FROM the TITHONIAN (UPPER JURASSIC) of MEXICO MARIE-CE´ LINE BUCHY1 1Museo Del Desierto, Saltillo, Coahuila, Mexico, ,Mcbuchy@Gmail.Com

J. Paleont., 84(1), 2010, pp. 149–155 Copyright ’ 2010, The Paleontological Society 0022-3360/10/0084-0149$03.00

FIRST RECORD OF OPHTHALMOSAURUS (REPTILIA: ICHTHYOSAURIA) FROM THE TITHONIAN (UPPER JURASSIC) OF MEXICO MARIE-CE´ LINE BUCHY1 1Museo del Desierto, Saltillo, Coahuila, Mexico, ,[email protected].

INTRODUCTION paleobiogeographical and paleoecological implications of this ROM THE Middle Jurassic on, the Tethys basin opened find are discussed. F westward; the existence of a Carribean corridor linking ABBREVIATIONS the European and Pacific realms now appears well supported CPC: Coleccioń Paleontolo´gica de Coahuila, housed at by comparison of marine reptile assemblages (e.g., Gasparini MUDE: Museo del Desierto, Saltillo, Coahuila, Mexico; SEP: and Fernańdez, 1997, 2005; Gasparini et al., 2000; Fernańdez Secretarıá de Educacioń Pu´blica; SMNK: Staatliches Museum and Iturralde-Vinent, 2000; Gasparini and Iturralde-Vinent, fu¨r Naturkunde, Karlsruhe, Germany; UANL-FCT: Univer- 2001, 2006; Gasparini et al., 2002). Marine transgression in sitad Autońoma de Nuevo Leoń, Facultad de Ciencias de la Mexico began during the Callovian, as evidenced by the Tierra, Linares, Nuevo Leoń, Mexico. evaporites of the Minas Viejas Fm. However, microfossils and invertebrate assemblages indicate that the Mexican Gulf SYSTEMATIC PALEONTOLOGY remained isolated from both the European Archipelago and Order ICHTHYOSAURIA de Blainville, 1835 the Pacific, at least temporarily, until the middle Berriasian; Family OPHTHALMOSAURIDAE Baur, 1887 the Florida uplift and southward movement of Yucatan were Genus OPHTHALMOSAURUS Seeley, 1874 proposed as possibly forming a barrier (Salvador et al., 1993; OPHTHALMOSAURUS CF.O.ICENICUS Seeley, 1874 Adatte et al., 1994, 1996; Goldhammer, 1999; Goldhammer and Johnston, 2001; Gasparini and Iturralde-Vinent, 2006). Referred specimen.—The partial skull and post-cranium is After almost a decade of field work and examination of housed at MUDE under accession number CPC 238 (formerly collections, the Late Jurassic marine reptile assemblage of CEP 1876, Buchy et al., 2006b; Figs 2–4). north-east Mexico confirms the conclusions drawn from Horizon and Locality.—CPC 238 is derived from the Early microfossils and invertebrates. Poorly diagnostic ichthyosaur Tithonian La Caja Formation at the Sierra El Jabalı´, next to remains, with various thalattosuchians, numerically dominate Gomez Farıás, Coahuila (Buchy et al., 2005b, 2006b, fig. 1, the assemblage. Sauropterygians are rare, mainly represented 2006d; Velasco Segura, 2005). Occurrence.—The material was collected during several field by large pliosaurids of unclear affinities, a few vertebrae campaigns between the years 1999 and 2005 by teams from the attributed to elasmosaurids, and a unique cryptoclidid. Turtles SEP and MUDE. Preparation was performed at SMNK by J. are yet to be reported (Frey et al., 2002; Buchy et al., 2003, Lo´pez Espinoza (SEP, MUDE) and the author. 2005b, 2006a–d; Buchy, 2007, 2008a, b; material currently Preservation.—Seen in left lateral view, CPC 238 exhibits under study). skull and mandible in articulation (Fig. 2), while the other side The conglomerates, sandstones and siltstones of the of the specimen comprises both coracoids close to anatomical Kimmeridgian-Berriasian La Casita Formation characterize connection, both scapulae, the left humerus, articulated the Late Jurassic deltaic and inner shelf paleoenviron- cervical centra, including fused atlas and axis, ribs and ments proximal to the Coahuila Peninsula in the area of phalanges (Fig. 3). As is visible in dorsal view (Fig. 4), in Saltillo and Monterrey, whereas the shales, siltstones, and the left and medial portions of the skull, the bones appear phosphorites of the coeval La Caja Formation indicate more mostly undistorted, even if slightly displaced or twisted, while distal outer shelf facies further to the south (Fig. 1; e.g., the right orbital region slid rostrally and the adjacent part of Goldhammer, 1999). the right mandible appears ‘‘folded.’’ The distal part of the Buchy et al. (2006b) indicate that at the Sierra El Jabalı´, rostrum was broken and tilted dorsally but the fragments close to the city of Gomez Farıás, about 70 km south of remained close to their original position. Additional material Saltillo, Coahuila (Buchy et al., 2006d, fig. 1), the Tithonian that cannot be articulated with the main block includes an section of the La Caja Formation includes a 1.5 m thick articular, ribs fragments, phalanges and vertebral centra. coquina layer qualified as a concentration Lagersta¨tte owing As is common for fossils from the locality (Buchy et al., to its richness in marine vertebrates. Following preliminary 2006b, d), the course of most sutures cannot be determined observations, the same authors suggest the coquina deposit due to the poor contrast between matrix and bone. A recent formed in a shallow subtidal mud bottom environment below break with subsequent extensive weathering runs subvertically storm wave base, as a result of reduced rate of sedimentation at the level of the external nares. As noted by McGowan and and condensation, possibly due to transgression. Motani (2003), the overlapping of the cranial bones in The aim of the present paper is to describe a partial ichthyosaurs makes it difficult to extrapolate the external ichthyosaur from the Sierra El Jabalı´, documenting the taxon course of a suture when the surface is damaged. Consequently, Ophthalmosaurus for the first time in Mexico. The specimen the respective participation of the premaxilla, maxilla, nasal, represents the first specimen referred to an ubiquitous taxon of prefrontal and lacrimal to the border of the naris, as well as marine reptiles from the Late Jurassic Mexican Gulf. The the contacts of these bones cannot be determined. 149 150 JOURNAL OF PALEONTOLOGY, V. 84, NO. 1, 2010

element of the palatal area. The lacrimoprefrontal suture commences just dorsal to this element; it undulates rostro- dorsally until it reaches a calcite vein after which it cannot be identified further. The caudal extent of the nasal in the interorbital area, and thus its contact with the frontal, cannot be distinguished. In the preorbital region, an internasal suture is visible; there, the medial surface of both nasals are depressed, the median dorsal surface of the rostrum therefore forming a gutter. The nasal is excluded from the rostrodorsal margin of the orbit by the prefrontal and postfrontal. The suture between the latter two bones appears located around the middle of the dorsal margin of the orbit. The caudal portion of the postfrontal and the other bones forming the temporal bar are damaged and/or obscured by overlying elements (neural arch, ribs). At the caudal-most extremity of the postfrontal, a sinusoidal line delimits the probable facet for the supratemporal. The supratemporal itself cannot be identified. The squamosal is probably visible as a superficial, triangular bone wedged between the caudoventral margin of the postfrontal and the caudodorsal margin of the postorbital. The rest of the suspensorium is obscured by a thick vein of calcite. A bone with a vertical, straight rostral margin and a convex dorsal margin is identified as the quadratojugal. The caudal margin of the quadrate is straight, subvertical. The medial margin of the postorbital was pressed outward by elements of the pectoral girdle; what is exposed in lateral view of the postorbital is mostly its medial postorbital flange. As a result the postorbital is now a boomerang-shaped bone forming the caudal margin of the orbit and the caudal half of its ventral margin, although its original extent and shape cannot be determined. Likely as a geometrical consequence of the shifting of the postorbital, the area of contact with the FIGURE 1—Paleogeographical map of the Late Jurassic (modified from jugal now protrudes into the orbit, with the caudal part of the Smith et al., 1994). Insert: paleobiogeographical map of northeastern Mexico during the