 GAFFNEY ET AL.: KURMADEMYS 13 it is broken posteriorly and its length is in- occipital. A shallow, median concavity lies determinate. between the paired tuberculae and is formed almost entirely by the basioccipital. EXOCCIPITAL Both exoccipitals are preserved and only PROOTIC lack the condylus occipitalis, although there Both prootics in ISI R152 are preserved; is some breakage around the posterior foram- the right one is nearly complete and the left ina. one has a partially eroded dorsal surface. The exoccipital contacts the supraoccipital The prootic is exposed on the dorsal and dorsally, the opisthotic dorsolaterally, the anterior surface of the otic chamber with the quadrate ventrolaterally, and the basioccipital following contacts: the parietal medially, the ventrally. The quadrate-exoccipital contact quadrate laterally, the supraoccipital posteri- occurs in all Bothremydidae and is absent in orly, and the pterygoid ventrally. There is no other pleurodires. prootic-opisthotic contact in Kurmademys. Dorsomedially the exoccipital forms the The foramen stapedio-temporale is formed in lateral and ventral margin of the foramen the prootic-quadrate suture. In contrast to all magnum. Ventromedially the exoccipital pre- other bothremydids, the foramen opens an- sumably participates in the formation of the terodorsally rather than anteriorly. It is visi- condylus occipitalis, but this structure is bro- ble in dorsal view in Kurmademys, but in ken off at its base in ISI R152, so the pres- other bothremydids it is barely or not visible ence or absence of the basioccipital in the in dorsal view. The position of the foramen condylus is not determinable. The foramen stapedio-temporale in Kurmademys is very jugulare posterius in Kurmademys is formed similar to that in pelomedusids and chelids. entirely by the exoccipital. The bone sur- The foramen nervi trigemini is formed by the rounds most of the foramen, but on each side prootic dorsolaterally, the parietal dorsome- the foramen is open laterally due to the pres- dially, and the pterygoid ventrally. The fo- ence of a narrow ®ssure. This ®ssure is dif- ramen is best preserved on the right side; the ferent in shape on both sides and may be due left foramen nervi trigemini is larger and has to breakage, in which case the original con- broken edges. dition of the foramen would be closed. Kur- On the ventral surface of ISI R152, the mademys has two foramina nervi hypoglossi prootic is exposed where the three bones (the as in all other pelomedusoids; they lie near pterygoid, basisphenoid, and quadrate) meet the base of the condylus occipitalis, ventro- (®g. 4). This is in the deepest part of the lateral to the foramen magnum. The more concavity formed by these bones. The con- dorsal foramen is formed entirely within the cavity is presumed to be for the pterygoideus exoccipital, but the more ventral one is muscle attachment. The prootic is exposed formed in the exoccipital-basioccipital su- and has a very similar shape on both sides, ture. so this is not interpreted as a consequence of preservation or an artifact. The prootic has a BASIOCCIPITAL distinct foramen, here interpreted as the fo- The basioccipital in ISI R152 is nearly ramen nervi facialis (VII), for the facial complete. A small amount of breakage is vis- nerve, always closely associated with the ible on each tuberculum basioccipitale and prootic ossi®cation. The form of the prootic the broken condylus occipitalis does not exposure in Kurmademys is not like that in clearly show basioccipital sutures. any other pleurodire. The primitive condition The basioccipital of Kurmademys has a of the ventral prootic exposure occurs in broad, transverse contact with the basisphe- chelids, pelomedusids, and Araripemys, all noid anteriorly. Posterolaterally the basioc- of which have a large prootic exposure with cipital contacts the quadrate, and posterodor- the foramen posterius canalis carotici interni sally there is a broad contact with the exoc- in the prootic. In all bothremydids, podoc- cipitals. The tuberculum basioccipitale is nemidids, and FR 4922 the prootic does not formed about equally by the quadrate and ex- contain the internal carotid as it does in chel- 14 AMERICAN MUSEUM NOVITATES NO. 3321 ids and pelomedusids. In FR 4922 the prootic tact with the pterygoids trends posterolater- is partially exposed in a narrow space be- ally and anteromedially, and the angle this tween the basisphenoid and quadrate, similar suture makes with the midline is similar to to the primitive position found in chelids and that in Foxemys and Zolhafah. At the antero- pelomedusids, but is distinctly posterior to lateral corner of the basisphenoid, between the prootic, as exposed in Kurmademys. It is the pterygoid and quadrate contacts, is a likely that the prootic exposure in Kurma- short contact with the prootic. In nearly all demys is not a retention of a primitive state, bothremydids the prootic is covered, so this but is an autapomorphy of this genus, prob- contact is unusual. The lateral margin of the ably related to the development of the pter- basisphenoid is a long, parasagittal contact ygoideus concavity that removed covering with the quadrate. This contact in Kurma- elements. Small areas of the prootic are var- demys is longer than in any other bothre- iably exposed in other bothremydid skulls mydid; Foxemys and Polysternon most close- due to erosion or the development of the con- ly approach it. Posteriorly the basisphenoid cavity. has a transverse contact with the basioccipital. In contrast to all other bothremydids, in OPISTHOTIC Kurmademys the foramen posterius canalis carotici interni is formed