Carnivora from the early Oligocene of the ' Phosphorites du Quercy' in southwestern France

2019 ● 41 ● 15 geodiversitas M e m o r i a l S t é p h a n e P e i g n é – C a r n i v o r e s o f t h e C e n o z o i c – M e m o e i g g r n i v n o z o i c g g Geodiversitas est une revue en flux continu publiée par les Publications scientifiques du Muséum, Paris Geodiversitas is a fast track journal published by the Museum Science Press, Paris Les Publications scientifiques du Muséum publient aussi / The Museum Science Press also publish: Adansonia, Zoosystema, Anthropozoologica, European Journal of Taxonomy, Naturae, Cryptogamie sous-sections Algologie, Bryologie, Mycologie. Diffusion – Publications scientifiques Muséum national d’Histoire naturelle CP 41 – 57 rue Cuvier F-75231 Paris cedex 05 (France) Tél. : 33 (0)1 40 79 48 05 / Fax : 33 (0)1 40 79 38 40 [email protected] / http://sciencepress.mnhn.fr © Publications scientifiques du Muséum national d’Histoire naturelle, Paris, 2019 ISSN (imprimé / print) : 1280-9659/ ISSN (électronique / electronic) : 1638-9395 Directeur De la publication : Bruno David, Président du Muséum national d’Histoire naturelle réDacteur en chef / Editor-in-chiEf : Didier Merle assistants De réDaction / AssistAnt Editors : Emmanuel Côtez ([email protected]) ; Anne Mabille Mise en page / PAgE lAyout : Emmanuel Côtez coMité scientifique / sciEntific boArd : Christine Argot (MNHN, Paris) Beatrix Azanza (Museo Nacional de Ciencias Naturales, Madrid) Raymond L. Bernor (Howard University, Washington DC) Alain Blieck (chercheur CNRS retraité, Haubourdin) Henning Blom (Uppsala University) Jean Broutin (UPMC, Paris) Gaël Clément (MNHN, Paris) Ted Daeschler (Academy of Natural Sciences, Philadelphie) Bruno David (MNHN, Paris) Gregory D. Edgecombe (The Natural History Museum, Londres) Ursula Göhlich (Natural History Museum Vienna) Jin Meng (American Museum of Natural History, New York) Brigitte Meyer-Berthaud (CIRAD, Montpellier) Zhu Min (Chinese Academy of Sciences, Pékin) Isabelle Rouget (UPMC, Paris) Sevket Sen (MNHN, Paris) Stanislav Štamberg (Museum of Eastern Bohemia, Hradec Králové) Paul Taylor (The Natural History Museum, Londres) couverture / covEr : Made from the figures of the article. Geodiversitas est indexé dans / Geodiversitas is indexed in: – Science Citation Index Expanded (SciSearch®) – ISI Alerting Services® – Current Contents® / Physical, Chemical, and Earth Sciences® – Scopus® Geodiversitas est distribué en version électronique par / Geodiversitas is distributed electronically by: – BioOne® (http://www.bioone.org) Les articles ainsi que les nouveautés nomenclaturales publiés dans Geodiversitas sont référencés par / Articles and nomenclatural novelties published in Geodiversitas are referenced by: – ZooBank® (http://zoobank.org) 601 GEODIVERSITAS • 2019 • 41 (15) © Publications scientifiques du Muséum national d’Histoire naturelle, Paris. www.geodiversitas.com urn:lsid:zoobank.org:pub:9DD3CC29-3AEA-44B8-8E8F-6AD882DF5B1C Bonis L. de, Gardin A. & Blondel C. 2019. — Carnivora from the early Oligocene of the ‘Phosphorites du Quercy’ in southwestern France, in Bonis L. de & Werdelin L. (eds), Memorial to Stéphane Peigné: Carnivores (Hyaenodonta and Carnivora) of the Cenozoic . Geodiversitas 41 (15): 601-621. https://doi.org/10.5252/geodiversitas2019v41a15. http://geodiversitas.com/41/15. ABSTRACT The early Oligocene carnivorans from the Quercy phosphorites (southwestern France) studied herein come from three localities: Mas de Got, La Plante2 and Valbro, all in MP 22 of the European Paleogene Mammal stratigraphy scheme. These localities contain a rich carnivoran fauna with weaselsized to tiger-sized species in diverse families, especially Amphicynodontidae (with four species of Pachycynodon and two of Amphicynodon) and Nimravidae. New taxa are described (Wangictis n. gen., Peignictis pseudamphictis n. gen., n. sp.). We note the absence of taxa often present in the sites of the Quercy phosphorites, such as the family Amphicyonidae and, in the Feloidea, the genera Stenoplesictis Filhol, 1880 and Palaeoprionodon Filhol, 1880. The fauna of the three localities is typical of the early Oligocene prior to a faunal turnover in the middle Oligocene. Future research in the Quercy will give more precision to the timing and mode of this faunal event. RÉSUMÉ Carnivores de l’Oligocène inférieur des ‘Phosphorites du Quercy’, sud-ouest de la France. Les carnivores de l’Oligocène inférieur des phosphorites du Quercy (sud ouest de la France) examinés dans cette étude proviennent de trois localités Mas de Got, La Plante2 et Valbro, toutes dans l’unité MP 22 de l’échelle mammalienne du Paléogène. Elles contiennent une riche faune de carnivores, dont les tailles s’étagent depuis celle d’une belette jusqu’à celle d’un tigre, à travers diverses familles, en particulier celles des Amphicynodontidae (avec quatre espèces de Pachycynodon et deux d’Amphicynodon) et des Nimravidae. De nouveaux taxons sont décrits (Wangictis n. gen., Peignictis pseudamphictis n. gen., n. sp.). Nous notons également l’absence de certaines formes très souvent présentes dans les sites fossilifères des phosphorites du Quercy, comme la famille des Amphicyonidae et, parmi les Feloidea, les genres Stenoplesictis Filhol, 1880 et Palaeoprionodon Filhol, 1880. La faune de ces trois localités est typique de l’Oligocène inférieur avant un changement qui se produira vers le milieu de l’Oligocène. Les futures recherches dans le Quercy devront préciser la date et les modalités de cet évènement. Louis de BONIS Axelle GARDIN Cécile BLONDEL Palevoprim: laboratoire de Paléontologie, Évolution, Paléoécosystèmes, Paléoprimatologie, Bâtiment B35 TSA51106, 6 rue Michel Brunet F-86073 Poitiers cedex 9 (France) [email protected] [email protected] [email protected] Submitted on 10 February 2019 | accepted on 16 April 2019 | published on 10 September 2019 Carnivora from the early Oligocene of the ‘Phosphorites du Quercy’ in southwestern France KEY WORDS Caniformia, Feliformia, Amphicynodontidae, Ursidae, Mustelidae, Nimravidae, sabre-toothed carnivorans, Paleogene, MP 22, lectotypification, new combination, new genera, new species. MOTS CLÉS Caniformia, Feliformia, Amphicynodontidae, Ursidae, Mustelidae, Nimravidae, carnivores à canines-ensabre, Paléogene, MP 22, lectotypification, combinaison nouvelle, genres nouveaux, espèce nouvelle. 602 GEODIVERSITAS • 2019 • 41 (15) Bonis L. de et al. INTRODUCTION The Quercy area is situated in southwestern France, near the city of Cahors. A large part of this area is covered by Jurassic limestone in which erosion hollowed fissures, caves and galleries to form a large karstic system. Through time, there was a succession of erosion and infilling by sediments. The sediments contain phosphate deposits and, during the last third of the 19 th century, they were worked to extract the ‘phosphorite’, which was used as a fertilizer for cultivation and farming. In several localities the sediments were rich in diverse categories of plant, invertebrate and vertebrate fos - sils. In particular, countless thousands of vertebrate remains, comprising skulls, teeth and bones, were unearthed by the workers from many quarries lying across an area of hundreds of km2. These fossils were sold or given to different people, amateurs and professionals, so that the fossils are now scattered in collections across the whole world. The geological age of the specimens extends from the early Eocene (Astruc et al. 2000) to the early Miocene (Sigé et al. 1991). Nevertheless, most of the fossils were deposited from the late Eocene to the late Oligocene but are today all mixed together. Therefore, the numerous paleontological studies of the old collections during the 19th could not work out problems of evolution and concluded that the fossils were mixed at every locality (see Bonis 2011 for more complete references). The same has been the case concerning the Carnivora in more recent publications (Teilhard 1915; Piveteau 1931, 1942, 1962; Bonis 1966, 1971, 1981; Ginsburg 1966, 1979; Lange 1969, 1970; Wolsan & Lange-Badré 1996; Peigné & Bonis 1999; Peigné 2000, 2001, 2003), in which new genera and species were described but did not clear up the different lineages. At that time, many people thank thought all the fossil bearing fissures were emptied. Another period began during the 1930th when new researches (Gèze 1938a, b) demonstrated that two Quercy localities had retained their sediment infills, including fossil fauna. Each of these fossil faunas was homogeneous, without any mixing. From the middle of the 1960s, a team of researchers from some French universities (Montpellier UM, Paris 6 [UPMC], Lyon 1 [Claude Bernard], Poitiers and the Muséum national d’Histoire naturelle, Paris) decided to carry out paleontological field campaigns in the Quercy area. Over the years, the results were published, bearing out the homogeneity of the sites (Vianey-Liaud & Legendre 1986) and describing hundreds of species from several orders of mammals. Some of these Quercy localities played a significant role in the creation of the European mammal stratigraphic scheme for the Paleogene numbered from MP 1 to MP 30, with the type levels MP 17b (Perrière), MP 19 (Escamps), MP 23 (Itardies), MP 25 (Garouillas), MP 26 (Mas de Pauffié) and MP 28 (Pech du Fraysse) all coming from the Quercy area (see Biochrom’97 1997). Carnivores (Hyaenodonta and Carnivora) are present in most of the sites and have been the subject of several papers. In the present article we describe early Oligocene Carnivora specimens belonging to different taxa coming from localities of MP 22: La Plante2, Mas de Got, and Valbro. MATERIAL AND METHODS The studied material comes from several field campaigns in the Quercy. The fossils were obtained both by excavating and by washing-screening as well. Measurements were made by digimatic callipers to the nearest 1/100 mm and presented to the nearest 1/10. The new material was compared to the old collections of Quercy, containing most of the type specimens, as well as to fossils from other localities. The photographs were taken with a camera Nikon Coolpix 750 and processed in Adobe Photoshop. AbbreviAtions D deciduous upper molar; d deciduous lower molar; aac anterior (mesial) accessory cusp; aacd anterior (mesial) accessory cuspid; pac posterior (distal) accessory cusp; pacd posterior (distal) accessory cuspid; L length; w width; w1 mesial width; w2 distal width; tr trigonid; tl talonid; mts metastyle; MP Mammal Paleogene reference levels. Institutions MNHN.F Muséum national d’Histoire naturelle, Paris, collections de Paléontologie; NHMB Naturhistoriches Museum Basel; UM Université de Montpellier; UP Université de Poitiers. MP 22 Localities VD, VBO, VBOA Valbro; MGA, MGB Mas de Got; LPL La Plante2. Teeth We use capitals for the upper teeth and lower case for lower teeth. SYSTEMATICS Order CARNIVORA Bowdich, 1821 Suborder CANIFORMIA Kretzoi, 1943 Parvorder URSIDA Tedford, 1976 Family AmphicynodontidAe Simpson, 1945 Genus Amphicynodon Filhol, 1881 type species. — Cynodon velaunus Aymard, 1846 (by original designation). diAgnosis. — Amphicynodontidae with a relatively low skull with a faint sagittal crest slightly more developed distally; elongate muzzle; ossified bullae; shallow suprameatal fossa; alisphenoid canal present. Dental formula 3/3 I, 1/1 C, 4/4 P, 2-3/3 M. Simple premolars P1-P3 and p1-p3; short P4 with robust protocone and short metastyle; triangular M1 with tiny parastyle without para- 603 Oligocene carnivorans from Quercy GEODIVERSITAS • 2019 • 41 (15) conule but with metaconule; M2 more or less reduced; p4 with pacd or not; grouped and not very tall trigonid of m1, horizontal wear of the cuspids, metaconid as high as the paraconid; not very elongate m2 with paraconid present or not, metaconid as high or higher than the protoconid; small rounded m3. The genus is present in the old Quercy collections represented by several species but also in the new early Oligocene localities (Table 1). remArk Aymard (1846) created the species C. velaunus for a fossil from the early Oligocene locality Ronzon (France) that he described a few years later (Aymard 1850). He added another species, C. palustris Aymard, 1853 for a hemi-mandible slightly larger than that of C. velaunus. Later (1881) Filhol, considering that C. palustris was very different from C. velaunus, created the new genus Amphicynodon for it. Finally, the name Cynodon being pre-occupied by a fish, Trouessart (1904) proposed the genus name Aymardia. However, following the rules of nomenclature, Amphicynodon Filhol, 1881 has priority, although Teilhard (1915), without taking into account either Filhol or Trouessart, continued using Cynodon. Later, several species of Amphicynodon were described, principally from the old collections of the Quercy, thus lacking biostratigraphic support. We shall here focus on the specimens from MP 22. Amphicynodon typicus (Schlosser, 1888) (Fig. 1B) type specimen. — Neotype by designation (Cirot & Bonis [1992: 121]): left hemi-mandible with p4-m2 UP PC46 from Pech Crabit, MP 23. new mAteriAl. — Left-hemi-mandible UP VAL2; right hemimandible with m2 UM VD13. description The left hemi-mandible UP VAL2 (Fig. 1B) preserves the tooth series p2-m3. The corpus is elongate with a convex lower border. The ascending ramus and the distal part of the mandible are broken off, leaving only the rostral part of the masseteric fossa. The teeth are unworn; the single-rooted p1 is missing. The p2 is asymmetric, the distal portion being very elongate, p3, separated by a short diastema, is larger and less asymmetric. The p4, the least asymmetric premolar, is tall, and pointed with a pacd and the base of the crown slightly upturned mesially and distally. A faint cingulid underlines the base of the buccal side of m1; the protoconid, higher than the p4, is slightly bucco-lingually compressed; the paraconid, set slightly obliquely, extends the shearing blade of the trigonid; the conical, reduced metaconid is not visible in buccal view; a relatively short talonid displays a large and compressed hypoconid and a very low entoconid, in contact at their bases, creating a V-shaped talonid basin although most of the surface is occupied by the sloping lingual face of the hypoconid. This structure is seen in A. typicus. The protoconid and metaconid of m2, the former being slightly the larger, enclose a well-developed mesial fovea. The distal fovea, enclosed by the hypocristid and entocristid, is more elongate. The buccal portion of m3 is partially broken off, the tooth seems to have had a circular flat basin enclosed by a swollen cristid. Two main characters present in UP VAL2 and A. typicus are the pointed p4 and the trenchant talonid of m1. They seem sufficient to allow us to place the specimen in this species. The hemi-mandible UM VD13 is shallow and includes fragments of the roots of p3 and p4, alveoli of m1, m3 and an unworn m2. Its outline tapers distally. The preprotocristid of a well-developed protoconid runs to a tiny mesio-lingual bulge corresponding to the paraconid, which is separated by a notch from the metaconid. The latter, as high as the protoconid, is followed directly by the postmetacristid to join a globular hypoconid, enclosing a small but deep talonid basin. The trigonid occupies 2/3 of the crown. Amphicynodon sp. 1 (Fig. 1A) description Less complete than the former, another left hemi-mandible (UP VAL1) retains only p3-p4, m2 and alveoli of p1-2, m1, m3 (Fig. 1A). It is slightly larger than UP VAL2. The dentition is unworn, p1 and m3 are single rooted, and p3 is slender and asymmetric. The p4 is less asymmetric and includes a pacd, the crown terminating distally in a small upturn. Based on the alveoli the length of m1 is estimated to 8 mm. The structure of m2 is unusual with a large mesial fovea limited mesially by a cristid and, distally by the cristids of protoconid and metaconid, two pyramidal cuspids of similar size, with convex buccal and lingual and lingual faces respectively. There is a very long basin bordered by the hypocristid and entocristid, terminating in the hypoconid and entoconid respectively, followed by cristids distally closing the basin. An m2 that long is unusual in Amphicynodon. Without taking into account forms with an especially short m2 (A. velaunus and A teilhardi), the index (length m2/ length m1) × 100 is 75 (the length of m1 measured from the alveoli). In the sample from the locality Itardies (MP 23), the same index is 44-62, mean 52. It is 54 for the type specimen of A. crassirostris (Filhol, 1876), 56 for A. typicus (Schlosser, 1888), 48 for A. rossignoli (Filhol, 1882) but 75 for A. gracilis (Filhol, 1874). Thus, the latter has the same proportions as UP VAL1 and we may place this specimen near A. gracilis. Nevertheless, there are some differences insofar as A. gracilis is smaller, there is no pacd in p4 and all the premolars have a marked cingulid. Thus, this identification is inappropriate and UP VAL1 remains Amphicynodon sp. Amphicynodon sp. 2 (Fig. 1D) description A right M1 (UM VBO492) from Valbro (Peigné et al. 2014: fig. 22b) has an overall pattern indicating Amphicynodon 604 GEODIVERSITAS • 2019 • 41 (15) Bonis L. de et al. morphology. Thus UM VBO492, shares with the M1 of