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Chupacabrachelys complexus, n. gen. n. sp. (Testudines: Bothremydidae), from the Aguja Formation (Campanian) of West Texas

Thomas M. Lehmana; Steven L. Wickb a Department of Geosciences, Texas Tech University, Lubbock, Texas, U.S.A. b Division of Science and Resource Management, Big Bend National Park, Texas, U.S.A. Online publication date: 02 December 2010

To cite this Article Lehman, Thomas M. and Wick, Steven L.(2010) 'Chupacabrachelys complexus, n. gen. n. sp.

(Testudines: Bothremydidae), from the Aguja Formation (Campanian) of West Texas', Journal of Vertebrate Paleontology, 30: 6, 1709 1725 To link to this Article: DOI: 10.1080/02724634.2010.520782 URL: http://dx.doi.org/10.1080/02724634.2010.520782

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Journal of Vertebrate Paleontology 30(6):17091725, November 2010 2010 by the Society of Vertebrate Paleontology

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CHUPACABRACHELYS COMPLEXUS, N. GEN. N. SP. (TESTUDINES: BOTHREMYDIDAE), FROM THE AGUJA FORMATION (CAMPANIAN) OF WEST TEXAS
THOMAS M. LEHMAN*,1 and STEVEN L. WICK2 Department of Geosciences, Texas Tech University, Lubbock, Texas 79409, U.S.A., tom.lehman@ttu.edu; 2 Division of Science and Resource Management, Big Bend National Park, Texas 79834, U.S.A., tenbitsranch@hughes.net
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ABSTRACTChupacabrachelys complexus, n. gen. n. sp., is an unusual bothremydid pleurodire of the tribe Taphrosphyini found in the Campanian Aguja Formation in the Big Bend region of West Texas. The type example is one of the most complete bothremydid specimens known. Its skull and lower jaw are very narrow, triangular, and dorsoventrally compressed, with the coronoid process posterior to midlength of the jaw. The orbits are elongate anteroposteiorly with narrow extensions along the maxilla-prefrontal sutures, and rugose maxillary projections at the anterolateral corners of orbits. Its reduced temporal emargination, weak posteroventral ange on the squamosal, weak lateral extension of the squamosal along the quadrate suture, and posteroventral knob on opisthotic suggest afnity with Taphrosphyini. The shell has six neurals and a nuchal with sharp midline embayment, and is nearly identical to those of Chedighaii and Bothremys. Chupacabrachelys provides additional evidence for the dramatic evolutionary radiation of bothremydid turtles in tropical paralic environments during Late Cretaceous time.

INTRODUCTION Pleurodira or side-necked turtles are today a relict group of low diversity, restricted to freshwater habitats in the southern hemisphere. However, during Late Cretaceous and Paleogene time, pleurodires were taxonomically diverse, inhabited a variety of environments, and had a nearly worldwide distribution (Gaffney et al., 2006). Much of the early literature on fossil pleurodires focused on their relatively conservative shell morphology; it was only the recent discovery of many well-preserved skulls that revealed the magnitude of pleurodire diversity. The extinct family Bothremydidae was among the most widespread of pleurodire groups during their Late Cretaceous and Paleogene acme. Bothremydids have distinctive skulls with wide prefrontals, and in contrast to other pleurodires, have very little cheek emargination retaining a maxilla-quadratojugal contact. Many also possess jaws with unique triturating surfaces indicative of highly specialized feeding strategies. In a monographic study of bothremydids, Gaffney et al. (2006) recognized four distinct clades (Kurmademydini, Cearachelyini, Bothremydini, and Taphrosphyini), which they ranked as tribes. Fragmentary specimens of bothremydid turtles are common in coastal facies of the Upper Cretaceous Aguja Formation in the Big Bend region of West Texas. Most of these (e.g., TMM 43469-1) consist of isolated shell fragments, originally identied doubtfully as ?Taphrosphys (Lehman, 1985) and later as Bothremys (Lehman, 1997; Tomlinson, 1997). However, none of these many fragmentary specimens is sufciently complete to condently identify. In 2005 the authors recovered a complete bothremydid shell, and a second nearly complete skeleton with skull and jaws. These specimens represent a new genus and species of bothremydid, Chupacabrachelys complexus. It is likely that much of the fragmentary material recovered previously from the Aguja Formation may also belong to this species.

*Corresponding

author.

The type specimen of Chupacabrachelys complexus (TMM 45606-1) was collected on the Ten Bits Ranch, north of Study Butte, Texas (Fig. 1). It is remarkably complete, more so than many other Cretaceous bothremydids, and is missing only the hind limbs. As a result, in the present paper most of the axial and appendicular elements are described and illustrated from multiple perspectives. The referred specimen (TMM 45856-1) was recovered from the same stratigraphic interval as the type, about 24 km south in Big Bend National Park (Fig. 1). Other than the shell, the referred specimen has two elements (coracoid and humerus) that can be compared with the holotype. The shells of both specimens have most of the sutures evident on their visceral surfaces, but scute sulci can only be delineated in places. Both appear to be adult individuals; however, the type specimen is only about three-fourths the size of the referred specimen (measurements in Appendix 1). Although it seems likely that the referred specimen pertains to Chupacabrachelys complexus, the shells of some bothremydids (Chedighaii, Bothremys) are nearly identical to each other and to Chupacabrachelys complexus, and so we cannot be certain in the referral. Therefore, in the following description, the features of the referred specimen not clearly evident in the type (nuchal notch, form of mesoplastra) are noted separately. Chupacabrachelys complexus will be compared below primarily with the well-known bothremydids Chedighaii and Bothremys, which have shells remarkably like that of Chupacabrachelys, but radically different skulls. Geologic SettingThe Aguja Formation consists of an eastward-thinning wedge (135 to 285 m) of paralic and marine sandstone interbedded to the west with mudstone and lignite deposited in terrestrial coastal plain environments (Lehman, 1985; Fig. 1). The Aguja Formation is underlain by marine shale of the Pen Formation, and the two units intertongue to the east. Two major depositional sequences are recorded in these deposits, the lower of which is present only in the western Big Bend region. Lehman (1985) informally subdivided the Aguja Formation into several members. The basal sandstone member consists of progradational deltaic and littoral facies. It is overlain by the terrestrial lower shale member, which includes interbedded

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FIGURE 1. Locality map and cross-section showing stratigraphy of the Upper Cretaceous Pen and Aguja Formations in southwestern Texas. The stratigraphic positions of Texas Memorial Museum (TMM) Chupacabrachelys complexus collection sites on Ten Bits Ranch (XB) and near Rattlesnake Mountain (RM) in Big Bend National Park are shown. Symbols on stratigraphic columns show intervals with ostreid bivalves and Ophiomorpha burrows.

sandstone, carbonaceous shale, and lignite. A transgressive marine sandstone, the Rattlesnake Mountain sandstone member, overlies the lower shale (Lehman and Tomlinson, 2004). A westward-thinning marine shale unit (informally referred to as the McKinney Springs tongue of the Pen Formation) is interposed within the Aguja. Overlying this marine shale are deposits of the second depositional sequence of the Aguja, extensive over

the entire Big Bend region and adjacent Mexico. The Terlingua Creek sandstone member is overlain by the upper shale member, which is comprised of sandstone, mudstone, and carbonaceous shale accumulated in coastal environments. The type and referred specimens of Chupacabrachelys complexus were collected from the base of the upper shale member (Fig. 1), about 1 m (TMM 45856-1) to 5 m (TMM 456061)

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FIGURE 2. Quarry diagram showing holotype specimen of Chupacabrachelys complexus (TMM 456061) as preserved. Skeletal elements found beneath others are shaded. Current ripple cross-lamination 10 cm above bone-bearing horizon indicates northwesterly ow during deposition.

