# The early Oligocene Caniformia (Carnivora, Mammalia) from the standard level MP23 in the ‘Phosphorites du Quercy’, Occitanie, southwestern France

Louis de Bonis, Axelle Gardin, Gilles Escarguel

► To cite this version:

Louis de Bonis, Axelle Gardin, Gilles Escarguel. The early Oligocene Caniformia (Carnivora, Mammalia) from the standard level MP23 in the ‘Phosphorites du Quercy’, Occitanie, southwestern France. Geobios, 2025, 10.1016/j.geobios.2025.07.001. hal-05220646

HAL Id: hal-05220646

https://hal.science/hal-05220646v1

Submitted on 23 Aug 2025

HAL is a multi-disciplinary open access
archive for the deposit and dissemination of sci-
entific research documents, whether they are pub-
lished or not. The documents may come from
teaching and research institutions in France or
abroad, or from public or private research centers.

L'archive ouverte pluridisciplinaire HAL, est
destinée au dépôt et à la diffusion de documents
scientifiques de niveau recherche, publiés ou non,
émanant des établissements d'enseignement et de
recherche français ou étrangers, des laboratoires
publics ou privés.

<figure><img src="image_1.png" /></figure>

Distributed under a Creative Commons Attribution 4.0 International License
<figure><img src="image_2.png" /></figure>

Research Paper

# The early Oligocene Caniformia (Carnivora, Mammalia) from the standard level MP23 in the 'Phosphorites du Quercy', Occitanie, southwestern France ☆

Louis de Bonis<sup>a,*</sup>, Axelle Gardin<sup>a,b</sup>, Gilles Escarguel<sup>c</sup>

## ARTICLE INFO

**Article history:**

Received 9 December 2024

Accepted 10 July 2025

Available online xxxx

**Keywords:**

Biostratigraphy

Paleoenvironment

Palaeobiology

Grande Coupure

Phosphatières du Quercy

Konzentrat-Lagerstätten

## ABSTRACT

This article describes Caniformia carnivorans newly found in southwestern France fossiliferous localities called 'Phosphorites du Quercy' and dated to the MP biochronological scale standard level MP23 (reference locality: Itardies, 30.56 ± 0.18 Ma; early Oligocene). In Europe, the Eocene-Oligocene transition is marked by a significant paleobiological turnover known as Stehlin's 'Grande Coupure', representing a major shift in the evolutionary history of Caniformia. Before the 'Grande Coupure', a nearly singular, small and primitive Amphicyonidae was present in the late Eocene of Europe, while a wide diversity of taxa and morphologies emerged by the early Oligocene. When compared to the material from the 19th century 'Old Collections from the Quercy', the well-dated new material provides biochronological information to species first described without any time constraint, thus precising their position in the evolutionary process. We describe the most primitive musteloid taxon ever found in Europe. Other Caniformia taxa show ecological niches absolutely unknown in extant faunas, with horizontal wear facets in the check teeth. The most abundant taxon shows adaptations to climbing; the high diversity of small carnivores and glirid rodents supports the hypothesis of an at least partially forested environment. The high number of individuals including juveniles of species of amphicynodontids favours the hypothesis of individuals buried in their den and suggests a potential gregarious behaviour.

© 2025 The Author(s). Published by Elsevier Masson SAS. This is an open access article under the CC BY license (http://creativecommons.org/licenses/by/4.0/).

## 1. Introduction

The 'Causses du Quercy', in southwestern France, is a Jurassic limestone plateau which emerged during the Campanian-Maastrichtian, ca. 70 Ma (Mazières, 1956; Pélissié et al., 2021) and functioned as a karst from the early Eocene (Legendre et al., 1992) to, for the latest, the early Miocene (Sigé et al., 1991), with hundreds of karstic fissures being successively opened and filled by sediments. Most fissures contained a phosphatic mineral called phosphorite used in agriculture as a fertilizer. The 'Phosphorites du Quercy' were exploited between 1870 and 1907, leading to the discovery in many localities of vertebrate, invertebrate and plant fossils which were collected or purchased from the workers for private or official collections, constituting the so-called 'Old Collections from the Quercy' (Durand-Delga, 2006; Pélissié et al., 2021).

Unfortunately, all the specimens found at that time were mixed without any precise location and geologic age indication (Legendre et al., 1997; Pélissié and Sigé, 2006). Recorded for the first time in 1871 (Daubrée, 1871; Trutat, 1871), the first fossils from the Quercy were published one year later (Delfortrie, 1872; Gervais, 1872), then principally by Filhol (1873, 1876–77) and after them several authors until recently.

A new phase of paleontological research began in 1965 and continues today, 60 years later. A group of researchers from various French institutions (Montpellier university, UPMC, MNHN Paris, Lyon and Poitiers universities) carried out new excavations and demonstrated that almost every fissure with infillings in *situ* allows getting a homogeneous fauna dated biochronologically (Vianey-Liaud and Legendre, 1986). Fissures after fissures, the recovered faunas illustrate ~30 myr of vertebrate evolution and changes in the climate-environment-biodiversity relationships (Pélissié and Sigé, 2006). With more than 200 fossil assemblages described so far, the Quercy faunas took a large part in establishing the system of fossil Mammal Paleogene (MP) European standard levels, some

<footnote><sup>a</sup>Laboratoire de Paléontologie, Evolution, Paléoécosystèmes et Primatologie (PALEVOPRIM) UMR 7262 CNRS, Université de Poitiers, Poitiers, France</footnote>

<footnote><sup>b</sup>Department of Earth and Environmental Sciences, University of Michigan, Ann Arbor, USA</footnote>

<footnote><sup>c</sup>Université Claude Bernard Lyon 1, LEHNA UMR 5023, CNRS, ENTPE, F-69622 Villeurbanne, France</footnote>

<footnote>* Corresponding editor: Laurent Marivaux.</footnote>

<footnote>* Corresponding author.</footnote>

<footnote>E-mail address: louis.de.bonis@univ-poitiers.fr (L. de Bonis).</footnote>
sites being designated as reference localities for this biochronolog-
ical system (Rémy et al., 1987; BiochroM'97, 1997). They are also
used for other analyses including quantitative biochronology
(Legendre and Bachelet, 1993; Escarguel et al., 1997), paleobotany
(Francheschi et al., 2006), and macroevolution/macroecology
(Legendre, 1989; Escarguel and Legendre, 2006; Escarguel et al.,
2008; Weppe et al., 2023).

The Mammalian order Carnivora is of course present in the old
as well as the new collections (Bonis et al., 2019). The present arti-
cle aims to give a comprehensive study of the Caniformia carnivo-
rans from the MP23 localities of the Quercy (reference locality:
Itardies, early Oligocene, 30.56 ± 0.18 Ma; Escarguel et al., 1997).
In the order Carnivora, the sub-order Caniformia includes nowa-
days the families Canidae (dog, wolf), Ursidae (bear), Mustelidae
(wolverine, weasel), Procyonidae (raccoon), Ailuridae (red panda)
and Mephitidae (skunks).