Tithonian; Cb: Coahuila block; M: Monterrey; S: jugal dragged by the rotation of the rostral ramus of the Saltillo; scale bar 150 km (redrawn after Goldhammer, 1999). Note the postorbital. Consequently, the surface of the suborbital hypothetical barrier (‘‘Florida-Yucatan ridge,’’ see Gasparini and portion of the jugal is wrinkled like corrugated iron. The Iturralde-Vinent, 2006) formed by the Florida uplift and Yucatan, to caudal extent of the jugal is unclear. As preserved, it wraps the explain the observed endemism of invertebrates and microfossils (see text). caudoventral corner of the postorbital and extends further dorsally toward the quadratojugal. The surangular is exposed laterally further rostral than the Description.—The rostrum being incomplete, no biometric external naris. The angular is well-exposed laterally until at ratio can be determined for the skull (e.g., McGowan and least the level of the nare and appears to extend further Motani, 2003). The orbit is very large, and, taking into rostrally; its rostral-most extent, though, cannot be deter- account distortion the skull underwent, especially shifting of mined due to breakage. The caudal extent of the mandible is the postorbital, was probably subcircular. It accounts for most obscured by the jugal, a phalanx, and matrix. of the postnarial part of the skull in lateral view, the Within the matrix between the premaxilla and the dentary, postorbital region being reduced. The sclerotic ring is almost some teeth are poorly preserved. In the broken area ventral to as large as the orbit itself; its caudoventral portion is displaced the external naris, cross-sections through several teeth are by elements of the ?palate. In some places, the sclerotic plates visible. Further caudally, remnants of at least two upper teeth are covered by a whitish, translucent substance which is absent are laterally exposed over the dentary. These are small, from surrounding bones. conical, about 5 mm in basal diameter, and less than 10 mm in The premaxilla forms most of the rostrum. A 5 mm high height. Enamel is not preserved. groove runs for the entire preserved length of the rostrum at The atlas and axis are fused with the suture still apparent. about mid-height of the premaxilla. The nasal appears The third cervical centrum is not fused to the atlas-axis incomplete rostrally. The maxilla cannot be identified in the complex. The other preserved centra are craniocaudally weathered area ventral to the external naris. compressed (see Bardet et al., 1997; Ferna´ndez, 2007). The lacrimal is subtriangular, forming the rostral margin of The coracoids are preserved close to their original position, the orbit. A subcircular foramen is present in the lacrimal visible in ventral aspect; they bear a single, anterior notch. rostral to the orbit, possibly on the lacrimojugal suture, about Their medial margin is gently rounded. The acetabular margin 30 mm caudal to the external naris. What is here interpreted as is straight, as is seemingly the scapular facet. One scapula is the medial, preorbital portion of the lacrimal was apparently preserved next to the left coracoid and appears slender, but compressed laterally and now occupies the rostral-most few of its original margins are visible. A second scapula is extremity of the orbit. This bone could also represent an probably wedged between the left coracoid and mandible, still BUCHY—OPHTHALMOSAURUS FROM MEXICO 151

FIGURE 2—CPC 238, Ophthalmosaurus cf. icenicus: 1, specimen in left lateral view; 2, interpretative drawing of the specimen. Abbreviations: a: angular; atax: alas/axis; cor: coracoid; cv: cervical vertebra(e); d: dentary; en: external naris; f: frontal; h: humerus; j: jugal; l: lacrimal; ms: mandibular symphysis; mx: maxilla; n: nasal; na: neural arch; o: orbit; par: parietal; paf: parietal foramen; ph: phalanx; pmx: premaxilla; po: postorbital; pof: postfrontal; prf: prefrontal; pt: pterygoid; qj: quadratojugal; r: rib; sa: surangular; sc: scapula; sq: squamosal; st: supratemporal. Scale bar 50 mm. in anatomical contact with the left humerus. The