completely by the Both opisthotics are preserved in ISI basisphenoid, without participation of the R152; the right one is nearly complete and pterygoid. However, the foramen is very the left one is missing a small part anteriorly. close to the pterygoid suture, particularly on In dorsal view, the opisthotic has these the right side. The foramen posterius canalis contacts: supraoccipital anteromedially, the carotici interni in Kurmademys is also placed squamosal laterally, and the exoccipital pos- farther anteromedially than in any other teromedially. There is no prootic-opisthotic bothremydid. This could be explained mor- contact. The opisthotic in Kurmademys ends phologically by a reduced ossi®cation of the posteriorly at about the same level as the canalis caroticus internus posteriorly. The ca- squamosal; it does not extend posteriorly be- nalis in all bothremydids travels anterome- yond the squamosal as in pelomedusids, Ar- dially and slightly dorsally to enter the sella aripemys, and FR 4922. Ventrally the opis- turcica. If the canalis in a form like Bothre- thotic forms the roof and some of the sub- mys (which has the foramen posterius canalis divisions of the fenestra postotica. In ISI carotici interni placed far posterolaterally) R152 the opisthotic divides the fenestra pos- were to be exposed by the removal of bone totica into two portions: a more lateral one ventrally, the foramen would appear to mi- that seems to have contained the stapedial grate anteromedially along the path of the ca- artery and lateral head vein, and a more me- nalis caroticus internus. It is possible that this dial one with unknown contents. The medial condition could result from the development foramen has a ®nished dorsal margin, al- of a deep pterygoideus muscle concavity, though as preserved, the ventral margin is which is formed directly ventral to the ca- broken. Presumably the more medial fora- nalis caroticus internus. Although Kurmade- men just held cartilage as in living turtles mys has a distinct pterygoideus concavity, it with an open fenestra postotica (Gaffney, is relatively shallow compared to such forms 1979). as Foxemys, and Foxemys does not have the foramen posterius canalis carotici interni BASISPHENOID placed anteromedially. The basisphenoid is complete and well The dorsal surface of the basisphenoid in preserved in ISI R152. The cavum cranii is Kurmademys has the dorsum sellae and sella largely free of matrix and some of the dorsal turcica visible. The dorsum sellae overhangs surface of the basisphenoid is visible. the sella turcica and has the foramen anterius The basisphenoid in Kurmademys is not canalis carotici interni also hidden in dorsal strongly triangular as in other bothremydids view and lying at the posterolateral corner of but is more pentagonal. It is a relatively large the sella turcica. There is a small processus element, wider than long. The anterior con- clinoideus on the left side; the right one is 2001 GAFFNEY ET AL.: KURMADEMYS 15 broken. The shape and general proportions of the basisphenoid. Foxemys, Polysternon, Ro- the dorsum sellae and sella turcica are similar sasia, Zolhafah, and Bothremys can be united to those in Pelusios. The degree of overhang by having a foramen stapedio-temporale very of the dorsum sellae, however, is greater in close to the foramen nervi trigemini and a Kurmademys than it is in Pelusios. The ros- broad preorbital part of the skull, features ab- trum basisphenoidale is fused into a single sent in Kurmademys. structure, but its anterior end shows the two Kurmademys has characters in common ossi®ed trabeculae rather than the single ros- with the Bothremys Group, but the state of trum seen in Pelusios. There is no sign of a the current analysis shows only a few steps foramen nervi vidiani in the left sulcus cav- from Kurmademys as the sister group to the ernosus. The right side still has some matrix. Nigeremys Group plus Bothremys Group. The addition of as yet undescribed taxa will RELATIONSHIPS help to resolve its phylogenetic position. Kurmademys is a pleurodire because it has ACKNOWLEDGMENTS these synapomorphies of the group listed by Gaffney and Meylan (1988) as diagnostic for We wish to thank our associates in pleu- the Pleurodira: (1) processus trochlearis pter- rodiran studies, P. Meylan, H. Tong, and R. ygoidei present, (2) quadrate process below Wood, for their support and counsel on this cranio-quadrate space, (3) epipterygoid ab- ongoing project. Ed Heck did the photogra- sent, (4) foramen palatinum posterius behind phy and illustrations with his customary ex- orbit, and (5) pelvis suturally attached to car- cellence. We particularly appreciate the help apace and plastron (based on shell material of Judy Galkin in the preparation of the pa- from same locality ascribed to Kurmademys). per. It is a member of the Pelomedusoides (sensu S. L. Jain and A. Sahni provided infor- Broin, 1988; Meylan, 1996; Lapparent de mation and access to specimens, which we Broin and Werner, 1998; Tong et al., 1998), greatly appreciate. We thank the Indian Sta- which is equivalent to the Pelomedusidae in tistical Institute for ®eld logistics and sup- the classical sense (sensu Gaffney and Mey- port, permission to study the specimens, and lan, 1988) because it has these characters: (1) the National Geographic Society for funding nasals absent, (2) prefrontals meeting on this project. midline, and (3) splenial absent (based on lower jaws from same locality ascribed to REFERENCES Kurmademys). Kurmademys can be identi®ed Antunes, M. T., and F. de Broin as a member of the family Bothremydidae 1988. 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