The reference locality of the standard level MP23 is Itardies (Quercy), whose preliminary list recognises only three caniform species (BiochroM'97: 790-791). The other Quercynian localities of MP23 with known carnivoran remains are Gardiol 3, Mounayne, Pech Crabit, and Roqueprune 2, while no carnivoran remains have been found so far in Coulou, Mas de Gaston, Mêge and Raynal, also from the same standard level in the Quercy area.

**2. Material and methods**

The new material of fossil mammals from the 'Phosphorites du Quercy' is currently housed in the University of Poitiers, the University of Montpellier, and, partially, in the MNHN and the University of Lyon. We took all measurements with a Matsumoto calliper to the 1/100° mm given to the 1/10° mm (Fig. 1).

**Abbreviations:**

**Anatomy:** D, deciduous upper molar; d, deciduous lower molar; M, upper molar; m, lower molar; P, upper premolar; p, lower premolar; aac, anterior (mesial) accessory cusp; aacd, anterior (mesial) accessory cuspid; pac, posterior (distal) accessory cusp; pacd, posterior (distal) accessory cuspid; mcc, mesial cingulum cusp; dcs, distal cingulum shelf; mccd, mesial cingulid cuspid; dacd, distal accessory cuspid; dcsd, distal cingulid shelf.

**Measurements:** L, length; w, width; w1, mesial width of M1; w2, distal width of M1; tr, trigonid; tl, talonid; mts, metastyle.

**Biochronology:** MP, Mammal Paleogene standard level; MN, Mammal Neogene biozonation.

**Quercynian localities:** GAO, Gardiol 3; CAV, Cavalé; ITD, Itardies; MOU, Mounayne; PC, Pech Crabit; ROQ, Roqueprune 2; VAL, Valbro.

**Institutions:** BMNH, British Museum of Natural History, London, UK; MNHN, Muséum national d'Histoire naturelle Paris, France; NMB, Naturhistorische Museum Basel, Switzerland; UM, Université de Montpellier, France; UPMC, Université Pierre et Marie Curie (Paris 6), France; UP, Université de Poitiers, France.

**3. Systematic palaeontology**

Mammalia Linnaeus, 1758
Carnivora Bowdich, 1821
Caniformia Kretzoi, 1943
Arctoidea Flower, 1869
"Amphicynodontidae" Simpson, 1945.

**Remarks:** The type-genus of this taxon – either a ‘family’ or a “group” – is *Amphicynodon* Filhol, 1881. “Amphicynodontids” are primitive, generally small-sized arctoid carnivorans with slightly elongate skull showing moderate sagittal and lambdoid crests. They display a bony unpartitioned tympanic bulla principally built

by the ectotympanic (Type A: Hunt, 1974, 1998) and not much
expanded caudally. The dental formula is 3/3 Incisors, 1/1 Canines,
4/4 Premolars, 2-3/2-3 Molars, rare specimens of *Amphicynodon*
may bear 3 upper molars or 2 lower molars. The incisors are small,
on a straight line, I1 and I2 having almost the same size and I3
being slightly larger. The canines are moderately developed. P1,
P2 and P3 are simple and do not have accessory cusp. P1 is very
small and one-rooted; P2 and P3 are two-rooted. P4 is robust with
a large protocone in front of the mesial part of the paracone and
moderately extended posterior; two crests, mesial and distal, run
to the base of the paracone, the latter being prolonged distally as
a cingulum. The buccal cingulum projects mesially as a small para-
style with, sometimes, a tiny cusp. The paracone is large, its lingual
face is flat and the buccal is convex or sometimes displays a crest
from the tip to the cingulum, and is almost pyramidal; the metas-
tyle is short. M1 is triangular; the paracone, almost pyramidal, is
larger than the metacone; the buccal cusps are linked to the proto-
cone by the crests of the trigone and it is sometimes possible to
distinguish a metaconule on the distal one; the protocone is sur-
rounded by a large cingulum continuing along the whole crown.
M2 is oval, three-rooted, and displays almost the same pattern as
M1, although there is a larger size-difference between the buccal
cusps and there is no metaconule. The dentary is variable, between
slightly deep to shallow with a straight lower border to slightly
convex. The posterior mandibular process is situated below the
level of the tooth row, while the articular condyle is situated at
the level or slightly higher. The premolars are simple, except p4,
which bears a small pacd, and are two rooted except p1 which is
one-rooted. The m1 is robust, low and has a grouped trigonid,
the protoconid of which is taller than p4 and the metaconid is well
developed; the talonid is broad with an elongate hypoconid fol-
lowed by a cristid surrounding the rest of the crown with some-
times a lingual thickening indicating an entoconid; the talonid
basin is moderately deep. The third molar is a small round button
with a central basin surrounded by a continuous cristid.

Some other genera group with *Amphicynodon*. In Eurasia, *Pachycynodon Schlosser, 1888* differs by smaller m1 relative to the premolar row, p4 equal to or taller than the protoconid of m1. In Asia, there are *Amphicticeps Matthew and Granger, 1924* from the Oligocene of the Hsanda Gol Fm. and *Wangictis Bonis et al., 2019* from Ulanmannei, East of Saint-Jacques, Nei Mongol, China. Several genera described from Eocene, Oligocene and early Miocene layers of North America such as *Allocyon Merriam, 1930*, *Kolponomos Stirton, 1960*, *Parictis Scott, 1893*, *Subparictis Clark and Guensburg, 1972*, *Drassonax* Galbreath, 1953, and *Eoartos Wang et al., 2023* were considered as close to the amphicynodonts (Wang et al., 2023). The systematics of the group changed from one author to another one: *Canidae* (Aymard, 1846, 1850; Merriam, 1930; Simpson, 1945; Clark and Guensburg, 1972), *Ursidae* (Ginsburg, 1966; *Prothero and Emry, 1996*; *Hunt, 1998*; *Finarelli, 2008*; *Wang et al., 2005, 2023*; *Wang et al., 2023*), *Mustelidae* (Scott, 1993), *Amphicynodontidae* (Simpson, 1945 as *Amphicynodontinae*; *Prothero and Emry, 1996*; *Wang et al., 2023*), or *Procyonidae* (Stirton, 1960). The much-derived *Allocyon* and *Kolponomos* are not very helpful in recognizing the shared derived characters uniting the ‘*Amphicynodon* group’ with one of these families (Merriam, 1930; Stirton, 1960).

The auditory area with the tympanic bulla is the main area used
to separate families within the Carnivora. Type A may be consid-
ered the most primitive. It is present in extant Carnivora ursids, ail-
urids, otariids, odobenids, lutrines and mephitine mustelids (Hunt,
1974, 1998), without indicating any special link of ancestry; it is
the basic bulla of arctoids, and other families such as other muste-
lid and procyonid ancestors must have had the same bulla even
though the extant forms are different. Presently, amphicynodon-
tids are often considered as a subfamily of the ursid family, and
<figure><img src="image_3.png" /><figcaption>Fig. 1. Tooth measurements used in this work.</figcaption></figure>

they "are the ancestral ursid stock from which ultimately the other subfamilies evolved; hence "Amphicynodontinae" is a polyphyletic category retained for convenience until the members of the group are better known" (Hunt, 1998: p. 178). Some amphicynodontids were also possibly a stem group for other families than ursids, mustelids, mephitids, ailurids and procyonids. A cladistic analysis of the arctoids (Finarelli, 2008) brings *Amphicynodon* near *Ursus* on the base of 3 characters (12, 31, 40 in Finarelli, 2008):

* 12- Condyloid canal absent, but it is present in *Amphicynodon* (Cirot, 1992: fig. 26);

* 31- the bulla not expanded posteriorly - it is true but the bulla of *Amphicynodon* is more inflated than that of *Ursus* (derived character) and the no-expanded bulla is primitive;

* 40- P4 medial shelf moderately developed - it is true but it is a common characteristic.