latter is Bajocian of Argentina has a conspicuously small sclerotic ring exposed in dorsal view. It bears three distal facets, the middle compared to the orbital diameter, a feature possibly related to one the largest, the cranial one the smallest and oblique old individual age of the holotype (Fernańdez et al., 2005); its compared to the proximodistal axis. All phalanges present are post-orbital region is more developed than is CPC 238’s rounded. (Fernańdez, 1999, pers. com.). Nannopterygius (Kimmeridgian of England) possesses a humerus reduced in size, with only DISCUSSION two distal facets (McGowan and Motani, 2003). Unodosaurus Comparative Paleontology.—CPC 238 undoubtedly repre- from the Tithonian of Russia possesses large, robust teeth sents Ophthalmosauridae as diagnosed by McGowan and (Efimov, 1999; McGowan and Motani, 2003), although this Motani (2003) in the rostral extent of the angular in lateral genus was considered synonymous with Ophthalmosaurus by view. The orbit is large, the sclerotic ring accounting for most Maisch and Matzke (2000). Of the three distal facets of the of its surface, and the postorbital region is reduced. The humerus of Brachypterygius (Kimmeridgian-Tithonian of humerus has three distal facets. Moreover, what is preserved England and Russia) the middle one (for contact with the of CPC 238 corresponds to descriptions and diagnoses of intermedium) is the shortest; its teeth are robust and its orbit Ophthalmosaurus given by e.g., Seeley (1874), Appleby (1956), relatively small (McGowan and Motani, 2003). The phalanges Kirton (1983), Bardet et al. (1997), Maisch and Matzke (2000), of Aegirosaurus (Tithonian of Germany) are polygonal, and McGowan and Motani (2003). All differences can be the middle facet of its humerus is the smallest of the three attributed to preservation. (Bardet and Fernańdez, 2000). Caypullisaurus, from the CPC 238 clearly differs from other members of Ophthal- Tithonian of Argentina, has polygonal phalanges (Fernańdez, mosauridae in the following features: Mollesaurus from the 1997). Its humerus appears very similar to the humerus of 152 JOURNAL OF PALEONTOLOGY, V. 84, NO. 1, 2010

FIGURE 3—CPC 238, Ophthalmosaurus cf. icenicus: 1, specimen in ‘‘right lateral view’’ (see text); 2, interpretative drawing. Scale 50 mm. Abbreviations as in Figure 2.

Ophthalmosaurus, and McGowan and Motani (2003) suggest icenicus are geographic and stratigraphic.’’ Synonymy of the that the two genera may be synonymous. The phalanges of latter genus with Ophthalmosaurus has long been debated (see Platypterygius (Lower Cretaceous) are rectangular, its teeth McGowan and Motani, 2003) and appears corroborated by are robust, and its orbit is relatively small (McGowan and anatomical data. Specimens referable to Ophthalmosaurus Motani, 2003). were described from Argentina, although the specific attribu- Two species pertaining to Ophthalmosaurus are retained by tion is debatable (Rusconi, 1940; Gasparini, 1988; McGowan McGowan and Motani (2003: 115): the European type species and Motani, 2003; Ferna´ndez, 2007, pers. com.). During the O. icenicus together with O. natans for material from the New Late Jurassic, northeastern Mexico was situated at about the World formerly referred to the genus Baptanodon, noting that same distance from the Western Interior Seaway, the ‘‘[the] only differences reported here between O. natans and O. European Archipelago, and the Neuque´n Basin (e.g., Smith BUCHY—OPHTHALMOSAURUS FROM MEXICO 153