above the top of the Terlingua Creek sandstone. Most of the fossil vertebrates of the Aguja Formation, including the two specimens of Chupacabrachelys complexus, were collected from this intervalthe lower part of the upper shale member (reviewed by Rowe et al., 1992; Lehman and Busbey, 2007). Ammonite biostratigraphy (Waggoner, 2006), vertebrate biostratigraphy (Rowe et al., 1992), and radiometric age determinations (Befus et al., 2008) indicate that the upper shale member is Middle to Late Campanian (Judithian) in age. Exact locality information for both specimens is available at the Vertebrate Paleontology Laboratory of the Texas Memorial Museum in Austin, Texas. TaphonomyTMM 456061 was preserved in the base of a very ne-grained cross-laminated sandstone bed with thin discontinuous layers of carbonaceous clay (Fig. 1). No remains of any other vertebrates were found at the site. All parts of TMM 456061 were preserved on the same horizon, disarticulated, with most of the larger elements in a horizontal attitude; smaller elements were jumbled and inclined at varied angles (Fig. 2). The plastron was preserved upright, probably close to where the carcass had originally rested. Most of the small appendicular and axial elements were displaced to the north and west. The lower jaw was detached from the skull. The inverted carapace had separated into anterior and posterior halves along the third costals, covering the skull and other elements. The missing parts of the nuchal, and right anterior peripheral bones, were lost due to mod-

ern erosion; they were exposed when the specimen was discovered (Figs. 2, 3). There is no indication that predation or scavenging was responsible for dismemberment of the carcass. The skeletal elements show little distortion due to compaction, including the carapace and plastron, which preserve their natural curvature. The referred specimen (TMM 45856-1) was preserved in a thick claystone (Fig. 1). It consists of a nearly complete shell, preserved in articulation with the pelvis and resting on the plastron. There are several partially healed conical punctures and long linear gouges in the carapace (Fig. 3). The anterior end of the plastron was also broken and partly healed later in life. Judging from the broad form and spacing of the bite marks, the unsuccessful predator in this case was probably Deinosuchus, whose remains are preserved in the same facies of the Aguja Formation (e.g., Schwimmer, 2002). Only a few appendicular elements are preserved (humerus, coracoid, metatarsal, and a phalanx), protected inside the shell and resting on the visceral surface of the plastron. The posterior carapace margin is encrusted with small oyster spat, and parts of both carapace and plastron are crossed by ne arcuate grazing trails. Although the anterior end of the carapace retains its natural curvature, the rest of the shell is attened. The stratigraphic position of both TMM 45856-1 and TMM 456061, just above the paralic Terlingua Creek Sandstone, and the presence of ostreid bivalves and selachian remains in nearby deposits

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FIGURE 3. Chupacabrachelys complexus, n. gen. n. sp., carapace of TMM 456061 in A, dorsal and B, visceral views; and TMM 45856-1 in C, dorsal view; plastron of TMM 456061 in D, ventral view, and TMM 45856-1 in E, visceral view. Inset diagram shows restoration of both shells at common scale (see Fig. 11) and pattern of crocodilian bite marks on TMM 45856-1.

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FIGURE 4. Chupacabrachelys complexus, n. gen. n. sp., skull of TMM 456061 in A, dorsal, B, ventral, C, right lateral, and D, anterior views; and lower jaw in E, dorsal, F, right lateral, and G, medial views; broken surfaces (cross-hatched), areas covered by sediment matrix (white), and photographs for comparison. Inset diagram provides restoration of skull in dorsal view with extent of visible sutures (and mirror image) shown; asymmetry and distortion of left side removed. Abbreviations: ang, angular; art, articular; cor, coronoid process; den, dentary; fm, fossa meckelii; fr, frontal; in, internal nares; j, jugal; lab, labial ridge; lin, lingual ridge; ma, mandibular articulation; mx, maxilla; op, opisthotic; p, parietal; pmx, premaxilla; po, postorbital; pra, prearticular; prf, prefrontal; ptr, pterygoid trochlear process; q, quadrate; qj, quadratojugal; scm, sulcus cartilaginous meckelii; sym, mandibular symphysis; sq, squamosal.

(Fig. 1) suggest a coastal environment for Chupacabrachelys, and is consistent with the brackish water habitat interpreted for many other North American fossil pleurodires (Gaffney and Zangerl, 1968). Institutional AbbreviationsALAB, Alabama Museum of Natural History, Tuscaloosa; ANSP, Academy of Natural Sciences, Philadelphia; FMNH, Field Museum, Chicago; TMM, Texas Memorial Museum, Austin; YPM, Yale Peabody Museum, New Haven.

DESCRIPTION Order TESTUDINES Linnaeus, 1758 Infraorder PLEURODIRA Cope, 1864 Superfamily PODOCNEMIDOIDEA Cope, 1868 Family BOTHREMYDIDAE Baur, 1891 Tribe TAPHROSPHYINI Gaffney et al., 2006 CHUPACABRACHELYS, new genus

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JOURNAL OF VERTEBRATE PALEONTOLOGY, VOL. 30, NO. 6, 2010 The premaxillae protrude anteriorly beyond the triturating surface in ventral view, and have a coarsely pitted texture differing from that on the rest of skull. A similar pitted surface is found on the premaxillae of Labrostochelys (Gaffney et al., 2006) and in some extant pelomedusoids (Pelusios niger; Gaffney 1979) where it is associated in life with a eshy conical nose. The midline dorsal process of the premaxilla is very low. The nasals are absent, and the prefrontals meet on the midline, forming a forwardly projecting process that partially divides the nares. The orbitonarial bar is very narrow, much narrower than the diameter of the orbit. The orbits have a distinct ventral rim, and face as much laterally as dorsally, unusual traits for bothremydids, particularly so considering that the skull is somewhat compressed dorsoventrally. The orbits are very elongate anteroposteriorly and have narrow extensions along the prefrontal-maxillary sutures. The prefrontal-frontal suture is visible on the right side at the narrowest point between the orbits. The prefrontal comprises only a small part of the anteromedial border of the orbit. The frontal-parietal suture is visible in places near the midline posterior to the orbits. The full extent of frontals and parietals is unclear, but the frontals comprise a relatively greater part of the skull roof than in many bothremydids (e.g., Bothremys, Chedighaii; Fig. 5). Subtle lines converge posteriorly on the parietal part of the skull roof; these probably resulted from postmortem compression of the parietals over the braincase. There is a distinct sagittally oriented supraoccipital plate; however, it is unclear whether or not the supraoccipital extended onto the skull roof. The maxillae are oriented vertically and do not extend much dorsally onto the skull roof, a condition also unlike typical bothremydids where there is little change in slope from sides to the top of the skull. The depth of the maxillae beneath the orbits is relatively shallow. There are rugose projections of the maxillae near the anterolateral borders of orbits. These maxillary excrescences have not been described in any other turtle. Although these may have been accentuated by dorsoventral compression of the skull, they appear to be natural features, with texture that differs from the surrounding bone. Because the maxillary excrescences are associated with the anterior edge of the orbit, a possible interpretation is that they are related to the salt-excreting glands found in many marine reptiles (e.g., Fernandez and Gasparini, 2008). The triturating surface is very narrow, and there is no distinct lingual ridge. There is no bothremydid pit on the triturating surfaces, and jugals do not appear to be exposed on the palate. Fragments of the palatines are exposed, showing the posterior borders of the internal nares. The pterygoid trochlear process appears to be well developed, but the pterygoid fossa is very shallow and partly obscured by poor preservation and matrix. The occipital plate is very narrow dorsoventrally, but its other features are obscured. The maxilla and quadrate are not in contact; the quadratojugal intervenes between them, and there is only a slight cheek emargination. The limits of the jugal are evident on the right side; it is roughly equidimensional and comprises only a small part of the posterolateral border of the orbit. Similarly, the quadratojugal appears to have limited contact with the jugal and does not extend signicantly posterodorsally over the quadrate. The temporal fossa is narrow dorsoventrally, and the temporal emargination is slight. The dorsal border of the temporal fossa reaches its anterior-most extent along its lateral edge (e.g., as in Foxemys) rather than medially as is usual in bothremydids. The lateral border of the temporal fossa does not extend onto the squamosal and quadrate, and the quadratojugal appears to barely reach its edge. The postorbital is relatively long; it comprises a small part of the posterior border of the orbit and extends to the lateral border of the temporal fossa. The postorbital suture with the parietal is evident on both sides of the skull.