On the other hand, ursids are characterized by a lengthening of the molars present in the oldest known Ursinae, *Ursavus* from the European early Miocene. The amphicynodontids display a reverse trend shown by the reduction of M2 and, sometimes, that of the distal part of M1 in *Amphicynodon* (Bonis et al., 2019: fig. 1D)
and *Amphicticeps* (Matthew and Granger, 1924: fig. 4; Wang et al., 2005: figs. 2, 8) as well as m2 in *Amphicticeps* (Wang et al., 2005: fig. 8). Thus we agree that amphicynodontid is a polyphyletic group, but it seems preferable here to provisionally consider the group as “family-like” amphicynodontids until we can identify the putative ancestors of the other arctoid families. If not, several taxa would be familyless orphans.

Genus **Amphicynodon** Filhol, 1881

**Type species:** Cynodon velaunus Aymard, 1846 (by original designation).

**Diagnosis** (modified from Bonis et al., 2019): "Amphicynodontid" with a relatively low skull with a faint sagittal crest slightly more developed distally; moderately elongate muzzle; ossified bulla of type A; shallow suprameatal fossa; alisphenoid canal present, post-glenoid canal present. Dental formula 3/3I, 1/1C, 4/4P, 2-3/3M. Simple premolars P1-P3 and p1-p3; short P4 with robust protocone and short metastyle; triangular M1 with tiny parastyle, without paraconule but with metaconule; more or less reduced oval M2; p4 with pacd or not; grouped and not very tall trigonid of m1, metaconid as tall as the paraconid; not very elongate m2 with paraconid present or not, metaconid as tall or taller than the protoconid; small rounded m3.

**Remarks:** Whatever its taxonomic assignment at family or subfamily levels, *Amphicynodon* belongs to a quite homogenous group arriving in western Europe during Stehlin's 'Grande Couvre', i.e., at the beginning of the European Oligocene. As far as we know, they have never been firmly recorded in Eocene layers in western Europe. Amphicynodontids were the major group of small carnivores with a large number of species. This large number raises the problem of the different kinds of diets among the taxa. *Amphicynodon* is known in Asia with *A. teilhardi* (Matthew and Granger, 1924) in Mongolia. This species could be a migrant from western Europe (Wang et al., 2023) or maybe a vestigial population that stopped there its migration from east to west. More surprising is the presence of *Amphicynodon major* (Clark and Guensburg, 1982) in North America. *A. major* is close in size to *A. crassirostris*, the largest European species. Nevertheless, there is a great difference between the former and all European species: its metaconid of m1 is backwardly shifted but remains very large and keeps a contact with the paraconid (Wang et al., 2023: fig. 73C). There is a slight shifting in *A. gracilis* but the metaconid is not separated from the paraconid. Another North American carnivoran displays characters close to amphicynodont ones. *Drassonax harpagops* Galbreath, 1953 comes from the early Oligocene of USA. Its lower jaws have a striking similarity with those of *Amphicynodon leptorhynchus* (Wang et al., 2023: fig. 74). The lower cheek teeth do no significantly differ from those of the European ones. The upper cheek teeth of *Drassonax* display some morphological variations (Wang et al., 2023: figs. 74AB, 75AB). The P4 of the maxilla UNSM 122001 is slightly different and more robust than that of *Amphicynodon leptorhynchus* but M1 is clearly less extended buccolingually and M2 is more reduced. The M1 of UCM 45181 is also very different from that of *A. leptorhynchus*. Nevertheless, the presence of a link between European and some American fossils claimed by Wang et al. (2023) is highly probable. The type species was cited under the genus *Cynodon* (Aymard, 1846) and published later (Aymard, 1850) with the same name (Cirot 1992).

*Amphicynodon leptorhynchus* (Filhol, 1874)
*Figs. 2, 3 (A, B, F, G), 4*

**Lectotype by designation** (Cirot and Bonis, 1992: p. 24): Left hemimandible (p2-m3), MNHN QU 9188; unknown locality, Phosphorites of Quercy (Filhol, 1876: figs. 73-75, 85).

**Paraleptotype by designation** (Cirot and Bonis, 1992: p. 24): Left hemimandible (p2-m3), MNHN QU 9198; unknown locality, Phosphorites of Quercy (Filhol, 1876: fig. 78).