A. leptorhynchus (Filhol, 1876) found in the locality Itardies (Cirot & Bonis 1992), a similar aspect of the trigon, the presence of a metaconule and a large lingual cingulum (Fig. 1D). On the other hand, it is narrower distally than mesially, indicating a reduction of the distal part of the jaw. This character is present in A. velaunus and A. teilhardi, but the outline of the tooth is different, with a greater length of the buccal border in those two species. We do not know if in our material there is a lower dentition corresponding to UM VBO492 and thus we cannot assign a species name to this M1, although Amphicynodon typicus could be a possibility. Amphicynodon sp. 3 (Fig. 1C) new mAteriAl. — Right M1 UM VD49. description The right M1 UM VD49 (Fig. 1C) has a symmetrical outline like that of A. leptorhynchus (Cirot & Bonis 1992). Paracone and metacone are nearly identical in size and are on more or less a same longitudinal line. A thick cingulum runs along the buccal side without any style. The protocone is situated slightly lingually relative to the middle of the crown. The postprotocone and postmetaconule cristae fade out in contact with the distal face of the metacone. There is a huge lingual cingulum forming a true cusp. There are small wear facets on buccal cusps and large wear facets on metaconule, protocone and lingual cingulum. The morphology of this M1 differs from that of the left M1 UM VBO1212 (Peigné et al. 2014) which is more symmetrical, with both preprotocone and postprotocone cristae. But neither tooth can be firmly associated to a mandible of any named species. Genus Pachycynodon Schlosser, 1888 type species. — Pachycynodon crassirostris Schlosser, 1888 by original designation. description Schlosser created the genus Pachycynodon for a P4 and M1 from the Quercy phosphorites (Schlosser 1988: pl. IX, fig. 2, 7, 9 and pl. IX, fig. 4 respectively) without any biostratigraphic context. He thought that these specimens were conspecific with Cynodictis crassirostris Filhol, 1876 (1876: figs 67, 69 and 72) but, in fact the Filhol’s specimen, a mandible with c, p3, m1-m2, belongs to the genus Amphicynodon. Thus, the name crassirostris being conserved, the type species of the genus is Pachycynodon crassirostris. Later, several species were added to the genus and to the Amphicynodontidae, forming a group with robust dentition (more robust than that of Amphicynodon), mandible often deep, p4 as high or higher than the protoconid of m1, massive cuspids of the trigonid of m1, postmetaconid cristid sloping very obliquely toward the preentocristid, paraconid often present in m2. The robust P4 has a short metastyle and large plateau-like protocone. There is a trend in M1 toward an almost quadratic outline, without paraconule and disto-lingual metaconule and with a large cingulum around surrounding the protocone. The M2 is more or less reduced. The muzzle is elongated. Pachycynodon crassirostris Schlosser, 1888 type specimen. — Lectotype, right P4 in Schlosser 1888: 29, pl. IX, fig. 2 and 9 “Phosphorites du Quercy”, BSP 1879XV40, Museum of Munich, by subsequent designation of Peigné et al. 2014. description The type specimen of the species Pachycynodon crassirostris Schlosser, 1888, an isolated P4 comes from an unknown locality in the Quercy phosphorites, without any biostratigraphic indicators. This tooth is not frequent in the collections and, except for P. boriei, never associated with a mandible. The P4 is not a good diagnostic element for a species insofar as all type specimens of other species are mandibles and lower dentitions. Schlosser considered that the type could correspond to a mandible named Cynodictis crassirostris Filhol, 1882 but we don’t know the criteria he used (see above). Later (1899) he figured other specimens from Quercy, a mandible and an m1 (Schlosser 1899: pl. VIII, figs 1, 8), which seemed to him match the upper tooth considered as the type specimen. Teilhard figured a Quercy mandible from the Museum of Montauban (Teilhard 1915: pl. IV, fig. 11) as P. crassirostris. The latter was synonymised by Cirot (1992) because of mandibular morphology with another Quercy species, Cynodictis dubius Filhol, 1882 (1882: pl. VIII, figs 11-13) of which type specimen, a mandible, is housed in the MNHN Paris (no. MNHN.F.QU3231). However an examination of this specimen (by LdeB) shows that it belongs in the genus Pachycynodon but differs from the Montauban mandible by its smaller size, relatively lower cuspids of m1 trigonid, relatively longer m2 table 1. — Measurements (in mm) of teeth of Amphicynodon Filhol, 1881, from the alveoli. Amphicynodon p2L p2w p3L p3w p4L p4w m1L m1Ltr m1wtr m1wtl m2L m2wtr m2wtl m3L m3w UP VAL1 – – 6.68 3.33 6.98 3.33 (8.1) – – – 6.23 6.23 4.1 3.59 – UP VAL2 4.86 2.3 4.61 2.33 7.09 2.82 7.99 5.56 4.17 3.83 4.27 3.47 2.89 2.43 2.01 M1L M1w1 M1w2 UM VBO492 6.5 9.2 8.8 UM VD49 6.9 10.3 8.3 605 Oligocene carnivorans from Quercy GEODIVERSITAS • 2019 • 41 (15) A1 A3 B3 A2 B1 B2 C, D C D fig. 1. — A, Amphicynodon sp. 1: left hemi-mandible UP VAL2, in buccal (A1), lingual (A2), and occlusal (A3) views; B, Amphicynodon typicus: left mandible UP VAL1, in buccal (B1), lingual (B2), and occlusal (B3) views; C, Amphicynodon sp. 3: right M1, UM VD49, in occlusal view; D, Amphicynodon sp. 2: right M1, UM VBO492, in occlusal view. Scale bars: A, B, 10 mm; C, D, 5 mm. 606 GEODIVERSITAS • 2019 • 41 (15) Bonis L. de et al. and shallower mandibular corpus. Thus, we think that P. dubius is really a different species. Pachycynodon crassirostris was recorded in Valbro (MP 22) by Peigné et al. (2014: fig. 26a) from a worn mandible whose wear is almost horizontal, probably indicating a hypocarnivorous diet (Peigné et al. 2014: fig. 20), but see below for Pachycynodon amphictina n. stat. Currently, the lower dentition of P. crassirostris is unknown. Pachycynodon cf. boriei or curvirostris (Filhol, 1876) (Fig. 2A) new mAteriAl. — d3-m1 (UP LPL13). diAgnosis. — Large species of Pachycynodon (length of m1 = 12 to 13.5 mm), m1 with slightly open trigonid basin. The very robust p4 and high and spaced premolars seem to be characteristic of the species. Another specimen, smaller but with the same characters, figured under the name Cynodictis leymeriei (Filhol 1876: figs 55-57) could be a female of P. boriei. The occurrence of P. boriei in the new collections of the Quercy was indicated in La Plante2 (Cirot 1992) and definitively in Valbro (Peigné et al. 2014) from a fragment of an M1. The new specimens do not firmly verify the occurrence of this species in one of the new localities in the Quercy insofar as another species of Pachycynodon, P. curvirostris (Filhol, 1876), slightly smaller than P. boriei, differs in its elongated and constricted lower premolars, and smaller p4 relative to m1 (holotype, MNHN.F.QU9208; Filhol 1876: figs 52-54). Two other mandibles (MNHN.F.QU9210 and MNHN.F.QU9216) may be related to the same species. remArk Pachycynodon boriei (Filhol, 1874) was named and later figured (Filhol 1876: figs 33, 40) as “one of the largest Carnivora ever found into the phosphate localities“(LdeB translation). Filhol 1876: 72) chose as type a mandible (Filhol 1876: fig. 40) with an associated cranium (1876: fig. 34) that is probably from the same individual. description The d3 (Fig. 2A4, A5) is single-cusped but two-rooted; the crown is slightly dissymmetric, the distal part being larger than the mesial one. It is slightly longer than high with a small medial edge (“protostylid”) and a medium high, lingually situated pacd not along the sagittal cristid as is normally the case in premolars; both mesial and distal cristids are sharp and there is a small, flat and slightly upturned talonid around the distal crown (size: 6.3 × 2.9 mm). The d4 (Fig. 2A4, A5) is thin with a high protoconid that is clearly buccally convex but slightly lingually convex and moderately sloping distally. The very oblique paraconid appears smaller than the metaconid in lingual view; the latter is slightly distally located and has a triangular horizontal section. The metalophid is acute as are the premetaconid and postmetaconid cristid. The lingual face of the metaconid is slightly convex but the mesial and distal ones are almost flat. The talonid basin has a wide and flat bottom. The cristid obliqua joins the buccal part of the base of the protoconid to the moderately high and bucco-lingually compressed hypoconid. The postmetaconid cristid joins the preentoconid cristid without a notch; postentoconid and posthypoconid cristids reach a small hypoconulid distally. A d4 from Valbro was previously described by Peigné et al. (2014). It is smaller and differs