FIGURE 4—CPC 238, Ophthalmosaurus cf. icenicus: 1, specimen in dorsal view; 2, interpretative drawing. Scale 50 mm. Abbreviations as in Figure 2. et al., 1994). Since CPC 238 differs from the type species O. anatomy demonstrates its being close to Brachypterygius icenicus in no perceived significant feature, the most conser- (Buchy and Lo´pez Oliva, in press). vative option is to refer it to this species. Paleobiogeography.—CPC 238 represents the first, diagnos- Late Jurassic Mexican ichthyosaurs.—Ichthyosaur remains tic occurrence in the Late Jurassic Mexican Gulf of an from the Late Jurassic northern Mexican Gulf consist mostly ubiquitous taxon of marine reptile, whereas recent work had of isolated vertebral centra or clusters of centra (Buchy et al., indicated both endemism and a peculiar composition of the 2006c; material in collection at MUDE and UANL-FCT and assemblage (Frey et al., 2002; Buchy et al., 2003, 2005b, in private hands). Among those, two specimens from the La 2006a–d). This endemism appears coincident with local Caja/La Casita Formations were mentioned and attributed to paleogeography. Block tectonics and sea level fluctuations the genus Ichthyosaurus (Aguirre Garza et al., 2000; Velasco- during the Late Jurassic in northeastern Mexico caused an Segura, 2005), which is stratigraphically unlikely and anatom- irregular sea floor topography and variable restrictions or ically impossible to substantiate. subdivision of basins, coherent with observed endemism of It has been suggested that all ichthyosaur remains from the marine invertebrates until the middle Berriasian (e.g., Salva- area likely belong to Ophthalmosauridae (Buchy et al., 2006c). dor et al., 1993; Adatte et al., 1994, 1996; Goldhammer, 1999; In view of unpublished MUDE material recently prepared, Goldhammer and Johnston, 2001; Gasparini and Iturralde- consisting of vertebrae, girdle elements and rounded phalan- Vinent, 2006). Hence, CPC 238 possibly invalidates the ges, it is here considered likely that most do belong to the apparent endemism of Mexican marine reptiles in general. genus Ophthalmosaurus. An exception is a large partial This hypothesis would imply that thalattosuchian taxa skeleton (Buchy and Lo´pez Oliva, 2009) from a new Late described from the Mexican Gulf were not yet discovered Tithonian locality close to the type locality of the La Casita elsewhere, and that pending preparation and description of the Formation, whose mid-dorsal vertebrae, with a diameter of sauropterygians will indeed reveal their belonging to hitherto about 120 mm, exceed in size what is known for Ophthalmo- known taxa. However, it does not explain the absence of saurus (e.g., Kirton, 1983; Motani, 2005), and whose humeral turtles after decades of field collection in the various facies 154 JOURNAL OF PALEONTOLOGY, V. 84, NO. 1, 2010 known of the La Caja/La Casita/La Pimienta Formations Consejo Estatal de Ciencia y Tecnologıá Coahuila (grant spanning from Kimmeridgian to Berriasian (Buchy, 2007). COAH-2008-C04-37) and MUDE. Buchy et al. (2006d) suggested that turtles had not made it from Europe to the Mexican Gulf by the Kimmeridgian. REFERENCES Marine turtles are present in the Tithonian Neuqueń Basin of ADATTE, T., W. STINNESBECK, AND J. REMANE. 1994. The Jurassic- Cretaceous boundary in Northeastern Mexico. Confrontation and Argentina (e.g., Fernańdez and Fuente, 1993; Gasparini and correlations by microfacies, clay mineral mineralogy, calpionellids and Fernańdez, 1997, 2005), so their absence from the Tithonian ammonites. Geobios, 17:37–56. Gomez Farıás site is quite peculiar (Buchy, 2007). 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VIGNAUD,E.FREY,W.STINNESBECK, AND A. H. (UANL-FCT) and A. H. Gonza´lez Gonza´lez (MUDE). Part GONZA´ LEZ GONZA´ LEZ. 2005b. New occurrence of the genus Geosaurus of the present work was included in a Ph.D. dissertation (Thalattosuchia, Crocodyliformes) in the Tithonian (Upper Jurassic) of Mexico. Kaupia, 14:72. supervised by E. Frey (SMNK) and W. Stinnesbeck (Univer- BUCHY, M.-C., E. FREY,S.W.SALISBURY,W.STINNESBECK,J.G.LO´ PEZ sity of Heidelberg). The constant presence in the field of Jose´ OLIVA, AND M. GO¨ TTE. 2006a. An unusual pliosaur (Reptilia, ‘‘Pato’’ Lo´pez Espinoza (MUDE) was essential, and his Sauropterygia) from the Kimmeridgian (Upper Jurassic) of northeast- preparation skill is praised. Thanks are due to the SEP/ ern Mexico. Neues Jahrbuch fu¨r Geologie und Palaöntologie, 240:241– MUDE team, past and present. For access to literature and 270. BUCHY, M.-C., E. FREY,W.STINNESBECK, AND A. H. GONZA´ LEZ comments, thanks are due to M. 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