**Material:** *Itardies* (UP-ITD) 2 md c-m3, 2b skull, 8 md p2, p4-m2, 9 md p1-m2, 18 md m1-m2, 23 md p2-p4, m2-m3, 25 P3-M1, 27 md d3, 32 md p3-m2, 40 md p1-m1, 40 b md c-m3, 40c md p2-p3, 42 mx P3-M2, 50 crane M3, 51 crane M3, 78 md p1-m2, 110 mx P4-M2, 123 md p4, 124 P4-M1, 127 md i2-m3 weathered, 128 p3-m1, 194 md p2-m2, 206 md c p2, p4-m3, 207 P3-M2, 231 md c, p2-p3, 240 md p9-m1, 241 md c, p2-m3, 260 mx P4-M2, 265 md c-, p2-m3, 311 skull in 2, I1-M1, 313 skull, 315 md c-m3, 316 md c-m2, 318 mc m2-m3, 325 skull, 348 skull, 353 skull, 355 md i3-m3, 355 b skull + md, 360 skull, 361 md c-m2, 362 md p2-m3, 362 md p2-m3, 363 md c-m2, 364 md c, p2-m2, 368 md p3-m1 cas, m2m3, 370 md p2, p4, 371 md p4-m1, 400 P4-M2, 406 md c-m3, 425 M1 ou 2, 440 skull, 441 md c-m3, 467 P4, 470 md p4, m1-m2, 486 md p2-m2, 496 p4-m2, 497 md d3-d4, 520 skull, 533 skull, 534 skull, 535 md m1-m3, 555 md p3-m2, 561 skull, 564 md m2, 565 md m2, 566 M1, 569 md d c, p2, p4-m2, 571 md p1-m2, 574 P2-P3, 575 m1, 582 md i-p3, 584 skull, 589 max D3-D4-P4, 604 md p1-m3, 618 md p4-m2, 619 amphi without m3, 620 md p2-m3, 621 mdp2-m2, 634 md p2, 639 md p2-p3, 650 max P4-M2, 661 md p4, 688 M1, 589 max D3-D4-P4, 765 md p1, p3, 767 md m2, 769 md m1, 771 md m2, 650 max P4-M2, 765 md p1, p3, 767 md m2, 769 md m1, 771 md m2, 775 md m2 cas m2, 777 md p2-p4. +775 md m2 cas m2 777 md p2-p4, 783 md, 784 md m2-m3, 785 d3-d4, 788 M1, 790 P4-M1, 792 frg md m1-m2, 793 P2-P3, M1-M2, 797 frg md p4-m1, 798 P2-P3, M1-M2, 809 frg md m2-m3, 847 d4, 857 md p2-m2, 874 skull, 875 md, p1-m1 + skull, 895 md m1-m2, 898 md p4, m1-m2, 914 md p2-m2, s916 md m1-m2, 927 md p1-p4, 997 skull, 1017 skull + md, 1018 skull, 1021 md p1-m2, 1022 md m1-m2, 1030 max P2-M2, 1075 md c, p2-m3, 1077 md p2-m2, 1099 skull, 1102 skull, 1105 md m2, 1114 skull, 1119 skull, 1121 P3-M1, 1122 md c-p1, 1124 md p2-m3, 1126 md c, p2-m1, 1127 md m2 sans m3, 1137 M1, 1142 md c-m3, 1143 md p2, p4-m2, 1145 m ca, -MP4-M1d c-m3, 1146 md c, p3, m1, 1147 md p2-m2, 1148 md p4-m3, 1169 p4-m3, 1180 max C, D1-D4 lepto, 1186 P3-M2, 1189 P2-P4, 1190 md m1 m2, 1192 md m1-m3, 1193 P4, M2, 1194 P3-P4, 1195 md m1, 1203 md, 1204 md p3-m1, 1229 md m1, 1246 md m1, 1248 md m1 m2-m3, 1260 max P4-M2, 1261 skull, 1262 skull, 1264 md p2-p3 + m1, 1269 md p2-m, 1273 md c-m1, 1274 md m1-m21292 md p4-m2, 1403 m1, 1427 max M1-M2, 1431 m1, 1432 m1, 1445 P4-M2, 1450 P2-P4, 1452 d4, 1453 skull, 1541 md p2, p4-m2, 1590 md c-m3, 1591 P3, 1592 P3-M1, 1747 md m2, 1748 md m2-m3, 1751 m1, 1758 md p1-m1, 1762 md p3-m1. 1764 p4, 1765 md m2, 1766 md m1 cas stat nas 1782 P4, 1783 1784 max P4-M2, 1786 P4, 1788 M1, 1795 p3-4, 1803 usure, 1803 frg md m1, 813 p1, 1817 p, 1820 M2, 1821 P4, 1822 P4, 1824 P4, 1825 M2, 1826 p digested, 1827 M2, 1828 M2, 1829 m-M3, 1830 m1, 1831 m1, 1832 m1, 1833 m1, 1691 md d4, 1766 md m1 cas stat nas, 1960 skull, VF1045, VF1159, VF11259, VF11065, vf10360, VF10242, VF10001, VF10594a, VF10594b, VF10148, VF10863, VF11107, VF10202. **Mounayne** (UP-MOU) 15 p2-p4. **Pech-Crabit** (UP-PC) 2 md p4-m1. **Gardiol 3** (UM-GAO) 509 md m1, 465 m1, 466 M1, 467 M1, 468 M1, 468 M1, 470 M1, 494 m1, 473 mx P4-M1, 272 m1. **Roqueprune 2** (UP-ROQ) 8 m1, 16 D4, 33 P1, 33 P134 P4-M1.

**Measurements:** see Table 1.

**Diagnosis:** Medium-sized *Amphicynodon* (average body-mass estimated 1.9 kg); shallow dentary: deepness of the dentary at
<figure><img src="image_4.png" /><figcaption>**Fig. 2.** *Amphicynodon leptorhynchus*, ITD60. **A.** Skull in dorsal view. **B.** Skull in ventral view. **C.** Maxilla in ventral view. **D.** Back skull in ventral view. Scale bars: 1 cm.</figcaption></figure>

m1-m2 from 9.8 mm to 12.3 mm; two mental foramina, a small one beneath the mesial root of p2, another one beneath p3; lower border slightly convex; angular process long, thin and hollowed medially as a gutter for the muscle pterygoideus medius. Dental formula 3/3I, 1/1C, 4/4P, 2-3 or 3-3M. Lower premolars without marked cingulid except sometimes in p4, a small anterior spur and a distal cristid encircling a small basin; the latter displays also a small slightly buccally shifted pacd; straight and abruptly sloping p4 mesial edge. The m1 is relatively low and robust; the trigonid groups inflated cuspids; there is only a short cingulid along a part of the base of the buccal para-protoconid blade; the paraconid mesial edge is lingually obliquely directed; the metaconid is globular as tall or taller than the paraconid and contacting the latter to close the trigonid valley at its third of height; the hypoconid is bulky and extends to the middle of the distal face of the crown; its lingual face slopes gently to the buccal base of the entoconid cristid which is almost vertical, as high as the hypoconid and prolonged to the middle of the rear but stopping before contacting the hypoconid, thus the narrow talonid basin is open distally. There is a vertical, moderately developed shearing wear facet along the buccal faces of para- and protoconid, and, sometimes, horizontal facets on the top of the cuspids. The m2 is rectangular or quadrate; there is no paraconid, protoconid and metaconid of almost the same size, associated in front of the crown, but not linked by a cristid, with a minute mesial fossa or without mesial fossa; distinct hypoconid and entoconid posteriorly by a cristid closing the small talonid basin. One-rooted, round and small m3 with a central basin circled by a cristid. Skull relatively low with a faint sagittal crest slightly more developed distally; moderately elongate snout; ossified bulla of type A, shallow suprameatal fossa; alisphenoid canal and post-glenoid canal present; simple premolars P1-P3; short P4 with robust protocone and short metastyle; triangular M1 with tiny parastyle, without paraconule but with metaconule; more or less reduced and oval-shaped M2. Differs from *A. velaunum* (Aymard, 1846) by less robust premolars, weaker cingulum and cingulid, less pointed cusps and more symmetrical M1.

**Description:**

**Skull** (Fig. 2). Several specimens from Itardies allow a quite precise knowledge of the skull despite the bad preservation of some parts, especially the ventral posterior part which is often crushed and destroyed. It is elongated with a thin muzzle; the nasals are narrow and stop on a line when contacting the frontal; the zygomatic arcades are relatively thin and widely distant from the skull; there is a sagittal crest, moderately developed or faint, depending probably on sexual dimorphism, but always more marked distally and prolonged by the thicker and more extended lateral lambdoid crests. Laterally, the infra-orbital foramen appears slightly oval and placed above the P3-P4 limit; the anterior lacerate foramen is lowly situated; the glenoid process is only slightly curved, leaving a possibility of lateral movements to the mandibular condyle; the auditory meatus is large and rounded without any auditory tube; the mastoid process is relatively small as is the paroccipital process. In ventral view, the auditory bulla is inflated but we cannot distinguish the different elements (ectotympanic, rostral and caudal entotympanics). On rare specimens we can observe the alisphenoid canal, the postglenoid foramen, and some other foramina (Fig. 2). The suprameatal fossa is shallow but present (Schmidt-Kittler, 1981: fig. 1), as in other arctoids such as ailurids, mustelids, procyonids and, even if it is lost in adults, young ursids.