in its less pointed trigonid cuspids, lower protoconid and less transverse paraconid (size: 8 × 35 mm) The m1 is a germ out of its crypt. Despite a fissure, the crown is very well preserved (Fig. 2A1-A3). The grouped cuspids of the trigonid are massive, with the protoconid slightly higher than the other cuspids and a part of the metaconid slightly visible in buccal view. The disto-lingual face of the metaconid is triangular and flat. The distal faces of both protoconid and metaconid are in almost the same plane, sloping distally at an angle of about 45°. There is a large hypoconid occupying the entire talonid basin and gently sloping to the base of the entocristid, the latter being a series of buds in which it is impossible to distinguish an entoconid (size: 12.3 × 5.8). Judging by the size and morphology, the m1 of the mandible, UP LPL13 could be a small specimen of P. boriei but we cannot exclude, without more complete specimens and knowledge of the premolars, a large P. curvirostris. smAll species And sub-species of Pachycynodon Several taxa of quite small size (length of m1 between about 7 and 9 mm) of Pachycynodon have been described, all from the old collections of the Quercy phosphorites: Pachycynodon dubius (Filhol, 1882); Pachycynodon filholi Schlosser, 1888; Pachycynodon vulpinus Schlosser, 1899; Pachycynodon tenuis Teilhard, 1915; Pachycynodon aff. tenuis Teilhard, 1915; Pachycynodon tenuis ‘amphictoïde’ Teilhard, 1915; Pachycynodon filholi var. amphictina Teilhard, 1915; All these taxa constitute a homogeneous group in which it is easy to recognize the genus characters but difficult to interpret the tenuous differences between the component species, with size playing a role in the distinctions. The above list of taxa represents almost half of the small Pachycynodon specimens housed in the MNHN. Pachycynodon tenuis is considered by Teilhard (1915: 36) as a model for the origin of the group because of its small size, p4 higher than m1 and the thinness of the latter despite the swollen cuspids of the trigonid. Nevertheless we believe that the narrow m1 is due to the small size of the specimen, since other m1s, although larger, have the same or lower breadth/length index of m1. Other features are cited by Teilhard, such as the sharp-edged cuspids, the hollow talonid of m1, the m2 beginning to be rounded and with a distinct paraconid, but these features also are present in other species (see below), more especially as the paraconid is not all that distinct. To this we may add an entoconid that is less enlarged than the hypoconid but high, and a reduced mesial fovea in m2. Pachycynodon aff. tenuis (Teilhard 1915: pl. 4, fig. 5), where the tips of the p4 and p3 cuspids are broken off, is characterized by a high and pointed p2 separated from p3 by a short diastema. Pachycynodon tenuis ‘amphictoïde’ (Teilhard 1915: pl. IV, fig. 6) is a poorly preserved hemi-mandible with m1 and 607 Oligocene carnivorans from Quercy GEODIVERSITAS • 2019 • 41 (15) A1 A4 C1 C3 D1 E1 E3 E2 G D2 D3 C2 B1 B2 A5 B3 A2 A3 F fig. 2. — A, Pachycynodon boriei or curvirostris (Filhol, 1876): A1-A3, UP LPL13, right m1, in buccal (A1), lingual (A2), and occlusal (A3) views; A4, A5, d3-d4, in buccal (A4), and lingual (A5) views; B, C, Pachycynodon sp. 1; B, UP MGB9, left d4, in buccal (B1), lingual (B2), and occlusal (B3) views (stereo); C, UM VD16, right d4, in buccal (C1), lingual (C2) view, and occlusal (C3) views; D, Pachycynodon sp. 2, UM VD18, right d4; D1, lingual view, occlusal (D2), and buccal (D3) views; E, Pachycynodon cf. filholi, UM VD7, left hemi-mandible, in buccal (E1), lingual (E2), and occlusal (E3) views; F, Pachycynodon sp. 3, UM VD48 M1, in occlusal view; G, Pachy cynodon sp. 4, UM VD50 M1, in occlusal view. Scale bars: 5 mm. 614 GEODIVERSITAS • 2019 • 41 (15) Bonis L. de et al. in Teilhard’s figure, even if the parietal bones had been removed, probably to observe the brain endocast before Lange’s study and thus it truly belongs to a different genus. The same skull is also de facto holotype of a new species, M. piveteaui Lange, 1969, but what can we say about the species name? Viverra schlosseri is a nomen dubium and the other name, Plesictis robustus Pomel, 1848, was created for late Oligocene and early Miocene carnivores that are true Plesictis and is not available for this Quercy specimen. Unfortunately, Teilhard did use the binomen Plesictis robustus for fossils from the Quercy. In fact, true Plesictis existed in late Oligocene localities in Quercy but not in the early Oligocene ones. The first robustus is a hemi-mandible considered Plesictis robustus by Filhol (1877: 49, 50). Filhol later (1882) changed his mind and called this specimen Cynodictis leptorhynchus viverroides (1882: 59-62, pl. VI, figs 3-5, pl. VII, figs 7-10). These specimens were figured again by Teilhard (1915: pl. VIII, figs 9, 10) and are housed in the MNHN Paris. We designate here the mandible of Teilhard (1915: pl. VIII, fig. 9), MNHN.F.QU9233, as lectotype of Mustelictis robustus. The mandible corresponding to the skull is at present unknown, but another species, Mustelictis olivieri Bonis, 1997 was described from a skull associated with mandibular remains and may help in elucidating the morphology of the lower teeth of other species of Mustelictis. The new material includes a mandible probably belonging to the same genus as the type and adds new specimens of M. olivieri. Mustelictis cf. major (Teilhard, 1915) (Fig. 4A) type specimen. — Holotype by monotypy in Teilhard 1915: 60, pl. V, fig. 9; MNHN.F.QU9133. new mAteriAl. — UM VD17 part of right hemi-mandible with alveolus of p1, p2, fragments of p3 roots, p4-m1. description The right hemi-mandible VD17 is broken off in front of the p1 alveolus and behind the m1. The mandibular corpus is relatively shallow with a small mental foramen under the missing p3. The p2 is high, slender, elongate and asymmetric, similar in morphology to that of M. olivieri, although larger. The p4 is also high and slender, with a well-marked pacd and two upturned spurs mesially and distally. The m1 has a high and trenchant protoconid, a smaller paraconid and a reduced but not distally placed metaconid; the talonid displays a moderately trenchant hypoconid, a low and thin entocristid and a slightly concave basin that is open distally. These characters match those of a species of Mustelictis. Never theless, we are facing the recurring problem of identifying lower dentitions based on upper ones, and vice-versa. Mustelictis piveteaui is a skull and we do not know the mandible, whereas M. olivieri is too small. Some species have been identified as “Plesictis” in the Quercy and most of them would fit Mustelictis. Mustelictis crassirostris (Teilhard, 1915) was established on the specimen described by Filhol as Cynodictis crassirostris viverroïde (Filhol 1882: 58, 59) retaining the same name because the species Cynodictis crassirostris is still considered valid. Compared to the type specimen (MNHN.F.QU9135), VD17 differs in the shallower dentary, the relatively smaller p4 and the less trenchant talonid of m1. It also differs from M. robustus (Filhol, 1877) in the higher p2 and p4 relative to m1 and the pointed p2. The closest species seems to be M. major (Teilhard, 1915) which displays a pointed p2 and a similar p4. Thus, we identify this hemi-mandible as Mustelictis cf. major. A definitive identification will be probably possible with more complete specimens, especially with m2 (Table 3). Mustelictis aff. olivieri Bonis, 1997 (Figs 4C-E, G; 5F) type specimen. — Holotype: skull, UP MGB60, by author designation; paratype: hemi-mandible, UP MGB7. new mAteriAl. — Left m1, LPL11; left fragment of hemi-mandible, UP LPL12; right m1, UM VBOA3-4; fragment of right hemi-mandible p2-p4, UM VD12; left P4, VBO494. remArks The holotype and paratype of the species come from Mas de Got (MP 22). A skull and a hemi-mandible (paratype) were figured by Bonis (1997: figs 1, 2). New research has recovered additional specimens in other localities. description The premolars are present in UP LPL12 and UM VD12 and all of them have cutting mesial and distal edges. The p2 is dissymmetric, the mesial part being smaller than the distal table 3. — Measurements (in mm) of lower teethof Mustelictis Lange, 1969. Mustelictis p2L p2w p3L p3w p4L p4w m1L m1w m2L m2w M. olivieri MGB7 2.8 1.3 3.3 1.4 3.8 1.6 5 2.5 2.4 1.8 MGB8 – – 3.2 1.4 3.8 18 5.2 2.7 – – MGB9–––––––––– M. aff. olivieri LPL11 – – – – – – 5.6 2.5 – – LPL12 – – – – 4.1 1.9 5.4 2.4 3 2 VD12 3.1 1.3 3.2 1.5 4.1 1.8 – – – – M. cf. major VD17 4.43 2.22 – – 5 2.16 8.19 5.15 3.8 3.3 615 Oligocene carnivorans