A Eutherian Mammal in the Latest Cretaceous of Vitrolles, Southern France
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A eutherian mammal in the latest Cretaceous of Vitrolles, southern France RODOLPHE TABUCE, MONIQUE VIANEY−LIAUD, and GÉRALDINE GARCIA Tabuce, R., Vianey−Liaud, M., and Garcia, G. 2004. A eutherian mammal in the latest Cretaceous of Vitrolles, southern France. Acta Palaeontologica Polonica 49 (3): 347–356. In Europe, the fossil record of the eutherian mammals is very scanty for the Late Cretaceous, as only two genera, docu− mented by isolated teeth, are presently recorded in France and in Spain. Both genera, Labes and Lainodon, are considered to be representatives of the “zhelestids”, a paraphyletic unit regarded as being at the origin of Cenozoic ungulates within the Ungulatomorpha clade. We here describe Valentinella vitrollense gen. et sp. nov. from Vitrolles la Plaine (Maas− trichtian, southern France). This species, represented by fragmentary remains of lower and upper dentitions, is tentatively assigned to the “zhelestids” according to the hypoconulid−entoconid twinning and the antero−posteriorly short trigonid on m1–3. The occlusal surfaces are obliterated by dental attrition, but Valentinella could be an evolved “zhelestid”, more de− rived than Labes and Lainodon by its fully compressed trigonid. Valentinella is similar to Gallolestes by other derived characters such as a crushing specialization of the teeth, associated with a probably molariform p4 (or dp4) and slightly re− duced m3. The enamel microstructure, showing a radial prismatic pattern combined with a reduced interprismatic matrix, in which cristallites are oriented at about 45° to the prisms axes, appears compatible with the ancestral morphotype for all ungulates; although no synapomorphy can be proposed for the ungulatomorphs. Key words: Mammalia, Eutheria, “Zhelestidae”, enamel microstructure, Cretaceous, France. Rodolphe Tabuce [[email protected]−montp2.fr], Monique Vianey−Liaud [[email protected]−montp2.fr], Labora− toire de Paléontologie, Institut des Sciences de l’Evolution, cc064, Université Montpellier II, place Eugène Bataillon, 34095 Montpellier cedex 05, France; Géraldine Garcia [geraldine.garcia@univ−poitiers.fr], Laboratoire de Géobiologie, Biochronologie et Paléontologie Humaine, Faculté des Sciences Fondamentales et Appliquées, Université de Poitiers, 40 avenue du Recteur Pineau, 86022 Poitiers cedex, France. Introduction The eutherian mammals recovered from these sites are still poorly documented, they are only known by fragmentary or According to present paleontological knowledge, the Late isolated complete teeth. Eutherians are undoubtedly docu− Cretaceous is a key period for the evolution of placental mented only in Champ−Garimond, Taveiro, Laño, and Quin− mammals, as the first representatives of the extant supra− tanilla del Coco. ordinal groups originated at that time (McKenna and Bell During excavations of dinosaur bones and eggshells in an 1997; Archibald et al. 2001). In Europe, Late Cretaceous Upper Cretaceous locality near Vitrolles (southern France) mammalian history is poorly understood because of the pau− (Garcia 1998; Garcia and Vianey−Liaud 2001), three worn city of continental deposits yielding mammals. Thus, the fos− and crushed mammalian specimens have been recovered. sil record in this area is very scarce in contrast to other The mammalian specimens are in association with other ver− holartic provinces (e.g., western North America, Gobi tebrate remains, some ratite eggshells, and typical Rognacian Desert, and Uzbekistan in Asia). Only a few localities are molluscs (Lychnus matheroni) (Garcia et al. in preparation). known, all of them located in the Iberian−Armorican and The Rognacian is a regional stratigraphic unit correlated with Transylvanian islands that made up part of the European the upper Campanian plus Maastrichtian by the magneto− Archipelago (Rage 2002): Champ−Garimond (Campanian, stratigraphy and the dinosaur eggshells distribution (Garcia France) (e.g., Sigé et al. 1997), La Neuve (Campanian, Fran− and Vianey−Liaud 2001). The occurrence of dinosaur egg ce) (Garcia et al. 2000), Laño (?Campanian, Spain) (e.g., clutches, belonging to the Megaloolithus mamillare oospecies, Gheerbrant and Astibia 1999), Taveiro (Late Campanian or restricts the age of the site to the Maastrichtian. Besides, the Maastrichtian, Portugal) (Antunes et al. 1986), Peyrecave position of the locality, below the Rognac Limestone, sug− (Maastrichtian, France) (Gheerbrant et al. 1997), Quintanilla gests an early Maastrichtian age for Vitrolles la Plaine (Table del Coco (Maastrichtian, Spain) (Pol et al. 1992), Pui (Maas− 1, Fig. 1). trichtian, Romania) (e.g., Rădulescu and Samson 1997), The mammalian specimens belong to a new eutherian ge− Toteşti−Baraj (Maastrichtian, Romania) (Codrea et al. 2002), nus that seems to be related to the “zhelestids”. This para− and Nălaţ−Vad (Maastrichtian, Romania) (Smith et al. 2002). phyletic family is considered as closely related to Cenozoic Acta Palaeontol. Pol. 49 (3): 347–356, 2004 http://app.pan.pl/acta49/app49−347.pdf 348 ACTA PALAEONTOLOGICA POLONICA 49 (3), 2004 limestone marl sandstone Vitrolles limestome (early Paleocene) Rognac Rognac limestome (Maastrichtian) Vitrolles Fossilerous level Mediterranean sea MARSEILLE 10 m Fig. 1. A. Location of the Vitrolles la Plaine site (Arc Basin, Bouches−du−Rhône, southern France). B. Stratigraphic section through the fossiliferous layers. archaic and modern ungulates within the Ungulatomorpha clade (Archibald 1996; Nessov et al. 1998; Archibald et al. Systematics 2001). The new genus here described is therefore a putative critical representative of one of the very few superordinal Class Mammalia Linnaeus, 1758 groupings of modern mammals known before the Creta− Infraclass Eutheria Gill, 1872 ceous−Paleogene boundary. Supergrandorder cf. Ungulatomorpha Archibald, Storage.—The specimens described in this paper are housed 1996 in the collections of the Institut des Sciences de l’Evolution, Family cf. “Zhelestidae” Nessov, 1985 Université Montpellier II (France) with the abbreviation Genus Valentinella nov. ISEM/VLP; the second part of the abbreviation refers to Type species: Valentinella vitrollense sp. nov. Vitrolles la Plaine. Known range: Maastrichtian, Arc Basin, southern France. Table 1. Faunal list from the Vitrolles la Plaine site (Maastrichtian, Diagnosis.—That of the type species. southern France). Etymology.—Dedicated to Xavier Valentin (University of Bony fish Poitiers, France), who discovered the specimens. Ginglymodi Lepisosteidae indet. Valentinella vitrollense sp. nov. Teleostei Figs. 2, 3. Sparidae indet. Holotype: ISEM/VLP−2, damaged right dentary with p3–m3. Squamates Squamata indet. Referred material.—ISEM/VLP−4: fissured fragment of a Turtles right dentary with a damaged ?p4, associated with two very Pleurodira fragmentary adjacent teeth. ISEM/VLP−3: a fractured maxil− Bothremydidae lary fragment with roots of right and left canines and a right cf. Polysternon ?P2. Crocodilians Diagnosis.—Valentinella presents the classic “zhelestid” fea− Eusuchia tures: the teeth are characterized by developed crushing func− Alligatoroidea cf. Musturzabalsuchus tion, the paraconid is lingually or sublingually positioned with a clear appression to the metaconid, the entoconid and the Dinosaurs Saurischia hypoconulid are twinned, and the talonid is expanded labio− Theropoda indet. lingually. Valentinella differs from the Asian “zhelestids” by Sauropoda the molarization of the ultimate premolar, and is distinct from Titanosauridae indet. Avitotherium and Gallolestes by its simple, bulbous p3 and by Mammals its larger metaconid and entoconid on lower molars, respec− Eutheria tively. Valentinella differs from Labes and Lainodon in hav− Valentinella vitrollense gen. et sp. nov. ing a mesio−distally compressed trigonid. TABUCE ET AL.—CRETACEOUS EUTHERIAN MAMMAL FROM FRANCE 349 Etymology.—From Vitrolles, name of the town near the crushed. The tooth is strongly worn, suggesting, like the fossiliferous site. occlusal surfaces observed in m1–2, a developed crushing Measurements.—(in mm): ISEM/VLP−2: p3 (L = 2.3; W = function of the teeth during dental attrition. On the talonid, 1.5); p4 (or dp4) (L = 2.3; W = 2); m1 (L = 3.4; W = 2.35; Ltri which is shorter than the trigonid, the remaining area of the = 2; Ltal = 1.4; Wtri = 2; Wtal = 2.35); m2 (L >3.5; W = 2.9); postcristid does not suggest a strong distal development of a m3 (L = 3.1; W >2). ISEM/VLP−4: ?p4 (Ltri = 1.5; Ltal = 1.2; hypoconulid. Wtri = 1.35; Wtal = 1.9). ISEM/VLP−3: right canine (H >2.5; ISEM/VLP−4 is a very damaged specimen which dis− diameter of the root = 1.7); left canine (H >2.3; diameter of plays fragments of both ?p3 and ?m1, plus a better pre− the root = 1.6); ?P2 (L = 2.5; W = 2.0) served ?p4. Only the lingual wall of the ?p3 is preserved, the length of this tooth being superior to 1.5 mm, as roughly Description.—During diagenesis and compaction of the sed− estimated from the two roots. The mesio−distal axis of the iment, the teeth were distorted along parallel cracks caused ?p4 shows a 90° labial rotation relative to the ?p3. The tooth by plant roots, and/or between talonids and trigonids, and be− is probably two−rooted, although the roots are not well−pre− tween teeth. These movements have produced oblique defor− served on the specimen. The crown appears to be high mations. (about 2 mm, on the disto−lingual side), but because of poor On ISEM/VLP−2 (Fig. 2), the dentary is almost an outline enamel preservation, it is difficult to identify the position of of a jaw fissured in the marl,