**Upper dentition** (Table 1). The three incisors are small and on a slightly arched line. Their size increases from I1 to I3 and all are mesiodistally flattened; their morphologies are slightly different, the crowns of I1 and I2 being narrow and shovel-like while that of I3 is pointed and slightly curved. The three have a lingual talon, that of I3 being relatively larger and pointing as a cusp. The upper canine is separated from I3 by a diastema; the crown is only slightly curved with an oval section; there is no mesial or distal cutting ridge (ITD1704).

P1 is close to the canine but separated from the next premolar by a large diastema; it is small, one-rooted, and asymmetric with a
<figure><img src="image_5.png" /></figure>

<figure><img src="image_6.png" /><figcaption>**Fig. 3.** **A, B, F, G. Amphicynodon leptorynchus.** A: right hemimandible, c-m3, ITD355, in buccal (1), lingual (2) and occlusal (3) views; B: right hemimandible, c-m2, ITD361, in buccal (1), lingual (2) and occlusal (3) views; F: juvenile left maxilla, D1-D4, ITD1180, occlusal view; G: Right hemimandible, d3-d4-m1, ITD365, in buccal (1) and lingual (2) views. **C, D. Amphicynodon typicus.** C: right hemimandible, p3-m2, PC46 (holotype), in buccal (1), lingual (2) and occlusal (3) views; D: left hemimandible, p4-m1, ITD1220, in buccal (1), lingual (2) and occlusal (3) views. **E. Amphicynodon gracilis.** hemimandible, p2-m3, ITD1149, in buccal (1) and lingual (2) views. **H. Amphicynodon** sp., left maxilla, P4-M2, ROQ34, in buccal (1) and occlusal (2) views. Scale bars: 1 cm.</figcaption></figure>
L. de Bonis, A. Gardin and G. Escarguel

Geobios xxx (xxxx) xxx

<figure><img src="image_7.png" /><figcaption>**Fig. 4.** Postcranial elements of *Amphicynodon leptorhynchus* from Itardies. **A.** Scapula, ITD-VF-10425. **B.** Humerus, ITD-VF-11259. **C.** Ulna, ITD-VF-11259. **D.** Radius, ITD-VF-11065. **E.** Manus, ITD-VF-10360. **F.** Coxal bone, ITD-VF-10242. **G.** Femur, ITD-VF-10001. **H.** Tibia, ITD-VF-10594a. **I.** Fibula, ITD-VF-10594b. **J.** Calca-neus, ITD-VF-10148. **K.** Talus, ITD-VF-10863. **L.** Navicular, ITD-VF-11107. **M.** Cuboid, ITD-VF-10202. Scale bars: 1 cm.</figcaption></figure>

small distal shelf; a smooth sagittal ridge runs mesiodistally. P2 and P3 are two-rooted; both larger than P1 – especially the latter –, they are more symmetric and thicker, especially P3, where there is no pcc. P4 is robust, with a well-marked but low, mesially projected parastyle; the metastyle is short; protocone wide, a high ridge running from the protocone all along the paracone and the metastyle as a cingulum; a short ridge runs to the mesial face of the paracone and a high another one, directed distally and closing a shallow elongate basin, vanishes along the metastyle; a massive protocone contacts the middle of the lingual face of the paracone while two other ridges close two small basins, mesially and distally.

The paracone of M1 is larger than the metacone, both being almost conical with slight mesiodistal ridges; the large protocone and thick protocrista and metacrista enclose a small and shallow trigone basin; a metaconule may be observed in some unworn teeth. There is lingually a large, half-moon shaped cingulum. M2 is smaller than M1 and oval, the buccal portion being slightly narrowed; the protocrista runs to the paracone and the metacrista runs distally to a clear metaconule. There is a large lingual cingulum.

**Mandible.** The corpus mandibulae is relatively shallow and elongate, tapering mesially and corresponding to the shape of the muzzle (Fig. 3(A, B)). There are two mental foramina below p2 and p3, respectively. Distally there is an elongate angular process. There is a moderate masseter fossa and the ascending process is relatively high, narrow and has a rounded or an axe-shaped top.

**Lower dentition (Table 1).** The incisors are rarely present in the mandible. They are very small and compressed each other against the canine; their size increases from the tiny i1 to i3; they are pointed with a small basal lingual cuspid for i1 and i2 and two for i3. The canine is large and more curved than the upper one; the mesial surface is flattened with an anterior vertical groove, the distal one being more convex, both separated by a thin lingual ridge.

The size of the premolars increases along the row; especially between the one-rooted p1 and p2; the first three premolars are simple, slightly asymmetric with a small distal shelf. The p4 is less asymmetric and possesses a dacd followed by a shallow small basin. The premolars may be contacted together or separated by small diastemas.

The carnassial is low and robust; the metaconid contacts the paraconid until mid-height; the more or less deep talonid is surrounded by the cristids of hypoconid and entoconid; there is no paraconid in m2; the metaconid is as large or larger than the protoconid, both being preceded by a small and shallow basin closed by a mesial cristid; in the talonid, a basin is encircled by the cristids of the hypoconid and entoconid; there is a variation in the morphology of the small and rounded m3 in some specimens where we observe a clearly indicated protoconid and metaconid followed by a monocuspid talonid while in other ones there are only cristids.

**Deciduous dentition.** Several remains of milk dentition in the locality of Itardies are allocated to *A. leptorynchus* on the basis on the size (Table 1) and of the morphology of m1 which is sometimes present (Fig. 3(F, G)). The d2 is asymmetrical with a medial part shorter than the distal one and a steeply inclined mesial border while the distal part has a gently inclined distal border. It is thin, without cingulid but a small accd. The d3 is elongated, thin and trenchant; the main cuspid (protoconid) is bucco-lingually compressed except for a vertical lingual pillar; there are a large accd and a large lingually situated pacd and at the distal basis, a small shelf with a shallow basin closed by a low cristid. The d4 has a slightly open trigonid with a metaconid well separated from the protoconid and the paraconid; in the talonid, the entoconid is elongated and higher than the hypoconid and its buccal border is steeply inclined while the lingual border of the hypoconid is less inclined; both cristids are interrupted by notches and separated from a large hypoconulid.

Upper dentition remains with D3-D4 are assigned to *Amphicynodon* based on size and overall trend of the dentition. A specimen, ITD1180, is extremely rare, and may be unique among the early Oligocene carnivorous, allowing to observe the full cheek tooth raw and the top of the adult canine of a young cub (Fig. 3(F)). D1 and D2 are very small – smaller than P1 and P2 – and simple; the former is asymmetric, the distal half being longer than the mesial one, while D2 is more symmetric. D3 has a pointed paracone, convex buccally and almost flat lingually; there is a large
<caption>**Table 1**</caption>

Measurement table of lower and upper teeth of various species of *Amphicynodon* from different early Oligocene localities of the 'Phosphorites du Quercy' (southwestern France).