Type SpeciesChupacabrachelys complexus, n. sp., by monotypy. DiagnosisAs for the species, by monotypy. EtymologyThe chupacabra (Spanish for goat sucker) is a mythical creature in contemporary Mexican-American legend said to feed on livestock in the border region of Texas and Mexico (e.g., Brown, 2008). The skull of Chupacabrachelys resembles that of a mangy coyote believed to be responsible for chupacabra sightings in South Texas during 2008. CHUPACABRACHELYS COMPLEXUS, new species DiagnosisA large bothremydid pleurodire of the tribe Taphrosphyini with the following unique features: skull and lower jaw very narrow (width/length = 0.63), compressed (height/length = 0.36) and triangular (jaw rami meet at 40 ), coronoid process posterior to midlength of jaw, orbits elongate anteroposteiorly with narrow extensions along maxilla-prefrontal sutures, rugose maxillary projections at anterolateral corners of orbits, low temporal emargination, weak posteroventral ange on squamosal, weak lateral extension of squamosal along quadrate suture, posteroventral knob on opisthotic; and differing from Labrostochelys in having a broader prefrontal, shorter projection of the premaxilla anterior to the labial ridge, external narial opening incompletely divided by the prefrontal and premaxilla, larger orbit, shorter posterior extension of the squamosal, weak lateral tubercle on the squamosal, mandibular condyle only slightly anterior to the occipital condyle, and more prominent pterygoid trochlear process. EtymologyIn recognition of The Complex tour performance of the Blue Man Group, which provided the authors with many hours of entertainment during collection and preparation of the type and referred specimens. Type SpecimenTMM 456061 consists of skull, lower jaw, a nearly complete shell (missing anterior end of nuchal and rst through third peripherals on the right side), atlas and parts of three cervical vertebrae, rst through fth thoracic vertebrae, 10 caudal vertebrae, both scapulae and coracoids, right humerus, both ulnae, right radius, three metacarpals, both innominates, six metatarsals, right astragalus, seven phalanges (limb and position uncertain), and eight ungual phalanges. Referred SpecimenA complete shell (TMM 45856-1) with right and left innominates in articulation, left coracoid, left humerus, metatarsal, and phalanx. Cranial Skeleton SkullThe skull is nearly complete; only the left quadratojugal and part of the right quadrate are missing, but sediment matrix obscures most of the palate and braincase. The preservation is insufcient to reveal many details of the cranial sutures, but a few are apparent (Fig. 4). The entire skull is mildly compressed dorsoventrally. As a result, the posterodorsal ends of the maxillae project slightly above the corners of the orbits (apparently an unnatural condition) and the left orbit is fractured along its posterior border. The left maxilla and premaxilla are slightly dislocated and tilted ventromedially. However, the postmortem compression is mild and does not distort the basic features of the skull. Its general form is very narrow and triangular, unusual for a bothremydid. The preorbital part of the skull is particularly narrow. In basic proportions, Chupacabrachelys is closest among bothremydids to Labrostochelys in having a low width to length ratio (0.63; lowest among bothremydids), and a low height to length ratio (0.36; lowest among bothremydids). The preorbital length to total length ratio (0.28) is also relatively high, and the temporal emargination is reduced, as in Taphrosphyini (Fig. 5). The depth of the cheek emargination is low as in most bothremydids, and particularly so compared to most other pelomedusoids.

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FIGURE 5. Comparison of selected Bothremydidae skulls in dorsal view, standardized to comparable length. Scales on right show four ratios of standard skull measurements for bothremydids: Araiochelys hirayamai (Ah), Bothremys cooki (Bc), Bothremys maghrebiana (Bm), Chupacabrachelys complexus, n. gen. n. sp. ( Cc), Cearachelys placidoi (Cp), Foxemys mechinorum (Fm), Kurmademys kallamedensis (Kk), Labrostochelys galkini (Lg), Rhothonemys brinkmani (Rb), Taphrosphys congolensis (Tc), and Taphrosphys ippolitoi (Ti). Markers on scales for Taphrosphyini are shown in gray, skull reconstructions are modied from Gaffney et al. (2006), and ratios are based on measurements given by Gaffney et al. (2006).

The squamosals form relatively short, horn-like processes that extend posterior to the opisthotics. There appears to be a weak posteroventral ange on the squamosal hornmore subtle than in other Taphrosphyini (e.g., Labrostochelys)and the posterior process of the opisthotic has a marked ventral knob as in species of Taphrosphys (T. sulcatus, T. ippolitoi, and T congolensis; Gaffney et al., 2006). The borders of the quadrate are well dened, but the interior of the tympanic cavity is obscured by sediment matrix. The tympanic cavity is open and not highly restricted posteriorly. The mandibular condyle lies anterior to the occipital condyle. Lower JawThe right ramus of the lower jaw is complete, but the postdentary elements are missing on the left side (Fig. 4). The general form of the jaw is narrow and triangular, with rami meeting at an angle of 40 (lines drawn along the midline of triturating surfaces). This basic form differs substantially from the broad Ushaped jaw typical of Bothremydini (64 to 87 ; Fig. 6). The narrow, triangular form is generally similar to that in some Taphrosphyini. For example, the jaws in Rhothonemys brinkmani and Taphrosphys congolensis (Gaffney et al., 2006) are narrow and triangular; however, in both species the rami diverge at a greater angle (85 and 60 , respectively). The lower jaw of Labrostochelys

is unknown, but based on the skull it was likely very narrow and triangular (47 ). In Chupacabrachelys, the coronoid process is very low, such that the entire jaw has a low prole in lateral view (height at coronoid process/length of jaw = 0.19). The low prole differs from typical Bothremydini (height/length range = 0.26 to 0.35), but is comparable to some Taphrosphyini such as Rhothonemys brinkmani (0.21) and Taphrosphys congolensis (0.14; Gaffney et al., 2006). The anterior part of the jaw is proportionally longer in Chupacabrachelys (length anterior to coronoid process/total length of jaw = 0.6) than in other bothremydids (range = 0.38 to 0.58). Consequently, the coronoid process is situated posterior to the midpoint of the jaw, and the posterior half is short, not expanded as in typical Bothremydini (Fig. 6). The dentaries in Chupacabrachelys are fused along an extended symphysis with no suture visible and no projection at the tip (Fig. 4). The triturating surface is narrow with no symphyseal ridge on the midline, and remains constant in width posteriorly. The lingual and labial ridges are roughly parallel and of similar height, but the labial ridge fades posteriorly and the lingual ridge rises above the labial ridge approaching the coronoid process. The lingual ridges meet to enclose a sharp V-shaped symphyseal trough (not an elevated U-shaped symphyseal wedge sensu

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FIGURE 6. Comparison of selected Bothremydidae lower jaws in lateral and dorsal views, standardized to comparable length with triturating surfaces (gray) and mandibular articulation (cross-hatched) highlighted. Scales on right show four ratios of standard jaw measurements and variation in angle between jaw rami for bothremydids: Araiochelys hirayamai (Ah), Bothremys cooki (Bc), Bothremys maghrebiana (Bm), Bothremys sp. FMNH PR247 (Bs), Bothremydini indet. AMNH 29989 (Bi), Chendighaii barberi (Cb), Chupacabrachelys complexus, n. gen. n. sp. ( Cc), Cearachelys placidoi (Cp), Foxemys mechinorum (Fm), Kurmademys kallamedensis (Kk), Labrostochelys galkini (Lg), Rhothonemys brinkmani (Rb), Taphrosphys congolensis (Tc), and Taphrosphys ippolitoi (Ti). Markers on scales for Taphrosphyini are shown in gray, jaw reconstructions are modied from Gaffney et al. (2006), and ratios are based on measurements given by Gaffney et al. (2006).

Gaffney and Forster, 2003). There are no triturating pits or posterior expansions of the triturating surface as in typical Bothremydini. The contacts of the dentary with postdentary elements are difcult to delineate; the sutures between angular, surangular, and articular appear to be completely fused. The dentaries are widely exposed on the lateral surface of the jaw, relative to the coronoid and surangular. Preservation is inadequate to reveal the presence or location of the foramen nervi auriculotemporalis. The sulcus cartilaginis meckelii is a distinct groove reaching anteriorly almost to the symphysis. The prearticular and angular are separated by a narrow opening that extends posteriorly from the sulcus cartilaginis meckelii (Fig. 4), and the two elements do not share a contact. Although this separation appears to be natural, it might be due to postmortem compression; the angular is ared outward on the lateral side of the jaw. The splenial is absent. The dorsal opening of fossa meckelii is a narrow slit, poorly exposed in dorsal view. There is a low midline longitudinal ridge on the mandibular articulation surface. The retroarticular process is well developed and posteriorly directed (not ventrally directed as in