from Quercy GEODIVERSITAS • 2019 • 41 (15) one and having a more sloping mesial edge, the distal one finishing by a small upturned spur at its base. The p3, less dissymmetric than p2, displays a mesial spur; distally there is a small talonid with a small fovea surrounded by a low cristid; there is also a small pacd at mid-height on the distal edge (Fig. 4G1, G2). The p4 is similar to p3 but is larger. The carnassial is very similar to that of the type of M. olivieri but the talonid is less narrow. The m2 is larger than in the type in both absolute size and relative to m1; it has a complete trigonid with high protoconid and metaconid and small but clear paraconid, and a narrow talonid (Fig. 4B). The isolated P4 (VBO 494) figured by Peigné et al. (2014: fig. 22a) is close to that of the type specimen from Mas de Got, with a mesio-lingually elongate protocone finishing by a conic cusp, a buccal cingulum and a small mesial bulging representing a parastyle (Fig. 5F). These remains are close to the material of M. olivieri (Fig. 4F1, F2) but the small differences lead us to be cautious about the identification. They could be due to a small difference in the geological age between two localities of MP 22. Family indet. Peignictis n. gen. urn:lsid:zoobank.org:act:D0A7BD46-4A31-4346-B31A-2EC645AF3720 type species. — Peignictis pseudamphictis n. sp. by monotypy. diAgnosis. — Small Mustelida. Shallow corpus mandibulae, deep masseteric fossa distally to the level of m2 but more shallow in its upper part and gently diminishing; broad ascending ramus at its base but the distal border being oblique forward, it is narrowing up to the condyle before being broken off. Condyle situated high; small, medially advanced and slightly down directed angular process. High trigonid of m1 with protoconid far higher than the other cuspids and with an almost vertical distal face; protoconid and paraconid provide a clear buccal shearing surface; metaconid pointed and higher than the paraconid but not especially reduced or distally displaced. etymology. — From latin ictis (weasel) and dedicated to the late Stéphane Peigné for his work on Carnivora. Peignictis pseudamphictis n. gen., n. sp. (Fig. 4H) urn:lsid:zoobank.org:act:C96DF1A3-816B-4BD7-9635-28223CFDA804 type specimen. — Holotype: posterior part of hemi-mandible, UM VD1. diAgnosis. — That of the genus type locAlity. — Valbro (Lot, France). geologicAl Age. — Early Oligocene (MP 22) etymology. — Parallelism with Amphictis in the lack of m3 and elongate m2. description UM VD1 from Valbro is a piece of hemi-mandible broken in front of m1 and with an unworn m1 and alveoli of m2, (length of m1 = 5.3 mm; w of trm1 = 2.8 mm; w of tlm1 = 2; height of the corpus under m1-m2 = 5.1; length of m2 from the alveoli = 2.5 mm). The ascending ramus, whose tip is broken off, is broad at the level of the condyle but narrowing upwards because the distal border is mesially oblique (Fig. 4H1-H3). The large and deep masseteric fossa extends mesially to the level of m2 and dorsally it diminishes gently without any trace of a boundary. The condyle is situated high (38 mm higher than the base of the angular process compared with 28 mm higher in Mustelictis olivieri of similar size). The angular process is flat, mesially situated relative to the condyle and not distally projected. The m1trigonid is high with a vertical distal wall, while the protoconid and paraconid form a shearing blade. The pointed metaconid is higher than the paraconid but not clearly reduced nor distally displaced. The talonid basin is flat and surrounded by a low uninterrupted cristid in which we cannot distinguish the cuspids. Buccally, there is a well-marked notch between protoconid and hypocristid. Two alveoli, the mesial being the larger, indicate the length of m2. It is very long for an early Oligocene carnivoran. This is a parallelism with the geologically younger Amphictis. However, the overall morphology of the latter is different in having a lower and less pointed m1 trigonid. Thus, VD1 exhibits some contradictory characters, e.g., a cutting trigonid associated with a grinding talonid. We don’t know any evolutionary history of this structure. The closest genus to Peignictis n. gen. is Mustelictis, which differs in having a slightly lower m1 trigonid, a more shearing m1 talonid and a shorter m2. Suborder FELIFORMIA Kretzoi, 1945 Family nimrAvidAe Cope, 1880 Nimravus Cope, 1879 type species. — Machaerodus brachyops Cope, 1878 by original designation. remArks The genus was created by Cope (1879) for the species M. brachyops from the Oligocene of Nebraska, United States (Cope 1878). One year later, he changed the generic name to Nimravus. The type species is widespread in North America in the middle to late Oligocene (Toohey, 1959). A European species, Nimravus intermedius (Filhol 1872a) has characters that are so close to those of N. brachyops that Toohey wrote, “I cannot, with the available materials and excellent figures in Piveteau’s study, distinguish any consistent morphological differences between N. brachyops and N.intermedius”. (Toohey 1959: 95). In Europe, Nimravus was present in early and middle Oligocene strata. 616 GEODIVERSITAS • 2019 • 41 (15) Bonis L. de et al. Nimravus intermedius (Filhol, 1872) (Fig. 5E, H-J) t ype specimen . — Holotype by monotypy: left hemi-mandible with m1, MNHN.F.QU9551, in Filhol 1873. n ew mAteriAl . — Right p4 UP LPL5; left P4 UP LPL4; right m1 UP LPL1711. remArks This species is common in the old Quercy collections and constitutes the largest sample of Quercy nimravids. It is present in the three MP 22 localities, La Plante2, Mas de Got and Valbro. The number of specimens recovered from all the Quercy localities provides an overall view of the species and a good idea of the intra-specific size variation (see Peigné 2003). For example, in lion m1 length (data from Turner 1984) the extreme values are 25.2 and 30.8 mm for a mean of 28.1, the index (difference divided by mean) × 100 = 19.9. In N. brachyops the indices are 19.9 for P4 and 23.8 for m1. In N. intermedius the indices are 32.7 (P4) and 50.35 (m1), both higher than in N. brachyops. A major part of the sample of N. intermedius is recorded in the old Quercy collections without any biostratigraphic data and extends over several million years. One explanation may be a progressive change in the size during this time, with the oldest specimens being smaller than the more recent ones. An increase in size of the p4 was also noted (Peigné et al. 2014: 28). Perhaps future researches will confirm the increase in size in N. intermedius, although in North America a larger range of time did not show a similar increase in N. brachyops. description The p4 LPL5 is relatively small and low (L =11.7, w = 4.2 mm) when compared to other specimens of N. intermedius (range: L = 10.9-19.4, w = 4.5-19.2 mm in Peigné 2003). The pacd is larger than the aacd and the buccal base is underlined by a clear cingulid (Fig. 5H1, H2). The relatively small size of a lower Oligocene tooth could confirm the probable increase of size at younger localities. The P4 (UP LPL4) with L = 18.8 mm, w = 9.85, Lmts = 7, wmst = 4.7, is within the range of the smaller P4 of N. intermedius (Fig. 5E, I). The lower carnassial (UP LPL1711), with a length of 15.7 mm (Fig. 5J1-J3), is slightly smaller than the smallest m1 (L = 16 mm) in Peigné (2003). Genus Dinailurictis Helbing, 1922 type species. — Dinailurictis bonali Helbing, 1922 by original designation. remArks The genus was created by Helbing (1922) for fossils found in La Tuque (Lot & Garonne, France) in a molassic continental level which could be dated to the middle Oligocene. The remains consisted of a large scimitar-like upper canine, a P3 and a P4 most likely from the same animal. He added to the same sample a root of a large upper canine from Moissac whose size is similar to that of the La Tuque canine, but the geologic age of which is unknown because there are several levels from the middle and late Oligocene to the early Miocene in the Moissac area. Later, a fragment of maxilla with P3-P4 from the old Quercy collection was described as Nimravus intermedius major Piveteau, 1931. The latter author did not cite Helbing and probably was not aware of his article. The specimen was revised by Ginsburg (1979) and considered as the type species of the new genus Quercylurus Ginsburg, 1979. The main difference between the two genera was the size, Quercylurus being larger than Dinailurictis. The other characters (elongated protocone of P4, irregular minute pleating of the mesial crest of the paracone, m1 more robust) are linked to the larger size (slight