<table><thead><tr><th>Amphicynodon<br>leptorhynchus</th><th>Lower<br>teeth</th><th>cl</th><th>cw</th><th>p1L</th><th>p1w</th><th>p2L</th><th>p2w</th><th>p3L</th><th>p3w</th><th>p4L</th><th>p4w</th><th>m1L</th><th>m1trg</th><th>m1w</th><th>m2L</th><th>m2w</th><th>m3L</th><th>m3w</th></tr></thead><tbody><tr><td rowspan="10">Itardies</td><td>mean</td><td>4.28</td><td>2.78</td><td>2.82</td><td>1.66</td><td>4.37</td><td>2.06</td><td>4.97</td><td>2.27</td><td>6.01</td><td>2.88</td><td>8.37</td><td>3.88</td><td>3.99</td><td>4.02</td><td>3.42</td><td>2.51</td><td>2.37</td></tr><tr><td>min</td><td>3.6</td><td>2.2</td><td>2.2</td><td>1.3</td><td>3.7</td><td>1.6</td><td>4.2</td><td>1.9</td><td>5.1</td><td>2.5</td><td>7.0</td><td>3.2</td><td>3.3</td><td>3.3</td><td>2.8</td><td>2.0</td><td>1.9</td></tr><tr><td>max</td><td>5.1</td><td>3.4</td><td>3.5</td><td>2.2</td><td>4.9</td><td>2.4</td><td>5.7</td><td>2.9</td><td>7.0</td><td>3.6</td><td>8.6</td><td>4.5</td><td>4.5</td><td>4.7</td><td>4.1</td><td>3.1</td><td>2.9</td></tr><tr><td>n</td><td>31</td><td>32</td><td>45</td><td>46</td><td>65</td><td>65</td><td>70</td><td>70</td><td>79</td><td>79</td><td>100</td><td>100</td><td>100</td><td>91</td><td>91</td><td>67</td><td>67</td></tr><tr><td><strong>Upper<br>teeth</strong></td><td><strong>CL</strong></td><td><strong>Cw</strong></td><td><strong>P1L</strong></td><td><strong>P1w</strong></td><td><strong>P2L</strong></td><td><strong>P2w</strong></td><td><strong>P3L</strong></td><td><strong>P3w</strong></td><td><strong>P4L</strong></td><td><strong>P4w</strong></td><td><strong>M1L</strong></td><td><strong>M1w</strong></td><td><strong>M2L</strong></td><td><strong>M2w</strong></td><td></td><td></td><td></td></tr><tr><td>mean</td><td>4.60</td><td>3.50</td><td>2.70</td><td>1.60</td><td>4.38</td><td>2.10</td><td>5.30</td><td>2.70</td><td>7.67</td><td>5.71</td><td>5.67</td><td>7.92</td><td>3.77</td><td>5.78</td><td></td><td></td><td></td></tr><tr><td>min</td><td>3.8</td><td>2.4</td><td>2.1</td><td>1.2</td><td>3.5</td><td>1.7</td><td>4.4</td><td>2.1</td><td>6.6</td><td>5.0</td><td>4.9</td><td>6.7</td><td>3.3</td><td>4.7</td><td></td><td></td><td></td></tr><tr><td>max</td><td>5.4</td><td>4.6</td><td>3.2</td><td>2.0</td><td>5.4</td><td>2.5</td><td>6.4</td><td>3.1</td><td>8.5</td><td>6.7</td><td>6.6</td><td>8.9</td><td>4.4</td><td>6.9</td><td></td><td></td><td></td></tr><tr><td>n</td><td>25</td><td>25</td><td>30</td><td>29</td><td>41</td><td>42</td><td>47</td><td>47</td><td>46</td><td>46</td><td>47</td><td>47</td><td>43</td><td>43</td><td></td><td></td><td></td></tr><tr><td><strong>Lower<br>milk<br>teeth</strong></td><td><strong>Ld2</strong></td><td><strong>wd2</strong></td><td><strong>Ld3</strong></td><td><strong>wd3</strong></td><td><strong>Ld4</strong></td><td><strong>wd4</strong></td><td><strong>Lm1</strong></td><td><strong>wm1</strong></td><td></td><td></td><td></td><td></td><td></td><td></td><td></td><td></td><td></td><td></td></tr><tr><td>mean</td><td>3.60</td><td>1.40</td><td>4.80</td><td>2.00</td><td>5.70</td><td>2.60</td><td>7.90</td><td>3.90</td><td></td><td></td><td></td><td></td><td></td><td></td><td></td><td></td><td></td><td></td></tr><tr><td>min</td><td>3.5</td><td></td><td>4.4</td><td>1.6</td><td>5</td><td>2.2</td><td>7.6</td><td>4</td><td></td><td></td><td></td><td></td><td></td><td></td><td></td><td></td><td></td><td></td></tr><tr><td>max</td><td>3.7</td><td></td><td>5.3</td><td>2.4</td><td>6.2</td><td>2.9</td><td>8.3</td><td>3.7</td><td></td><td></td><td></td><td></td><td></td><td></td><td></td><td></td><td></td><td></td></tr><tr><td>n</td><td>2</td><td>1</td><td>7</td><td>7</td><td>13</td><td>13</td><td>2</td><td>2</td><td></td><td></td><td></td><td></td><td></td><td></td><td></td><td></td><td></td><td></td></tr><tr><td><strong>Upper<br>milk<br>teeth</strong></td><td><strong>D1L</strong></td><td><strong>D1w</strong></td><td><strong>D2L</strong></td><td><strong>D2w</strong></td><td><strong>D3L</strong></td><td><strong>D3w</strong></td><td><strong>D4L</strong></td><td><strong>D4w</strong></td><td></td><td></td><td></td><td></td><td></td><td></td><td></td><td></td><td></td><td></td></tr><tr><td>mean</td><td>3.29</td><td>1.12</td><td>3.91</td><td>2.24</td><td>5.22</td><td>3.8</td><td>3.96</td><td>4.49</td><td></td><td></td><td></td><td></td><td></td><td></td><td></td><td></td><td></td><td></td></tr><tr><td>min</td><td></td><td></td><td></td><td></td><td>4.97</td><td>3.49</td><td>3.65</td><td>3.98</td><td></td><td></td><td></td><td></td><td></td><td></td><td></td><td></td><td></td><td></td></tr><tr><td>max</td><td></td><td></td><td></td><td></td><td>5.61</td><td>3.98</td><td>4.55</td><td>5.28</td><td></td><td></td><td></td><td></td><td></td><td></td><td></td><td></td><td></td><td></td></tr><tr><td>n</td><td>1</td><td>1</td><td>1</td><td>1</td><td>3</td><td>3</td><td>3</td><td>3</td><td></td><td></td><td></td><td></td><td></td><td></td><td></td><td></td><td></td><td></td></tr><tr><td><strong>Amphicynodon<br>typicus</strong></td><td>Lower<br>teeth</td><td><strong>p4L</strong></td><td><strong>p4w</strong></td><td><strong>m1L</strong></td><td><strong>m1trg</strong></td><td><strong>m1w</strong></td><td><strong>m2L</strong></td><td><strong>m2w</strong></td><td></td><td></td><td></td><td></td><td></td><td></td><td></td><td></td><td></td><td></td><td></td></tr><tr><td rowspan="4">Pech Crabit and<br>Itardies</td><td>mean</td><td>6.00</td><td>3.10</td><td>8.66</td><td>6.10</td><td>4.34</td><td>4.51</td><td>2.80</td><td></td><td></td><td></td><td></td><td></td><td></td><td></td><td></td><td></td><td></td><td></td></tr><tr><td>min</td><td>5.41</td><td>2.7</td><td>8.44</td><td>6.00</td><td>4.15</td><td></td><td></td><td></td><td></td><td></td><td></td><td></td><td></td><td></td><td></td><td></td><td></td><td></td></tr><tr><td>max</td><td>6.67</td><td>3.52</td><td>8.59</td><td>6.21</td><td>4.54</td><td></td><td></td><td></td><td></td><td></td><td></td><td></td><td></td><td></td><td></td><td></td><td></td><td></td></tr><tr><td>n</td><td>2</td><td>2</td><td>2</td><td>2</td><td>2</td><td>1</td><td>1</td><td></td><td></td><td></td><td></td><td></td><td></td><td></td><td></td><td></td><td></td><td></td></tr><tr><td><strong>Amphicynodon<br>gracilis</strong></td><td>Lower<br>teeth</td><td><strong>p1L</strong></td><td><strong>p1w</strong></td><td><strong>p2L</strong></td><td><strong>p2w</strong></td><td><strong>p3L</strong></td><td><strong>p3w</strong></td><td><strong>p4L</strong></td><td><strong>p4w</strong></td><td><strong>m1L</strong></td><td><strong>m1trg</strong></td><td><strong>m1w</strong></td><td><strong>m2L</strong></td><td><strong>m2w</strong></td><td><strong>m3L</strong></td><td><strong>m3w</strong></td><td></td><td></td></tr><tr><td rowspan="2">Itardies</td><td>mean</td><td>3.52</td><td>3.24</td><td>3.97</td><td>2.30</td><td>4.30</td><td>2.45</td><td>5.20</td><td>2.60</td><td>7.30</td><td>4.60</td><td>3.60</td><td>3.60</td><td>2.90</td><td>1.60</td><td>1.80</td><td></td><td></td></tr><tr><td>n</td><td>1</td><td>1</td><td>2</td><td>2</td><td>3</td><td>3</td><td>3</td><td>3</td><td>3</td><td>3</td><td>3</td><td>3</td><td>3</td><td>2</td><td>2</td><td>2</td><td></td><td></td></tr><tr><td><strong>Amphicynodon<br>sp.</strong></td><td>Upper<br>teeth</td><td><strong>P4L</strong></td><td><strong>P4w</strong></td><td><strong>M1L</strong></td><td><strong>M1w1</strong></td><td><strong>M1w2</strong></td><td></td><td></td><td></td><td></td><td></td><td></td><td></td><td></td><td></td><td></td><td></td><td></td><td></td></tr><tr><td>Roqueprune 2</td><td>mean</td><td>8.62</td><td>7.24</td><td>5.40</td><td>8.17</td><td>7.37</td><td></td><td></td><td></td><td></td><td></td><td></td><td></td><td></td><td></td><td></td><td></td><td></td><td></td></tr><tr><td></td><td>n</td><td>1</td><td>1</td><td>1</td><td>1</td><td>1</td><td></td><td></td><td></td><td></td><td></td><td></td><td></td><td></td><td></td><td></td><td></td><td></td><td></td></tr></tbody></table>