Podocnemididae), and has a small foramen (foramen posterius chorda tympani) on its dorsal surface. Postcranial Skeleton Cervical VertebraeParts of four cervical vertebrae are preserved (Fig. 7). These are (1) the atlas, lacking the right half of the neural arch; (2) the centrum of a mid cervical; (3) a complete posterior cervical (centrum and neural arch were preserved separately, but the two t together and appear to comprise a single vertebra); and (4) the centrum of the eighth cervical (preserved separately from the neural arch of the eighth, which remains in articulation with the carapace). The atlas centrum is amphicoelus and appears to be fused with the intercentrum (Fig. 7A). A remnant notochordal pit is present on the posterior face of the centrum. There is a thin keel on the ventral surface, expanded to form a nub on the posterior end. Only the left half of the neural arch is preserved. Its anterior end forms part of the articular facet for the occipital condyle. The lateral surface appears to have a remnant articulation surface for

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FIGURE 7. Chupacabrachelys complexus, n. gen. n. sp., vertebrae of TMM 456061; atlas in A, posterior, B, left lateral, and C, ventral views; centrum of ?fourth cervical in D, posterior, E, left lateral, and F, ventral views; ?seventh cervical in G, posterior, H, left lateral, and I, ventral views; centrum of eighth cervical in J, posterior, K, left lateral, and L, ventral views; anterior caudal vertebra in M, left lateral and N, ventral views. Inset line diagrams show the atlas, seventh cervical, and caudal vertebrae of Podocnemis expansa (modied from Williams, 1950; Gaffney, 1990) reduced to common height.

an atlantal rib. The posterior end bears a small, medially directed postzygopophysis. The centra for the mid and posterior cervicals are all procoelous, taller than wide, and compare well in general form with those of Podocnemis expansa (Williams, 1950). None of the articulation surfaces of the centra appear to have saddle joints but all have a strong ventral keel expanded at anterior and posterior ends to form an accessory ball-socket articulation, similar to those in Podocnemis expansa (Williams, 1950). The posterior articulation of the eighth cervical centrum is more expanded laterally than the others (Fig. 7L). The anteroventral rim of each centrum bears low parapophyseal protuberances. All of the centra have oval depressions on their lateral surfaces. The diapophyses are comprised of both centrum and neural arch, and are relatively short compared to Podocnemis expansa (Williams, 1950). The neural arch of the complete posterior cervical (Fig. 7G) is relatively taller than in Podocnemis expansa, but the postzygapophyses are elevated, close together at the posterior end of the neural spine, and separated by a deep pit, as in the seventh cervical of Podocnemis expansa (Williams, 1950). Thoracic VertebraeThe rst two thoracic vertebrae are preserved in articulation with the carapace, the third is preserved separately, the fourth and fth are in articulation, and the remainder are not preserved. All of the thoracic vertebrae have at articulation surfaces and compressed hourglass-shaped centra. The anterior end of the rst thoracic centrum is more expanded laterally for articulation with the rib heads than any of the others.

The fourth and fth centra are twisted as a result of a pathological shell abnormality (doubling of the fth costal on the left side). Caudal VertebraeParts of at least 10 caudal vertebrae are preserved; all have procoelus articulations. The proximal caudal shown here (Fig. 7M) is typical of the entire series, and much like the fth caudal of Podocnemis expansa illustrated by Gaffney (1990). ScapulaThe scapula (Fig. 8) matches the description for Taphrosphys sulcatus (as given by Gaffney, 1975) with a longer cylindrical dorsal process, slightly curved medially, and shorter ventral (acromial) process tapering distally. The angle of 92 between the two processes is closer to Taphrosphys (98 ) than Podocnemis (77 given by Gaffney, 1975). The left scapula was preserved in articulation or nearly so with the rst thoracic rib on costal 1 (Fig. 2). CoracoidThe coracoid (Fig. 8) is almost perfectly intermediate in form between those of Podocnemis expansa and Taphrosphys sulcatus (as compared by Gaffney, 1975). The shaft is broad and C-shaped in cross-section, and relatively short (shorter than in Podocnemis, but not as short as in Taphrosphys), broad and curved (more so than in Podocnemis, but not as curved as in Taphrosphys), and with a large scapular articulation (larger than in Podocnemis, but not as large as in Taphrosphys). The coracoid in the referred specimen (TMM 45856-1) is identical to that of the holotype. HumerusThe humerus (Fig. 8) is similar to that of Podocnemis expansa (illustrated by Gaffney, 1990), Bothremys barberi

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FIGURE 8. Chupacabrachelys complexus, n. gen. n. sp., forelimb elements of TMM 456061; left coracoid in A, lateral, B, ventral, and C, dorsal views; left scapula in D, anterior and E, lateral views; right humerus in F, dorsal, G, ventral, and H, anterior views; right radius in I, dorsal, J, ventral (showing proximal articulation surface), and K, medial views; left ulna in L, dorsal (showing proximal articulation surface), M, ventral, and N, medial views. Inset diagrams compare humerus, coracoid, articulated ulna and radius of Chupacabrachelys complexus with Taphrosphys sulcatus (modied from Gaffney, 1975), Chendighaii barberi (from Zangerl, 1948), and Podocnemis expansa (from Gaffney, 1990) reduced to common length. Abbreviations: acr, acromion process; bic, bicipital tendon attachment; cor, articulation for coracoid; ect, ectepicondylar foramen; gle, glenoid; int, articulation for intermedium; lat, lateral process; lig, radio-ulnar ligament attachment; med, medial process; ole, olecranon process; sca, articulation for scapula; scp, scapular spine; uln, articulation for ulnare

(CNHM P27405; illustrated by Zangerl, 1948), and Taphrosphys sulcatus (YPM PU 18707; illustrated by Gaffney, 1975). Its features are intermediate between the slender form with a short medial process and narrow distal end observed in Podocnemis and Bothremys, and the more robust form, with a wide medial process and wide distal end found in Taphrosphys. Although the distal end appears to be more expanded than in Bothremys, the ratio of distal width to total length is 0.3 (as in Podocnemis; Gaffney, 1975, and in Bothremysbased on illustration given by Zangerl, 1948) but not as wide as in Taphrosphys (0.47). Other differences are also subtle. The head is relatively small and more in line with the shaft (as in Bothremys), rather than large and set out at a right angle to the shaft (as in Taphrosphys and Podocnemis). The val-

ley between medial and lateral processes on the ventral side is not as deep as in Taphrosphys and Podocnemis. The acute ridge extending from the base of the medial process to capitellem on the ventral side of shaft noted by Zangerl (1948) in Bothremys is lacking. Although the area around the ectepicondylar foramen is broken, it is better preserved in the humerus found with the referred specimen (TMM 45856-1). In both cases, the foramen appears to be a more fully enclosed groove, rather than the shallow groove in Bothremys, but not as deep as in Podocnemis. There are slightly depressed, ovoid muscle insertion scars on the humeral shaft at the base of the deltopectoral crest (for deltoideus), and at the base of the lateral process adjacent to the head (for latissimus dorsi).