allometry) or are also present in Dinailurictis. Dinailurictis bonali Helbing, 1922 (Fig. 5A-D) t ype specimen . — Lectotype, left upper canine (NMB-AGN 618) NHMB, from La Tuque, France, was designated by Kretzoi (1929: 1320). new mAteriAl. — UM VD46, fragment of lower canine; UM VBOA3-15, distal right humerus; UM VBOA3-17, distal left ulna; UM VBOA3-16, proximal of piece right femur. description The lower canine (UM VD46), of which part of the root and part of the crown are broken off (Fig. 5D), displays the large size and crenulated trenchant distal carena that is typical of a large sabre-toothed cat. Eofelis, Nimravus, and Eusmilus are too small for this canine. We think that it belongs to Dinailurictis bonali. UM VBOA3-15, distal right humerus, has only a part preserved, the epitrochlear arch is lacking (Fig. 5A). The size is intermediate between lion and leopard. The medial lip of the trochlea is less projecting than in modern large cats. It is may be an indication of a different way of running, Dinailurictis being less cursorial, a projected lip fixing better the elbow joint in cranio-caudal movements. The cranial fossa for receiving the olecranon process of the ulna is quite deep. The size fits Dinailurictis bonali. UM VD-45, a distal left ulna (Fig. 5C), whose size is intermediate between lion and leopard, is characterized by a high and sharp interosseous crest. UM VBOA3-16, a proximal right femur (Fig. 5B), consists of a small part of femur whose size is also intermediate between those of leopard and lion. The inter-trochanteric crest is well developed and the trochanteric fossa is deep. The small trochanter projects well laterally. Genus Eusmilus Gervais, 1876 type species. — Machaerodus bidentatus Filhol, 1872 by original designation. 617 Oligocene carnivorans from Quercy GEODIVERSITAS • 2019 • 41 (15) A1 C J1 L1 L2 M1 M2 J2 J3 K1 K2 D E H1 H2 I F G A2 B fig. 5. — A-D, Dinailurictis bonali Helbing, 1922: A, right distal humerus UM VBOA3-15 in cranial (A1), and caudal (A2) views; B, piece of right proximal femur, UM VBOA3-16, cranial view; C, left distal ulna, UM VD45, cranial view; D, left lower canine, UM VD46, buccal view; E, H-J, Nimravus intermedius; E, left P4 UP LPL4, in occlusal view; H, right p4 UP LPL5, in buccal (H1), and lingual (H2) views; I, left P4, UP LPL4, in buccal view; J, right m1, UP LPL1711, in buccal (J1), lingual (J2), and occlusal (J3) views; F, Mustelictis aff. olivieri, left P4 UM VBO494, in buccal view; G, Pachycynodon cf. dubius? m2, UM VBOA3-9, in occlusal view; K, L, Eusmilus bidentatus, right upper canine, UM VD3, in lingual (K1), and buccal (K2) views; L, right P4, UM VD2, in buccal (L1), and lingual (L2) views; M, Eofelis edwardsi, right upper canine, UM VBO454, in lingual (M1), and buccal (M2) views. Scale bars: A, B, C, D, I, M, 5 mm; E, H, J, K, L, 10 mm. 618 GEODIVERSITAS • 2019 • 41 (15) Bonis L. de et al. Eusmilus bidentatus (Filhol, 1872) (Fig. 5K, L) type mAteriAl. — Lectotype, edentulous hemi-mandible, MNHN.F.QU9475, from the Quercy phosphorites, was designated by Peigné & Brunet (2001: 660). new mAteriAl. — Fragment of upper canine UM VD3; right P4 UM VD2. rermArks Part of the tip of an elongate and bucco-lingually much compressed canine with mesial and distal trenchant and crenulated carenae is characteristic of a sabre-toothed carnivore (Fig. 5K1, K2). It is impossible to know if it is a milk tooth or a permanent one because these are very similar (Peigné & Brunet 2001). Too small to belong to Dinailurictis, smaller and more elongated than that of Nimravus, it fits the genus Eusmilus. The latter, first described as Machairodus (Filhol 1872a) includes two European species, E. villebramarensis Peigné & Brunet, 2001 from early Oligocene deposits (MP 22) and E. bidentatus (Filhol, 1872) from undated old collections of Quercy and from Soumailles (MP 21), where a complete skeleton was recovered (Ringeade & Michel 1994a, b). The canines of the first species, whether permanent or milk teeth, are larger (Peigné & Brunet 2001) and UM VD3 is closer to E. bidentatus. An isolated P4 (Fig. 5L1, L2) of which the protocone is broken off corresponds to a medium-sized feloid-like carnivoran (L = 15 mm; Wmts = 5.2; Lmts = 6.7). It has a robust parastyle and an elongate metastyle. It differs from Proailurus, which is smaller and lacks a large parastyle. Eofelis edwardsi (Filhol, 1872) is too small and does not have a parastyle. The P4 of the larger Eofelis species, E. giganteus Peigné, 2000, is unknown but probably did not have parastyle, like in the smaller species. Nimravus does not have parastyle and Dinailurictis is too large. The size and the robust parastyle indicate Eusmilus. By its size we may refer this carnassial to E. bidentatus. Taking into account the occurrence of the large E. villebramarensis in Villebramar (MP 22), which is not far away, we conclude that two different species were present in the same area and at the same time. Genus Eofelis Kretzoï, 1938 type species. — Pseudaelurus edwardsi Filhol, 1872. Eofelis edwardsi (Filhol, 1872) (Fig. 5M) t ype mAteriAl . — Lectotype, hemi-mandible, MNHN.F.QU9539, phosphorites du Quercy, designated by Ginsburg 1979: 45. new mAteriAl. — UM VBO454, right upper canine. remArks This small feloid-like carnivoran was noted by Filhol (1872a) based on a mandible and described by the same author (Filhol 1872b) based on several specimens. Peigné (2000), in a revision of the genus, identified a total of 52 remains of E. edwardsi in many collections but only three from the upper jaw, with only one canine. Until now we had not found any remains of Eofelis in more than fifty years of excavations in the Quercy. Thus, the new discovery UM VBO4511 is significant. This canine is slightly curved, bucco-laterally compressed (mesio-distal diameter = 8.8 mm, bucco-lingual = 5.6 mm; ratio L/W = 1.57) the height from the tip of the crown to the base of the root is 37.3 mm (Fig. 5M1, M2). The buccal surface of the crown is slightly convex and the lingual one is quite flat. There are slight crenulations or serrations on the distal edge as indicated by Filhol (1872b: 6) for the lower canine of E. edwardsi and Peigné (2000: 657) for the lower and upper canines of the same species. The ratio L/W for the upper canine is close to that given by Peigné (2000: table 1) for E. edwardsi (1.5). The size is slightly greater than that of a specimen figured by Ginsburg (1979: fig. 10a, b) but the shape is similar. Another species, E. giganteus Peigné, 2000, is much larger and although its upper canine is unknown, cannot fit our specimen. Thus, we attribute UM VBO454 to E. edwardsi which is firmly dated for the first time. Family indet. Genus Palaeogale Meyer, 1846 type species. — Mustela minuta Gervais, 1848-1852 by original designation. Palaeogale sectoria (Gervais, 1848-1852) remArks The history of Palaeogale is complicated (see Bonis 1981). At present, two species are recognized in Western Europe: the Oligocene P. sectoria and the early Miocene P. minuta. The former was identified in Valbro (Peigné et al. 2014: fig. 22C) from a P4. It is at present the oldest known occurrence of the species. CONCLUSION The guild of early Oligocene carnivorans in the phosphorites of Quercy corresponds to a quite complete fauna, from the small Palaeogale sectoria probably occupying a niche similar to that of a weasel, to the tiger-sized Dinailurictis bonali. In size from smallest to largest, from a weasel-like to a tiger-like: Palaeogale sectoria Peignictis pseudamphictis n. gen., n. sp. Mustelictis olivieri Mustelictis cf. major Pachycynodon cf. filholi Pachycynodon cf. dubius 619 Oligocene carnivorans from Quercy GEODIVERSITAS • 2019 • 41 (15) Pachycynodon amphictina n. stat. Amphicynodon sp. Amphicynodon typicus Pachycynodon boriei-curvirostris Cephalogale sp. Eofelis edwardsi Eusmilus bidentatus Nimravus intermedius Dinailurictis bonali The species of Pachycynodon have a dentition with inflated cusps or cuspids and a special type of wear. The shearing part of the teeth, particularly the protoconid-paraconid blade of m1, is not especially worn buccally but rather at the tip of the cuspids, more or less horizontally, like the dentition of omnivorous carnivorans such as the extant palm civets. The diet of the small Pachycynodon was probably based in a large part on fruits and seeds, even if insects or small vertebrates contributed to add some protein to the diet. This ecology implies with high probability a tropical forest where this kind of food is abundant and where these small carnivorans would help in the maintenance of the system by dispersion of the seeds. These results strengthen