pointed parastyle, slightly basined at the base of the paracone, while the metastyle is small; there is a protocone slightly basined and directed lingually from the middle part of the crown; there is a lingual cingulum forming the top of the protocone and the parastyle. D4 is triangular and its mesial width is shorter than the distal one; there is a buccal shelf with a cingulum giving rise to a parastyle and a metastyle, the latter being larger than the former and linked to the metacone by a small crest; the pyramidal paracone is taller than the conic metacone, both linked to a low protocone by two thick paracrista and metacrista without any trace of conule. Between the thick crests of the trigone, the central basin is small but deep.

**Appendicular skeleton.** The anatomy of the appendicular skeleton of *A. leptorhynchus* from Itardies (Fig. 4) has been described in detail in comparison to several extant carnivorans and analysed to infer its locomotor adaptations and its paleoecology (Gardin et al., 2021, 2022).

The mean body mass of *A. leptorhynchus* is evaluated at 1.9 kg and has been inferred from the measurements taken on the postcranial elements; bringing this animal close to the stature of the extant stone martens or genets (Gardin et al., 2021, 2022). All the locomotor adaptations of *A. leptorhynchus* converge on proficient climbing skills (Gardin et al. 2021, 2022) of which here is a summary. Contrary to terrestrial, especially to cursorial carnivorous, its limbs movements were not constrained in a parasagittal

plane by the shape of its articulations but had a great range of mobility. Indeed, *A. leptorhynchus* was able to move its shoulder, elbow, wrist, hip, and ankle articulations in a 3D-space. The attachment areas for retractors of the humerus, the flexors and adductors of the forelimb, and the adductors and hamstring muscles of the hindlimb seem to indicate that these muscles played a very important role in the mode of locomotion of *A. leptorhynchus*. Indeed, they allowed this small carnivoran to grip a vertical substrate very firmly, with the palm and foot facing toward the support; and carry its body in its posture against gravity. Thus, it was able to control its movements, to propel its body upwards or to retain it when descending headfirst, as do the extant arboreal carnivorans such as the lesser panda or the kinkajou. The morphology of the olecranon suggests that *A. leptorhynchus* adopted a crouched posture during the stance phase, which allowed him to lower his centre of gravity near the support, thus increasing its stability when climbing. The anatomy of the bones of its forearm suggests good pronation-supination ability. The digits of *A. leptorhynchus* were long, widely spaced and very mobile (especially the thumb), able to close on the palm of the hand. Moreover, the morphology of the middle and distal phalanges of *A. leptorhynchus* demonstrate the ability to partially retract its claws and control their movement, both for the hand and the foot. Like some extant arboreal carnivorans, *A. leptorhynchus*' ankle articulation allowed it to rotate its foot, with the plantar aspect lying in a medial or dorsal position,
a movement called hindfoot reversal. Using a firm grasp on a branch with its well-developed digital flexor, *A. leptorhynchus* was also probably able to perform hindlimb suspension. This skill was useful for this climber, as it allowed it to hold its body against gravity, by the feet, when it descended a vertical support. Thanks to the mobility of its limbs, *A. leptorhynchus* could use its hands and claws not only to firmly grip the branches around it during locomotion, but also to catch food or to pin its prey to the ground to kill it. Several other postcranial features shared with half-bounding carnivorans suggest strong hindlimb extensors favouring powerful extension movement allowing to perform jumps and rapid accelerations.

Based on those results, Gardin et al. (2021, 2022) bring the probable lifestyle of *A. leptorhynchus* closer to that of current small skilful arboreal carnivorans such as the lesser panda, the African palm civet, the kinkajou, the stone marten or the genet, which spend most of the time in trees.