LEHMAN AND WICKBOTHREMYDID TURTLE FROM TEXAS Ulna and RadiusBoth the ulna and radius are similar in form to Podocnemis expansa (illustrated by Gaffney, 1990), although relatively shorter and with more expanded distal ends (Fig. 8). The proximal end of the radius has a cup-shaped humeral articulation surface, slightly larger in diameter than the shaft. There is a prominent attachment site for the superior radio-ulnar ligament just below the humeral articulation (Gaffney, 1990). The proximal end of the ulna has a very shallow sigmoid notch and olecranon extending only slightly above the humeral articulation. The bicipital tubercle (Walker, 1973) lies close to the humeral articulation surface. The broad carpal articulation surface has two distinct faces, one directed medially for the intermedium, the other distally for the ulnare. The broad and attened distal expansions of radius and ulna suggest that Chupacabrachelys may have had a webbed manus as in some extant and fossil pelomedusoids (e.g., Fielding et al., 2005). However, only a few metacarpals are preserved, and these are not elongate compared to Podocnemis expansa. Tarsus and MetatarsalsThe preserved parts of the tarsus and pes are remarkably similar in form to those of Podocnemis expansa (illustrated by Gaffney, 1990). Elements of the right hind foot were found disarticulated but closely associated (Fig. 9). The astragalus has a complex, proximal saddle-shaped articulation surface and a gently convex distal surface. There are several isolated distal tarsals, one of which may be distal tarsal 4. This tarsal is cylindrical in form with articulation facets on four sides, and a shallow concavity on the presumed anterior surface. Metatarsals I through IV, with placement interpreted by gradual increase in length and similarity in form with those of Podocnemis expansa (Gaffney, 1990), have curved shafts and asymmetric proximal ends that would overlap in articulation. The distal articulation surfaces are well developed on metatarsals I through III, but less distinct on IV. Metatarsal V is a square plate with a distinct articulation facet for distal tarsal 4 and a prominent knob for articulation with phalanx V-1. Of the isolated phalanges, only phalanx V-1 can be identied; it is comparable in length to metatarsal IV but slender and curvedvery much like that of Podocnemis expansa (Gaffney, 1990). UngualsThe ungual phalanges are broad, thin, and paddleshaped as in Taphrosphys sulcatus (Gaffney, 1975). All were preserved separately and cannot be condently assigned to fore- or hind limb or digit; however, examples span a range in size and shape (Fig. 9). Some are relatively narrow and slightly constricted below the proximal articulation surface; others are broad and more constricted at the articulation surface. Based on the similar arrow-shaped unguals in Araripemys barretoi (Meylan, 1996), the larger broad unguals likely belong to digit I, II, or III, whereas the smaller narrow ones are from digit IV or V. The wide and shallow articulation surfaces suggest that the unguals were capable of only a limited range of exion. PelvisBoth innominates are preserved in TMM 456061, the left side completely (although the ischium was found detached). Both are also preserved in articulation in the referred specimen (TMM 45856-1), but dorsoventrally crushed. The general form of the pelvis is very much like Podocnemis expansa (illustrated by Gaffney, 1990). The ilium and pubis are united, with only a faint suture visible between them within the acetabulum (Fig. 10). The prepubis appears to be shorter than in Podocnemis expansa, but the end is missing or unnished. However, the prepubis is completely preserved in the companion specimen. It is a short remnant triangular process as restored here (Fig. 10), and as in Bothremys barberi (FMNH P27370; Zangerl, 1948). The prepubic processes did not meet on the midline, unless they were extended in cartilage. Similarly, the posteroventral part of the ischium is expanded to form an arcuate articulation with the plastron. In the referred specimen (TMM 45856-1), the plastral articulation of the ischium extends anteromedially, but does not meet the opposite ischium along the midline. The iliac surface for articulation with

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FIGURE 9. Chupacabrachelys complexus, n. gen. n. sp., hind foot elements of TMM 456061 in dorsal view; A, right metatarsal V, B, distal tarsal ?IV, C, right astragalus, D, right phalanx V-1, E, left metatarsal IV, F, left metatarsal III, G, left metatarsal I, H, several ungual phalanges showing variation in form. Inset diagram shows reconstruction of right hind foot in dorsal view, based on Podocnemis expansa (modied from Gaffney, 1990). On metatarsal V articulations are shown for distal tarsal IV (dt IV), metatarsal IV (mt IV), and phalanx V-1 (p V-1).

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FIGURE 10. Chupacabrachelys complexus, n. gen. n. sp., left innominate of TMM 456061 in A, lateral, B, anterior, and C, posterior views with ischium removed; ischium in D, posterior and E, anterior views. Inset diagram compares outline of articulation surface of right ilium with carapace (arrow indicates midline, anterior to top) in Chupacabrachelys complexus, Chendighaii barberi (CNHM P27370; from Zangerl, 1948), and Podocnemis expansa (AMNH 62947; from Gaffney, 1990). Abbreviations: act, acetabulum; il, articulation for ilium; isc, articulation for ischium; pub, articulation for pubis.

the carapace is narrow, and does not extend medially as far as in Podocnemis expansa (Gaffney, 1990). This articulation surface completely lacks the medial extension that overlaps the suprapygal. The iliac articulation in Bothremys barberi has a broad medial expansion; however in some specimens (e.g., FMNH P27370, illustrated in articulation with the carapace by Zangerl, 1948), the dorsal process of the ilium does not overlap the suprapygal, whereas in others it does (FMNH P27372; Zangerl, 1948). CarapaceThe carapace in TMM 456061 is almost complete, missing only part of the anterior margin (most of the nuchal) and the edge of the right side (rst through fourth peripherals). The carapace was preserved in two parts (anterior and posterior) with third costals and bridge peripherals loose (Fig. 11). The two parts are uncrushed and preserve their natural curvature, and when joined together require that the anterior end tilt downward at an angle of about 50 relative to the rear (Fig. 11). The entire carapace is about as wide as it is long. When in articulation with the plastron, there is a much wider gape at the front than at the back of the shell. The general form of the carapace, and the number and relationships of each of its elements, is nearly identical to that in Bothremys barberi (Gaffney et al., 2006) even in subtleties (e.g., the slight inection of the posterior margin of the carapace at the ninth peripheral, although this is at the tenth peripheral in Bothremys). The form and positions of axillary and inguinal buttress scars also mirror Bothremys barberi (Gaffney et al., 2006). The rst neural is rectangular and only slightly longer than second; the eighth and ninth peripherals are largest, with tenth and eleventh much smaller. There are six neurals; although the sixth is somewhat asymmetric in TMM 456061, this is likely the result of abnormal doubling of the fth costal on the left side (this also produced an abnormal doubling of the inguinal buttress on that side). Similar shell abnormalities have been described in some specimens assigned to Bothremys (e.g., a small anomalous bone interceding between mesoplastron and hyoplastron in ANSP 15902; Gaffney and Zangerl, 1968) and are common in turtles generally (Zangerl, 1969). The sixth through eighth costals meet on the midline. The iliac scar overlaps most of the boundary between seventh and eighth costals, but does not extend at all onto the suprapy-

gal. This can be conrmed by articulation with the pelvis (also preserved; Fig. 10) and for both the type and referred specimens. This slight difference in the iliac scar, and the greater declination of the anterior carapace margin allowing the plastron to project slightly anterior of the carapace, are subtle differences between the shells of Chupacabrachelys and Bothremys barberi, but the extent of individual variation in these features is unknown. In TMM 456061, the fourth and fth peripherals lack a lateral ange; ventral and dorsal faces meet at a rounded obtuse angle, very much like those described in the same peripherals in some specimens of Taphrosphys sulcatus (e.g., YPM PU 18707 described by Gaffney, 1975). Although the ventral and dorsal plates of these peripherals meet at a more acute angle in the referred specimen (TMM 45856-1), this may be another minor point of difference with the carapace in Bothremys barberi. A deep and sharply dened nuchal notch is present in the referred specimen, but this area is not preserved in the holotype. The depth of the nuchal notch varies among specimens assigned to Bothremys; it is very subtle and narrow in YPM 3608 but broad and pronounced in FMNH PR 247. The scale sulci are not well preserved, but appear also to be generally consistent with Bothremys barberi (Gaffney et al., 2006). The form of the rst vertebral scale is evident in the referred specimen (TMM 45856-1) and parts of the second through fth can be delineated in the type specimen. There is an asymmetric scale that intercedes between the fourth and fth vertebrals in the type; however, this may be a result of the abnormal doubling of the fth costal on the left side (Fig. 11). Comparable abnormalities in scale pattern are observed in some specimens assigned to Bothremys (e.g., an anomalous scale interceding between the rst vertebral and rst marginals in YPM 3608; Gaffney and Zangerl, 1968). Sulci for the marginal scales are visible along the posterior part of the carapace in both type and referred specimens, and are restricted to the peripherals. There are two shallow oval pits (likely parasite attachment scars) on the visceral side of the pygal and suprapygal in TMM 456061 (Fig. 11). Zangerl (1948) described similar holes in a carapace of Podocnemis alabamae (= Bothremys barberi; FMNH P 27372) that he attributed to bone erosion as a result of parasitic infection.

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FIGURE 11. Chupacabrachelys complexus, n. gen. n. sp., preserved elements of shell in TMM 456061 with carapace in A, dorsal and B, visceral views showing cross-sections of bridge peripherals; and plastron in C, ventral and D, visceral views. Inset diagrams show anterior lobe of plastron in E, TMM 456061 and F, TMM 45856-1 compared with several specimens of Chendighaii barberi (from Zangerl, 1948; Gaffney and Zangerl, 1968); restoration of shell in G, dorsal, H, ventral, and I, right lateral views, based on both TMM 456061 and 45856-1, showing sutures and scute sulci observed on both specimens (with mirror image); abnormalities and asymmetry removed.

The ornamentation of the carapace consists of the typical pelomedusoid pattern of ne reticulate and discontinuous branching polygonal grooves. The ne grooves are not deeply incised (as in some specimens of Taphrosphys; Gaffney, 1975) and are only obvious in oblique illumination. In the larger referred specimen (TMM 45856-1), the ornamentation is more sharply dened.