the conclusions from the study of the flora (Franceschi et al. 2006) regarding the Mediterranean and sub-tropical characters of the plants. The number of species of carnivorans (13) and that of Hyaenodonta (2 or 3), all carnivorous even if some of them could have been partially herbivorous, indicates the richness of the prey. A tropical forest seems to be the right environment to contain a large diversity of available prey. Few of the Orders of putative prey have been studied but one of them, Rodentia, encompassed 11 species in the locality Valbro (Peigné et al. 2014). If we add the artiodactyls, small like the Cainotheriidae (Blondel 2005), or larger, the reptiles (Augé 2006; Claude & Tong 2006), amphibians and birds (Mourer-Chauviré 2006) that are normal inhabitants of forests, the painting is complete. However, we should note the absence of taxa, which are present in old Quercy collections. For the small carnivorans, if the absence of scarce taxa like Haplogale Filhol, 1882 or Palaeoprionodon Filhol, 1880 is not surprising, that of Stenoplesictis Filhol, 1880 is astonishing because it is common in other Oligocene localities. It is the same for the family Amphicyonidae which includes more than 15 genera and more than 20 species (double if we take into account the sub-species; Springhorn 1977). The explanation could be ecological for the three studied localities, with amphicyonids perhaps living in more open environments although their skeleton, except that of the Daphoeninae, indicates that they were not good runners (Argot 2010). Another possibility is temporal and we may suppose they were later immigrants in Western Europe: all would “late travellers of the great Stampian migration” (Viret 1929). The early Oligocene carnivoran fauna is intermediate between the upper Eocene one with a few taxa, such as Paramiacis, Cynodictis and Simanphycyon, and the assemblages of middle and late Oligocene. Future research will try to pinpoint up the precise level of the changes between MP 22 and MP 26. Acknowledgements This article is greatly indebted to all the people who worked in the Phosphorites of Quercy fossil bearing localities during half a century. The bio-stratigraphic framework making it possible to follow the Oligocene fossil lineages through time is due particularly to the study by Monique Vianey-Liaud. The process of digging, washing and screening is continuing today under the leadership of Thierry Pélissié and Gilles Escarguel. We thank very much Sabine Riffaut for the blocking out of the photo backgrounds and the composition of the plates. We sometimes used the unpublished data of Edith Cirot in this article. We do not forget the great help of Christine Argot to access the collections in the MNHN Paris. We thank very much Manuel Salesa and Monique Vianey-Liaud who greatly improved the manuscript, and Lars Werdelin for the English corrections. REFERENCES Argot c. 2010. — Morphofunctional analysis of the post cranium of Amphicyon major (Mammalia, Carnivora, Amphicyonidae) from the Miocene of Sansan (Gers, France) compared to three extant carnivores: Ursus arctos, Panthera leo and Canis lupus. Geodiversitas 32 (1): 65-106. https://doi.org/10.5252/g2010n1a2 Astruc J.-g., escArguel g., mArAndAt b., simon-coinçon r. & sigé b. 2000. — Floor-age constraining of a tectonic paroxysm of the Pyrenean orogen. Late middle Eocene mammal age of a faulted karstic filling of the Quercy phosphorites, southwestern France. Geodinamica Acta Paris, 13: 271-280. https://doi. org/10.1016/S0985-3111(00)01047-0 Augé m. 2006. — Répartition des tailles chez les lézards des phosphorites du Quercy. Strata, série 1, 13: 69-78. AymArd A. 1842-1846. — Essai monographique sur un nouveau genre de mammifère fossile trouvé dans la haute vallée de la Loire et nommé Entelodon. Annales de la Société d’Agriculture, Sciences, Arts et Commerce du Puy 12: 227-267. https://gallica.bnf.fr/ ark:/12148/bpt6k408328n/f230.item AymArd A. 1850. — Du Cynodon, mammifère carnassier fossile trouvé dans les calcaires marneux de Ronzon, près du Puy. Annales de la Société d’Agriculture, Sciences, Arts et Commerce du Puy 15: 92-122. https://gallica.bnf.fr/ark:/12148/bpt6k2293835/f95.item biochrom’97 1997. — Synthèses et tableaux de corrélations in Acts of the Congress Biochrom’97. Mémoires et Travaux de l’EPHE 21: 769-805. blAinville h. m. d. de 1841-1864. — Ostéographie ou description iconographique comparée du squelette et du système dentaire des mammifères récents et fossiles pour servir de base à la zoologie et à la géologie. Tome 2. Carnassiers. J. B. Baillère, Paris. https:// doi.org/10.5962/bhl.title.127351 b londel c. 2005. — New data on the Cainotheriidae (Mammalia, Artiodactyla) from the early Oligocene of south-western France. Zoological Journal of the Linnean Society 144 (2): 145-166. https:// doi.org/10.1111/j.1096-3642.2005.00166.x bonis l. de 1966. — Arrière-crânes et moulages endocrâniens de Carnivores fossiles. Annales de Paléontologie 52 : 143-162. bonis l. de 1971. — Deux nouveaux carnassiers des Phosphorites du Quercy. Annales de Paléontologie 57 (1): 117-127. bonis l. de 1974. — Premières données sur les carnivores fissipèdes provenant des fouilles récentes dans le Quercy. Palaeovertebrata 6: 27-32. bonis l. de 1981. — Contribution à l’étude du genre Palaeogale Meyer (Mammalia, Carnivora). Annales de Paléontologie 67 (1): 37-56. 620 GEODIVERSITAS • 2019 • 41 (15) Bonis L. de et al. bonis l. de 1997. — Précisions sur l’âge géologique et les relations phylétiques de Mustelictis olivieri nov. sp. (Carnivora, Mustelidae), carnassier de l’Oligocène inférieur (MP 22) des phosphorites de Quercy (France). Geobios M.S. 20: 55-60. https://doi.org/10.1016/S0016-6995(97)80009-5 bonis l. de 2011. — A new species of Adelpharctos (Mammalia, Carnivora, Ursidae) from the late Oligocene of the “Phosphorites du Quercy” (France). Estudios Geológicos 67 (2): 179-186. https://doi.org/10.3989/egeol.40553.181 bonis l. de 2013. — Ursidae (Mammalia, Carnivora) from the late Oligocene of the “Phosphorites du Quercy” (France) and a reappraisal of the genus Cephalogale Geoffroy, 1862. Geodiversitas 35 (4): 787-814. https://doi.org/10.5252/g2013n4a4. https://doi.org/10.5252/g2013n4a4 bonis l. de, crochet J. y., rAge J. c., sigé b., sudre J. & viAney-liAud m. 1973. — Nouvelles faunes de vertébrés oligocènes des phosphorites du Quercy. Bulletin du Muséum national d’Histoire naturelle, 3ème série, Sciences de la Terre, 174: 103-113 (Sciences de la Terre 28). cirot e. 1992. — Étude phylogénétique de quelques genres d’Arctoidea de l’Oligocène eurasiatique. Comparaisons des données morphologiques et moléculaires. PhD University of Poitiers (Unpublished). cirot e. & bonis l. de 1992. — Révision du genre Amphicynodon, carnivore de l’Oligocène. Palaeontographica 220 (4-6): 103-130. cirot e. & bonis l. de 1993. — Le crâne d’Amphictis ambiguus (Carnivora, Mammalia) : son importance pour la compréhension de la phylogénie des Mustéloïdes. Comptes rendus de l’Académie des Sciences, Série 2, Mécanique, Physique, Chimie, Sciences de l’Univers, Sciences de la Terre 316: 1327-1333. https://gallica. bnf.fr/ark:/12148/bpt6k6316609v/f349.item clAude J. & tong h. 2006. — Les faunes chéloniennes fossiles du Quercy : mise à jour des connaissances. Strata, série 1, 13: 79-84. cope e. d. 1878. — Vertebrate fauna of the Miocene of Oregon. Paleontological Journal 30: 1-16. cope e. d. 1879. — On the genera of Felidae and Canidae. Proceeding of the Academy of Natural Sciences, Philadelphia 31: 168-194. https://www.jstor.org/stable/4060308 cope e. d. 1880. — On the extinct cats of America. The Amerian Naturalist 14 (12): 833-858. https://doi.org/10.1086/272672 filhol h. 1872a. — Sur les Carnassiers et les Chéiroptères dont on trouve les débris fossiles dans les gisements de phosphorites de Caylux, Frégols, Concots. Comptes rendus hebdomadaires des Séances de l’Académie des Sciences 75: 92-94. https://gallica.bnf. fr/ark:/12148/bpt6k30321/f94.item filhol h. 1872b. — Recherches sur les mammifères fossiles des dépôts de phosphate de chaux dans les départements du Lot, du Tarn, et de Tarn-et-Garonne. Première partie : carnassiers et chiroptères. Annales des Sciences géologiques 3 (7): 1-31. filhol h. 1873. — Sur les vertébrés fossiles trouvés dans les dépôts de phosphate de chaux du Quercy. Bulletin de la Société philo mathique de Paris, série 6, 8-10: 85-89. https://biodiversitylibrary.org/page/31641365 f ilhol h. 1876-1877. — Recherches sur les phosphorites du Quercy. Études des fossiles qu’on y rencontre et spécialement des mammifères. Bibliothèque de l’École des Hautes Études, Section des Sciences naturelles. Première