PlastronThe plastron is shorter than the carapace; however, when the carapace is restored to its natural curvature, the plastron extends slightly anterior to the front edge of the carapace (Fig. 11). The form of the plastron, the proportions and relationships of each of its elements, are nearly identical to that in Bothremys barberi (Gaffney et al., 2006). The anterior edge is semicircular; the bridge is longer than the anterior lobe and shorter

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JOURNAL OF VERTEBRATE PALEONTOLOGY, VOL. 30, NO. 6, 2010 physis. The shell in Chupacabrachelys resembles their Atlantic Coast subspecies (e.g., ANSP 15092), which has bridge peripherals with ventral plates about half as long as the dorsal plates, and a thick ventral ridge parallel to the posterior edge of the carapace. However, these features were not considered signicant when Gaffney et al. (2006) later tabulated shell criteria for nine specimens of Bothremys and Chedighaii. Gaffney et al. (2009) found that a shell condently attributed to Chedighaii (ALAB PV 2001.2) has a plastron with a straight anterior margin, swollen epiplastra, and a wide entoplastron with a straight posterior margin. Other shells (FMNH P27369, YPM 3608) share these features but were not preserved with diagnostic cranial material. In contrast, a shell condently attributed to Bothremys (FMNH PR 247) has a semicircular anterior plastron margin, thin epiplastra, and an entoplastron with a pointed posterior margin. Some isolated shells (FMNH 26055, 27370) also share these features. However, the holotype of Chupacabrachelys exhibits the smoothly rounded anterior plastron margin thought to be diagnostic of Bothremys, whereas the larger referred specimen exhibits the truncated plastron margin with swollen epiplastra thought to be diagnostic of Chedighaii (Fig. 11). This disparity lends credence to Zangerls (1948) notion that the latter condition may be attributable to advanced age. With the specimens presently available, it may not be possible to distinguish ontogenetic and individual variation from features having taxonomic signicance. Only the form of the entoplastron remains a possible discriminating feature; the entoplastron is relatively long and narrow in Chupacabrachelys, broad but posteriorly pointed in Bothremys, and abruptly truncated posteriorly in Chedighaii. DISCUSSION Gaffney et al. (2006) recently conducted an exhaustive phylogenetic analysis of side-necked turtles, based on 175 characters in more than 40 taxa, and focused particularly on bothremydids. Their analysis provides a comprehensive framework in which to evaluate the likely relationships of Chupacabrachelys, and as a result a similar investigation is unnecessary here. Moreover, their analysis makes it clear that Chupacabrachelys is a member of Bothremydidae, but its further relationships will remain unclear due to obscuration of several key characters (see below). Chupacabrachelys exhibits several features considered synapomorphic for Pleurodira in most phylogenetic analyses (e.g., Gaffney et al., 2006). These include the pterygoid trochlear process, the expanded dorsal process of the ilium situated close to the shell midline and in sutural articulation with the costals, the sutural articulation of the pubis and ischium with the xiphiplastron, and the anal notch in the plastron. Additional characters are shared with Eupleurodira: procoelous caudal articulations, neurals posterior to second are hexagonal in shape with anterolateral contacts shorter than posterolateral contacts, equidimensional mesoplastra not meeting on midline, axillary process of hyoplastron reaches third peripheral, elevated cervical postzygapophyses, and with Pelomedusoides: nasals absent, prefrontals meeting on midline, splenial absent, cervical vertebrae procoelous, anterior plastral lobe reaches anterior carapace margin. Collectively, these features indicate that Chupacabrachelys is a pelomedusoid pleurodire. Among pelomedusoids, Chupacabrachelys has clear afnities with family Bothremydidae. In their revised diagnosis for the family, Gaffney et al. (2006) relied exclusively on cranial characters; of these, several diagnostic bothremydid characters are evident in Chupacabrachelys, including (1) a maxilla-quadratojugal contact is present, in contrast to all other pleurodires (except Taphrosphyini); (2) the paroccipital process does not project posterior to the squamosal, in contrast to all other Pelomedusoides; and (3) the lingual ridge on the lower jaw is relatively high.

than the posterior lobe, with lateral margins tapering posteriorly; the anal notch is deep and U-shaped. Sutures for the mesoplastra are not visible in the holotype (obscured by a thin crust resistant to preparation), but they are well dened in the referred specimen (TMM 45856-1). The entoplastron is longer than wide in the type (as in some specimens of Bothremys barberie.g., ANSP 15902; Gaffney et al., 2006where in most cases the entoplastron is wider than long), but in the referred specimen (TMM 45856-1) the entoplastron is broader. The entoplastron in the companion specimen also has a medial ridge on its visceral surface (smooth in the type specimen). The epiplastra have slight bulges on their visceral surface; these are much more pronounced on the referred specimen, which also has a midline notch between the epiplastra (Fig. 11). The lateral margins of the posterior lobe have a slightly thickened lip that extends back from the inguinal buttresses. The pubic scar is oval; the ischiac scar is triangular and well forward of the posterior margin of the plastron. The right and left ischiac scars are joined by a low transverse ridge parallel to the border of the anal notch. The scale sulci are only partly visible in both type and referred specimens, but appear to also be like those in Bothremys barberi. The intergular scale extends onto the anterior part of the entoplastron, whereas the gular scale is small, triangular, and restricted to the epiplastron. The sulcus between humeral and pectoral scales is posterior to the epi-hyoplastral suture. The sulcus between pectoral and abdominal scales crosses the anterior part of the mesoplastron and posterior part of the hyoplastron, whereas the sulcus between abdominal and femoral scales crosses the anterior part of the hypoplastron, and the sulcus between femoral and anal scales crosses the xiphiplastron at a slight angle. Preservation is insufcient in both specimens to determine whether the pectoral scale reached the entoplastron. There is no obvious ornamentation on the external surface of the plastron. However, on the larger referred specimen (TMM 45856-1), the same pattern typical of the carapace extends onto the bridge and along the margins of anterior and posterior lobes of the plastron. Shell ProportionsThe shells of Bothremys and Chedighaii are strikingly similar to each other, and a considerable effort has been made to distinguish the two (Gaffney et al., 2006; 2009). The shell of Chupacabrachelys resembles both of these, and because isolated shells are commonly preserved without diagnostic cranial material, it is appropriate to reexamine shell features that may distinguish them. Zangerl (1948) provided a complete set of over 80 shell measurements for each of ve specimens later referred to Bothremys and Chedighaii (Gaffney et al., 2006; 2009). Zangerl (1948) standardized the shell measurements relative to the width of the nuchal for each specimen (measurement of character 100/maximum width of nuchal). A comparable set of measurements is given here (Appendix 1) for the type and referred specimens of Chupacabrachelys complexus, and standard ratios can be calculated in the same way. The referred specimen (TMM 458561) is slightly larger than any of the ve measured by Zangerl (1948) but is similar in size to several described later (ANSP 15902, Gaffney and Zangerl, 1968; ALAB PV2001.2, Gaffney et al., 2009). Nearly all of the 80 shell indices for both specimens of Chupacabrachelys, standardized to nuchal width, fall within the range reported for specimens measured by Zangerl (1948). Only two proportional features evident in both the type and referred specimens fall outside the range reported for Bothremys and Chedighaii shells. Standardized to nuchal width, these are (1) a relatively narrow entoplastron, and (2) a relatively deeper anal notch (length of posterior wings as given by Zangerl, 1948). Gaffney and Zangerl (1968) recognized three subtly different shell types, at that time hypothesized to represent Atlantic Coastal Plain, Gulf Coast, and Western Interior subspecies of Bothremys, and distinguished primarily on the basis of the relative lengths of the nuchal, mesoplastron, and epiplastral sym-

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LEHMAN AND WICKBOTHREMYDID TURTLE FROM TEXAS In addition, the neural series is reduced, extending only to the sixth costals (as in the bothremydids Bothremys and Chedighaii). These features indicate that Chupacabrachelys is a member of family Bothremydidae. Gaffney et al. (2006) subdivided bothremydids into several more exclusive groups (subfamilies, infrafamilies, tribes, and subtribes), in part following Baur (1891) and reecting informal groups recognized by previous authors (e.g., Antunes and Broin, 1988). Several features identied in the phylogenetic analysis of Gaffney et al. (2006) that are present in Chupacabrachelys are considered synapomorphic for Bothremydodda (tribes Bothremydini + Taphrosphyini) or shared by most members of this group, including (1) an open fossa meckelli (prearticular-angular contact is short, or perhaps lacking in Chupacabrachelys); (2) premaxilla protruding anteriorly beyond the labial ridge in ventral view; (3) prefrontals projecting forward to form a midline process; (4) rst costal more than twice the anteroposterior length of the second costal; and (5) a short anterior plastral lobe (although also found in the podocnemidid Bairdemys; Gaffney et al., 2006). Features of Chupacabrachelys shared with tribe Bothremydini include (1) the relatively long postorbital (though this may be correlated with reduced temporal emargination); (2) a wide oval pubic scar and triangular ischiac scar inset from the xiphiplastral edge; (3) shell ornamentation of ne forking and irregular grooves; and possibly (4) the embayed anterior margin of the nuchal (clearly evident in the referred specimen of Chupacabrachelys, but not preserved in the holotype). However, Chupacabrachelys lacks typical features of the Bothremys group (subtribe Bothremydina of Gaffney et al., 2006), such as (1) a very broad preorbital part of skull, (2) a maxilla-quadrate contact, and (3) paired conical pits on maxillary and dentary triturating surfaces. Features of Chupacabrachelys shared with tribe Taphrosphyini include (1) the narrow preorbital part of skull, (2) reduced temporal emargination, and (3) the squamosal with weak posteroventral ange and a weak tubercle along its suture with the quadrate (both unique though much better developed in typical Taphrosphyini than in Chupacabrachelys). However, Chupacabrachelys retains a maxilla-quadratojugal contact (lacking in Taphrosphyini), and lacks the jugal-quadrate contact (present in Taphrosphyini; Gaffney et al., 2006). The placement of Chupacabrachelys within infrafamily Bothremydodda (Gaffney et al., 2006) is therefore supported by several unambiguous characters, but it is not possible to clearly resolve its position within this group. Chupacabrachelys could be a member of subtribe Bothremydina that secondarily lost most of the key Bothremys group cranial features and acquired several features similar to those of Taphrosphyini. In support of this hypothesis, some of the diagnostic Bothremydina cranial features are also lost in Chedighaii, which has a shell nearly identical to that of Chupacabrachelys. Alternatively, Chupacabrachelys could be a basal member of tribe Taphrosphyini that acquired several of the cranial features (squamosal posteroventral ange and tubercle) of that clade, but retained a plesiomorphic shell morphology like that in Bothremydini. This also seems reasonable, particularly because shell features for the entire tribe Taphrosphyini are known only for Taphrosphys. Moreover, the unusal triangular cranial morphology of Chupacabrachelys, atypical of bothremydids, is found elsewhere only in Taphrosphyini. Of these two possibilities, the most likely seems that Chupacabrachelys is a basal member of tribe Taphrosphyini. Another member of this tribe, Labrostochelys, has broadly similar cranial features to those of Chupacabrachelys, and its shell morphology is unknown. Compared to Labrostochelys, the prefrontal in Chupacabrachelys is shorter, the premaxilla does not project as far anterior to the labial ridge, the external narial open-

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ing is not completely divided by the prefrontal and premaxilla, the orbit is relatively much larger, the posterior extension of the squamosal is shorter, the lateral tubercle on the squamosal is small, the mandibular condyle is not as far anterior to the occipital condyle, and the pterygoid trochlear process is more prominent. If Chupacabrachelys is a member of Taphrosphyini, then several purported cranial apomorphies of the clade (presence of jugal-quadrate contact, absence of maxilla-quadratojugal contact) and some shell features (plastron with broad posterior lobe, long narrow pubic scar, round ischiac scar at edge of xiphiplastron, carapace with deeply furrowed irregular polygonal ornamentation, lacking nuchal notch) were only acquired by more derived members of this tribe. Further detailed preparation or non-invasive imaging of the tympanic cavity, palate, and braincase in Chupacabrachelys might help better elucidate its tribal afliation. CONCLUSIONS Gaffney et al. (2006:652) remarked that pleurodires had a more extensive and more complex evolutionary history than has been realized and that they exhibit a diversity of morphologies indicating a remarkable diversity of feeding and sensory strategies. The discovery of Chupacabrachelys provides additional evidence of this diversity, and of the dramatic Late Cretaceous evolutionary radiation of bothremydids in particular. Although Chupacabrachelys has several key synapomorphies of Bothremydidae, and with infrafamily Bothremydodda, its tribal afliation is uncertain, largely because it has not been possible to free the tympanic cavity, palate, and braincase of its sediment matrix. The most likely hypothesis is that Chupacabrachelys is a basal member of tribe Taphrosphyini. Its shell morphology is nearly identical to those of Bothremydini (e.g., Bothremys, Chedighaii), but its long narrow skull, with slender triturating surfaces, posteriorly extended squamosal horns with subtle ventral anges and lateral tubercles, is like those of Taphrosphyini (e.g., Labrostochelys). The unusual rugose maxillary excrescences have not been described in other turtles. Although it is tempting to interpret these features as related in some way to salt-excreting glands, it would seem that similar osteological features should be present in other turtles that inhabit marine environments. Prior to the monographic study of Gaffney et al. (2006:652), most studies of bothremydid turtles had been based largely on shells that among pleurodires in general, and in Pelomedusoides in particular, is relatively conservative morphologically, masking the magnitude of pleurodire diversity. Chupacabrachelys provides a stunning example of this phenomenon. Its shell is so nearly identical to those of Bothremys and Chedighaii, both of which occur in coeval strata, that had they been found together they would have been assigned to the same taxa with little doubt. Their skulls differ so dramatically it is hard to believe they belong to animals with such similar shells. However, no other turtle family exhibits such extreme morphological variation as is found in the skulls of bothremydids (Gaffney, et. al., 2006). Chupacabrachelys provides an additional example of this remarkable variation. The occurrence of Chupacabrachelys also offers further substantiation for evident latitudinal biogeographic differences between northern and southern parts of the western interior province during Campanian time (e.g., Lehman, 1997). Bothremydid turtles are abundant in Campanian paralic strata along the Atlantic and Gulf coastal plain (Schwimmer, 2002) and in Texas. A few specimens have been found in Kansas (Gaffney and Zangerl, 1968) and in New Mexico (Gaffney et al., 2006), but they are currently unknown farther north (Brinkman, 2003). Bothremydid turtles are also abundant and diverse in Campanian strata of southern Europe and northern Africa (Gaffney et al.,

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Gaffney, E. S., and Zangerl, R. 1968. A revision of the chelonian genus Bothremys (Pleurodira: Pelomedusidae). Fieldiana Geology, 16:193239. Gaffney, E. S., Hooks, G. E., and Schneider, V. P. 2009. New material of North American side-necked turtles (Pleurodira: Bothremydidae). American Museum of Natural History, Novitates 3655, 26. pp. Gaffney, E. S., Tong, H., and Meylan, P. A. 2006. Evolution of the sidenecked turtles: the families Bothremydidae, Euraxemydidae, and Araripemydidae. American Museum of Natural History, Bulletin 300, 698. pp. Gaffney, E. S., Campbell, K. E., and Wood, R. C. 1998. Pelomedusoid side-necked turtles from Late Miocene sediments in southwestern Amazonia. American Museum of Natural History, Novitates 3245, 11. pp. Lehman, T. M. 1985. Stratigraphy, sedimentology, and paleontology of Upper Cretaceous (Campanian-Maastrichtian) sedimentary rocks in Trans-Pecos Texas. Unpublished Ph.D. dissertation, University of Texas at Austin, Austin, Texas, 300. pp. Lehman, T. M. 1997. Late Campanian dinosaur biogeography in the Western Interior of North America; pp. 223240 in D. L. Wolberg, E. Stump, and G. D. Rosenberg (eds.), Dinofest International, Proceedings of a symposium held at Arizona State University. Academy of Natural Sciences, Philadelphia. Lehman, T. M., and A. B. Busbey. 2007. Big Bend Field Trip Field Guide. Society of Vertebrate Paleontology, Fall 2007 Field Trip, 69. pp. Lehman, T. M., and S. L. Tomlinson. 2004. Terlinguachelys schbecki, a new genus and species of sea turtle (Chelonoidea: Protostegidae) from the Upper Cretaceous of Texas. Journal of Paleontology 78:11631178. Linneaus, C. 1758. Systema Naturae, 10th edition, Volume 1. Stockholm, 824. pp. Meylan, P. A. 1996. Skeletal morphology and relationships of the Early Cretaceous side-necked turtle, Araripemys barretoi (Testudines: Pelomedusoides: Araripemydidae) from the Santana Formation of Brazil. Journal of Vertebrate Paleontology 16:2033. Rowe, T., R. L. Cifelli, T. M. Lehman, and A. Weil. 1992. The Campanian Terlingua local fauna, with a summary of other vertebrates from the Aguja Formation, Trans-Pecos Texas. Journal of Vertebrate Paleontology 12:472493. Schwimmer, D. R. 2002. King of the Crocodylians. Indiana University Press, Bloomington, Indiana, 220. pp. Tomlinson, S. L. 1997. Late Cretaceous and early Tertiary turtles from the Big Bend region, Brewster County, Texas. Unpublished Ph.D. dissertation, Texas Tech University, Lubbock, Texas, 194. pp. Waggoner, K. J. 2006. Sutural form and shell morphology of Placenticeras, and systematic descriptions of Late Cretaceous ammonites from the Big Bend region, Texas. Unpublished Ph.D. dissertation, Texas Tech University, Lubbock, Texas, 398. pp. Walker, W. F. 1973. The locomotor apparatus of Testudines; pp. 1100 in C. Gans (ed.), Biology of the Reptilia, Volume 4, Morphology D. Academic Press, New York. Williams, E. E. 1950. Variation and selection in the cervical central articulations of living turtles. American Museum of Natural History, Bulletin 94:511561. Zangerl, R. 1948. The vertebrate fauna of the Selma Formation of Alabama. Part II, the pleurodiran turtles. Fieldiana: Geology Memoirs 3:2356. Zangerl, R. 1969. The turtle shell; pp. 311339 in C. Gans (eds.), Biology of the Reptilia, Volume 1, Morphology A. Academic Press, London. Submitted July 22, 2009; accepted June 3, 2010.