partie, 15 (4): 1-220. Deuxième partie 16 (1): 1-338. filhol h. 1880. — Sur la découverte de mammifères nouveaux dans les dépôts de phosphate de chaux du Quercy. Comptes rendus hebdomadaires des séances de l’Académie des Sciences 90: 1579-1580. https://biodiversitylibrary.org/page/3716419 filhol h. 1882. — Mémoires sur quelques mammifères fossiles des phosphorites du Quercy. Vialelle et Cie, Toulouse: 1-140. https:// doi.org/10.5962/bhl.title.86309 filhol h. 1890. — Description d’une espèce nouvelle de Viverra fossile. Bulletin de la Société philomathique de Paris, sér. 8 II: 139142. https://biodiversitylibrary.org/page/31965626 frAnceschi d. de, le gAll c., escArguel, g., hugueney m., legendre s., simon-coinçon r., pélissié t. & sigé b. 2006. — Une paléoflore des phosphatières karstiques du Quercy (sud-ouest de la France) : première découverte, résultats et perspectives. Strata, série 1, 13: 97-101. gèze b. 1938a. — Contribution à la connaissance des Phosphorites du Quercy. Bulletin de la Société géologique de France 5 (8) : 123-146. gèze b. 1938b. — La genèse des phosphorites du Quercy. Comptes rendus hebdomadaires des Séances de l’Académie des Sciences 206: 759-761. https://gallica.bnf.fr/ark:/12148/bpt6k3158p/f759.item gèze, b., rAge J. c., vergnAud-grAzzini c., broin f. de, buffetAud e., m ourer -c hAuviré c., c rochet J. y., s igé , b., s udre J., remy J. A., lAnge-bAdré b., bonis l. de, hArtenberger J. l. & viAney-liAud m. 1978. — La poche à phosphate de Sainte-Néboule (Lot) et sa faune de vertébrés du Ludien supérieur. Palaeovertebrata 8 (2-4): 167-326. ginsburg l. 1966.—– Les Amphicyons des Phosphorites du Quercy. Annales de Paléontologie (Vert.) 52 (1): 23-44. ginsburg l. 1979. — Révision taxonomique des Nimravini (Carnivora, Felidae) de l’Oligocène des Phosphorites du Quercy. Bulletin du Muséum national d’Histoire naturelle Paris 1 (1): 35-49. https://biodiversitylibrary.org/page/55498108 ginsburg l. & morAles J. 1995. — Zaragocyon daamsi n. gen. n. sp., Ursidae primitif du Miocène inférieur d’Espagne. Comptes rendus hebdomadaires des Séances de l’Académie des Sciences, série IIa, 321: 811-815. https://doi.org/10.1016/SO753-3969(98)80003-7 kretzoi m. 1929. — Materiale zur phylogenetischen Klassification der Aeluroiden. 10th International Zoological Congress, Budapest, 2: 1293-1355. l Ange b. 1969. — Un nouveau musteliné des Phosphorites du Quercy, Mustelictis piveteaui. Comptes rendus hebdomadaires des séances de l’Académie des sciences, Série D, Sciences naturelles 268: 2870-2872. https://gallica.bnf.fr/ark:/12148/bpt6k6293308f/f761.item l Ange b. 1970. — Mustelictis piveteaui, mustélidé nouveau des Phosphorites du Quercy. Annales de Paléontologie 56 (1): 3-16. helbing h. 1922. — Dinailurictis nov. gen., ein eigenartiger Felidentypus aus dem Oligocän. Eclogae geologicae Helvetiae Basel 16 (5): 568-575. hunt r. m. Jr 1996. — Biogeography of the order Carnivora, in gittlemAn T. L. (ed.), Carnivora Behavior Biology and Evolution. Cornell University Press, New York: 485-541. hunt r. m. Jr 2009. — Long-legged pursuit carnivorans (Amphicyonidae, Daphaeninae) from the early Miocene of North America. Bulletin of the American Museum of Natural History 318: 4-93. http://hdl.handle.net/2246/5948 mourer-chAuviré c. 2006. — The avifauna of the Eocene and Oligocene phosphorites du Quercy (France): an updated list. Strata, série 1, 13: 135-147. peigné s. 2000. — A new species of Eofelis (Carnivora: Nimravidae) from the Phosphorites of Quercy, France. Comptes Rendus de l’Académie des Sciences Paris 330 (9): 653-658. https://doi. org/10.1016/S1251-8050(00)00199-3 peigné s. 2001. — A primitive nimravine skull from the Quercy fissures, France. Implications for the origins and evolution of Nimravidae (Carnivora). Zoological Journal of the Linnean Society 132 (4): 401-410. https://doi.org/10.1111/j.1096-3642.2001. tb02467.x peigné s. 2003. — Systematic review of European Nimravinae (Mammalia, Carnivora, Ninravidae) and the phylogenetic relationships of Palaeogene Nimravidae. Zoologica Scripta 32: 199-229. https://doi.org/10.1046/j.1463-6409.2003.00116.x peigné s. & bonis l. de 1999. –—Le premier crâne de Nimravus (Mammalia, Carnivora) d’Eurasie et ses relations avec N. brachyops d’Amérique du Nord. Revue de Paléobiologie 18 (1): 57-67. peigné s. & brunet m. 2001. — Une nouvelle espèce du genre Eusmilus (Carnivora: Nimravidae) de l’Oligocène (MP 22) d’Europe. Geobios 34 (6): 657672. https://doi.org/10.1016/ S0016-6995(01)80027-9 621 Oligocene carnivorans from Quercy GEODIVERSITAS • 2019 • 41 (15) p eigné s., v iAney -l iAud m., p élissié t. & s igé b. 2014. –– Valbro : un nouveau site à vertébrés de l’Oligocène inférieur (MP22) de France (Quercy). I – Contexte géologique ; Mammalia : Rodentia, Hyaenodontida, Carnivora. Annales de Paléontologie 100: 1-45. https://doi.org/10.1016/j.annpal.2013.11.002 piveteAu J. 1931. — Les chats des Phosphorites du Quercy. Annales de Paléontologie 20: 107-163. piveteAu J. 1942 — Études sur quelques mammifères des Phosphorites du Quercy. I. Le genre Stenoplesictis. Annales de Paléontologie 30: 63-72. piveteAu J. 1962. — L’encéphale de Viverravus angustidens, Miacidé des Phosphorites du Quercy. Annales de Paléontologie 48: 165-175. r Age J.-c. 2006. — The lower vertebrates from the Eocene and Oligocene of the phosphorites du Quercy (France) an overview, Strata, série 1, 13: 161-173. rémy J. A., crochet J.-y., sigé b., sudre J., bonis l. de, viAneyliAud m., godinot m., hArtenberger J.-l., lAnge-bAdré b. & comte b. 1987. — Biochronologie des phosphorites du Quercy : mise à jour des listes fauniques et nouveaux gisements de mammifères fossiles. Münchner Geowissenschaftliche Abhandlungen A (10): 169-188. ringeAde m. & michel p. 1994a. — Une nouvelle sous-espèce de Nimravidae (Eusmilus bidentatus ringeadei) de l’Oligocène inférieur du Lot et Garonne (Soumailles, France) : étude préliminaire. Comptes rendus hebdomadaires des Séances de l’Académie des Sciences 318: 691-696. https://gallica.bnf.fr/ark:/12148/ bpt6k65095354/f89.item ringeAde m. & michel p. 1994b. — À propos de l’Eusmilus (Eusmilus bidentatus ringeadei Ringeade et Michel 1994) de Soumailles, canton de Duras (Lot et Garonne, France) : étude descriptive. Paleo 6: 5-37. schlosser m. 1888. — Die Affen, Lemuren, Chiropteren, Insectivoren, Marsupialier, Creodonten und Carnivoren des europäischen Tertiärs und deren Beziehungen zu ihren lebenden und fossilen aussereuropäischen Verwandten. Beiträge zur Paläontologie Österreich Ungarns und des Orients I. Theil 6: 1-224. II Theil 7: 1-162. schlosser m. 1899. — Ueber die Bären und bärenänlichen Formen des europäischen Tertiärs. Palaeontographica 46 (4) : 95-148. s igé b., A guilAr J. p. & m ArAndAt b. 1991. — Extension au Miocène inférieur des remplissages phosphatés du Quercy, la faune de vertébrés de Crémat (Lot, France). Geobios 23 (4): 497-502. https://doi.org/10.1016/S0016-6995(06)80251-2 springhorn r. 1877. — Revision fer alttertiären europäischen Amphicyonidae (Carnivora, Mammalia). Palaeontographica A 158 : 26-113. teilhArd de chArdin p. 1915. — Les carnassiers des phosphorites du Quercy. Annales de Paléontologie 9: 103-190. toohey l. 1959. — The species of Nimravus (Carnivora, Felidae). Bulletin of the American Museum Natural History 118: 71-112. http://hdl.handle.net/2246/1972 t rouessArt e. 1904. — Catalogus mammalium tam viventium quam fossium quinquinale supplementum. Vol. 1. R. Friedländer & Sohn, Berlin: 1-546, 224. https://doi.org/10.5962/bhl.title.61437 turner A. 1984. — Dental sex dimorphism in European Lions (Panthera leo L.) of the Upper Pleistocene: palaeoecological and palaeoethological implications. Annales Zoologici Fennici 21: 1-8. https://www.jstor.org/stable/23734092 viAney-liAud m. & legendre s. 1986. — Les faunes des phosphorites du Quercy : principes méthodologiques en paléontologie des mammifères; homogénéité chronologique des gisements de mammifères fossiles. Eclogae geologicae Helvetiae 79 (3): 917-944. http://doi.org/10.5169/seals-165856 viret J. 1929 — Les faunes de mammifères de l’Oligocène supérieur de la Limagne bourbonnaise. Mémoires de l’Université de Lyon 47: 1-305. wAng b. & Qiu z. 2003. — Notes on Early Oligocene Ursids (Carnivora, Mammalia) from Saint Jacques, Nei Mongol, China. Bulletin of the American Museum of Natural History 279: 116-124. https://doi.org/10.1206/0003-0090(2003)279<0163:C>2.0.CO;2 w olsAn m. & l Ange -b Adré b. 1996. — An arctomorph carnivoran skull from the Phosphorites du Quercy and the origin of procyonids. Acta Palaeontologica Polonica 41 (3): 277-298. Submitted on 10 February 2019; accepted on 16 April 2019; published on 10 September 2019.