2006), suggesting that their distribution was restricted to tropical waters, and did not extend to higher latitudes. ACKNOWLEDGMENTS The authors thank W. Langston, Jr., T. Rowe, and L. Murray of the Texas Memorial Museum for their assistance with specimens at the Vertebrate Paleontology Laboratory, D. Corrick and V. Davila of the Science and Resource Management staff of Big Bend National Park for their support of the authors eldwork in the Aguja Formation, and J. Wick and E. Lehman for their patient indulgence of the authors paleontological endeavors. E. Gaffney, F. OKeefe, and two anonymous reviewers provided helpful comments that substantially improved the manuscript. Illustrations are the work of T. Lehman. LITERATURE CITED
Antunes, M. T., and F. de Broin. 1988. Le Cretace terminal de Beira Littoral, Portugal: remarques stratigraphiques et ecologiques, etude complementaire de Rosasia soutoi (Chelonii, Bothremydidae). Ciencias da Terra 9:153200. Baur, G. 1891. Notes on some little known American fossil tortoises. Proceedings of the Academy of Natural Sciences of Philadelphia 43:411430. Befus, K. S., R. E. Hanson, T. M. Lehman, and W. R. Grifn. 2008. Cretaceous basaltic phreatomagmatic volcanism in West Texas: maar complex at Pena Mountain, Big Bend National Park. Journal of Volcanology and Geothermal Research 173:245264. Brinkman, D. B. 2003. A review of nonmarine turtles from the Late Cretaceous of Alberta. Canadian Journal of Earth Science 40:557 571. Brown, A. 2008. Haunted Texas: Ghosts and Strange Phenomena of the Lone Star State. Stackpole Books, Mechanicsburg, Pennsylvania, 122. pp. Cope, E. D. 1864. On the limits and relations of the Raniformes. Proceedings of the Academy of Natural Sciences of Philadelphia 16:181 183. Cope, E. D. 1868. On the origin of genera. Proceedings of the Academy of Natural Sciences of Philadelphia 20:242300. Fernandez, M., and Z. Gasparini. 2008. Salt glands in the Jurassic metriorhynchid Geosaurus: implications for the evolution of osmoregulation in Mesozoic marine crocodyliforms. Naturwissenschaften 95:7984. Fielding, S., Martill, D. M., and Naish, D. 2005. Solnhofen-style softtissue preservation in a new species of turtle from the Crato Formation (Early Cretaceous, Aptian) of northeast Brazil. Palaeontology, 48:13011310. Gaffney, E. S. 1975. A revision of the side-necked turtle Taphrosphys sulcatus (Leidy) from the Cretaceous of New Jersey. American Museum of Natural History, Novitates 2571, 24. pp. Gaffney, E. S. 1990. The comparative osteology of the Triassic turtle Proganochelys. American Museum of Natural History, Bulletin 194, 263. pp. Gaffney, E. S., and Forster, C. A. 2003. Side-necked turtle lower jaws (Podocnemididae, Bothremydidae) from the Late Cretaceous Maevarano Formation of Madagascar. American Museum of Natural History, Novitates 3397, 13. pp.

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APPENDIX 1. Measurements of the type and referred specimens of Chupacabrachelys complexus are given in millimeters. The skull and lower jaw measurements are taken in the manner of Gaffney et al. (2006:685, 690) and designated with the same capital letters. The shell measurements are given in the manner of Zangerl (1948:3638). , an approximate dimension due to distortion of the specimen or missing parts.

Skull (TMM 456061) Length, tip of snout to occipital condyle (A) Length, tip of snout to end of sagittal crest (I) Length, tip of snout to mandicular condyle (O) Height, mandibular articulation to skull roof (G) Height, occipital condyle to skull roof (K) Transverse width at cheeks (B) Transverse width at orbits (H) Minimum interorbital width (C) Anteroposterior length of orbit (D) Dorsoventral width of orbit (J) Transverse width of external nares (E) Lower jaw (TMM 456061) Sagittal length (B) Length of triturating surface (A) Width of ramus at coronoid process (C)
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Carapace (TMM 45856-1 and TMM 456061) 104 106 95 43 27 70 50 12 27 13 21 87 56 6 Sagittal length (along curve) Transverse width (at peripheral 7, along curve) Nuchal, sagittal length Neural 1, sagittal length Neural 2, sagittal length Neural 3, sagittal length Neural 4, sagittal length Neural 5, sagittal length Neural 6, sagittal length Suprapygal, length Pygal, length Nuchal, anterior width Neural 1, anterior width Neural 2, anterior width Neural 3, anterior width Neural 4, anterior width Neural 5, anterior width Neural 6, anterior width Pygal, anterior width Suprapygal, posterior width Pygal, posterior width Nuchal, maximum width Neural 1, maximum width Neural 2, maximum width Neural 3, maximum width Neural 4, maximum width Neural 5, maximum width Neural 6, maximum width Peripheral 1, inner length Peripheral 2, inner length Peripheral 3, inner length Peripheral 4, inner length Peripheral 5, inner length Peripheral 6, inner length Peripheral 7, inner length Peripheral 8, inner length Peripheral 9, inner length Peripheral 10, inner length Peripheral 11, inner length Peripheral 1, outer length Peripheral 2, outer length Peripheral 3, outer length Peripheral 4, outer length Peripheral 5, outer length Peripheral 6, outer length Peripheral 7, outer length Peripheral 8, outer length Peripheral 9, outer length Peripheral 10, outer length Peripheral 11, outer length Plastron (TMM 45856-1 and TMM 456061) Length, midline to anal notch Width anterior to bridge Width posterior to bridge Length of anterior lobe Length of posterior lobe on midline Length of epiplastral symphysis Length of entoplastron Length of hyoplastral symphysis Length of hypoplastral symphysis Length of xiphiplastral symphysis Depth of anal notch Width of entoplastron Length of hyo-hypoplastral suture Width of mesoplastron Length of mesoplastron Width of anal notch (between tips of xiphiplastra) 730 550 730 550 90 80 55 50 60 55 50 58 50 40 30 80 65 84 70 45 35 25 25 21 20 25 25 25 20 25 60 35 80 95 75 155 110 60 50 42 50 36 47 35 45 40 45 35 40 15 55 50 80 65 65 50 60 50 60 50 60 60 70 65 90 65 75 60 70 55 85 90 95 65 85 60 80 60 80 70 90 80 105 80 100 70 85 65 85 65 540 434 340 235 325 220 125 108 210 163 20 30 100 75 130 90 140 120 145 110 62 45 75 45 165 105 100 80 115 80