**Remarks:** The locality of Itardies was a small fissure with few preserved fossil-bearing sediments but it allowed the recording of a huge number of *A. leptorhynchus* remains including dozens of skulls, hundreds of mandibles and limb bones (Cirot, 1992; Cirot and Bonis, 1992; Gardin et al., 2021, 2022). It was possible to get a good idea of the size variation into a population of *Amphicynodon* (Table 1).

*Amphicynodon typicus* (Schlosser, 1888)
Fig. 3(C, D)

**Lectotype by designation** (Cirot and Bonis, 1992: p. 121): left hemi-mandible with p4-m2 (PC46).

**Type locality:** Pech Crabit, MP23, early Oligocene; Quercy, Occitanie, France.

**Material:** Pech-Crabit (UP-PC) 46 md p4-m2. Itardies (UP-ITD) 1220 md p4-m1.

**Age:** MP 23 (early Oligocene).

**Measurements:** see Table 1.

**Diagnosis:** Medium-sized *Amphicynodon*; p4 symmetrical, pointed, taller than the paraconid of m1, mesial edge gently sloping and slightly concave, developed pacd; grouped trigonid of m1, not elongate paraconid, metaconid taller than paraconid; basined and close distally talonid with well-marked hypoconid and low entoconid cristid; complete trigonid of m2 with metaconid larger than the protoconid; small m3.

**Remarks:** We add another specimen to *A. typicus*: a hemimandible from Itardies (ITD1220; Fig. 3(C, D)) displaying a high trigonid with a pointed metaconid of m1, pointed p4 higher than the paraconid of m1, and fitting the characters of this species.

The type-locality of the lectotype is Pech Crabit (Quercy, Occitanie, France), a large cavity with a fossil-bearing cliff of clay. The species name *typicus* comes from Schlosser (1887) for a hemimandible which, following this author, was similar to *Cynodictis dubius* Filhol, 1882, calling it *Plesiocyon typicus*. Nevertheless, Filhol's species belongs to *Pachycynodon* and Schlosser's specimen, borrowed by Teilhard de Chardin (1915: pl. 5, fig. 1), was lost; thus, a lectotype was created later (Cirot and Bonis, 1992).

*Amphicynodon gracilis* (Filhol, 1874)
Fig. 3(E)

**Holotype:** Well preserved-left hemimandible (MNHN-QU 9192) with p2-m1 (Filhol, 1876: fig. 337).

**Origin:** Phosphorites of Quercy, site unknown.

**Material:** Itardies (UP-ITD) 1149 md p2-m3, 1172 md p4-m1,? 230 skull.

**Age:** Unknown, probably Oligocene.

**Measurements:** see Table 1.

**Diagnosis:** Small amphicynodont with simple lower premolars except for p4 which may bear a small pacd and a small pcsd; a more or less marked cingulid runs on the buccal base of the crown of the cheek teeth; the main characters of m1, which differs from those of other amphicynodonts, are the thinner m1 (index 100 × width/length of m1 < 50; generally higher in other species); less oblique paraconid blade and faint mesial reduction of the metaconid which slightly opens the trigonid basin and is a little backward shifted; in the talonid, hypoconid high and slightly trenchant and lower entoconid, both join distally to close the talonid basin; no paraconid in m2, metaconid as large as the protoconid both being clearly separated with, in front, a small transversal cristid and narrow and shallow basin connected to the main talonid basin.

**Description:** Few specimens from Itardies display these characteristics and are assigned to *A. gracilis* (Fig. 3(E); Table 1), such as the hemimandible ITD1149 with p2-m3. A m1 of another hemimandible (ITD1172) has some characters of this species; it is slightly smaller and more worn than the type species; there is no pacd on p4, m1 is relatively narrow and fissured but the metaconid seems a little reduced and slightly backwardly shifted, while in the relatively narrow talonid, the height difference between hypoconid and entoconid is low. The presence of m2 allows adding its morphology in the diagnosis.

Another specimen, unfortunately much damaged (ITD230), is put into the same group. It is a skull with P3-M1, associated with a mandible with left well-preserved p2-p3 and p4-m2 broken off, and right p2-p3, p4 the top of which is broken off, damaged m1-m2, and m3. This specimen gives the opportunity to describe some upper teeth of this species. P3 is simple, slightly asymmetrical, without any accessory cusp; its buccal face is regularly convex while, in the lingual face, a vertical pillar much convex is preceded and followed by a flat surface, the crown being encircled by a cingulum; the P4 paracone is compressed bucco-lingually, there is a small cingular parastyle, a relatively long metastyle for an amphicynodont, and a mesio-lingually projected high and pointed protocone. M1, much weathered, is a slightly scalene triangle; the protocone is mesially situated and does not surpass the level of the protocone of P4. This morphology is close to that of *A. leptorhynchus* but M1 is a little shorter and enlarged. The size of *A. gracilis* is compatible with that of *A. leptorhynchus*; nevertheless, in a comparison all known specimens of *A. gracilis* m1 would be placed in the left part of a histogram concerning *A. leptorhynchus*.

**Remarks:** This species was called *Cynodictis gracilis* by Filhol (1874: p. 18) for a hemimandible coming from the phosphorites of Quercy without any description or figure. Then, Filhol (1876) described the specimen as *Cynodon gracilis* and gave a figure (Filhol, 1877: fig. 337, although there is a mistake in the caption). Filhol noted the differences between this hemimandible and that of *Amphicynodon velaunum* (Aymard, 1846) to conclude that it was a new species. The type specimen, which was the only specimen until now, lacks p1, m2 and m3, thus, we try to complete the diagnosis with new material that we assign here to this species.

*Amphicynodon* sp.
Fig. 3(H)

**Material:** Roqueprune 2 (UP-ROQ) 34, P4-M1.mx.

**Measurements:** see Table 1.

**Description:** We compare a piece of maxilla with P4-M1 coming from Roqueprune 2 (ROQ34) to a sample of *Amphicynodon leptorhynchus* from Itardies (Fig. 3(H); Table 1). The P4 is slightly larger than the largest specimens of Itardies. The cingulum is slightly stronger than in *A. leptorhynchus*; the paracone bears a vertical mesial notch; the protocone is elongated and basined but without cusp on its lingual extremity and with a ridge running into
the basin. M1 is more elongate bucco-lingually; the index w/
L = 0.86 while it can reach 1.34 to 1.5 in *A. leptorhynchus*. There
are parastyle and metastyle with two tiny cusps between them.
There is no trace of metaconule which is present in *A. lep-
torhynchus*. All these characters may distinguish the latter from
ROQ34, and the maxilla belongs to another species. Nonetheless,
insofar the species of *Amphicynodon* are based on mandibles, the
material is not enough to erect a new taxon.

Genus *Pachycynodon* Schlosser, 1887-1889.

**Type species:** *Cynodictis crassirostris* Filhol, 1876.

**Diagnosis** (partially from Bonis et al., 2019): large size range (from ca. 7 to 14 mm for the length of m1), thick skull bones in large forms, more or less elongated snout, dentition more robust than that of *Amphicynodon*, often deep dentary under the molars; p4 as tall or taller than the protoconid of m1; the index 100 × p4 length/m1 length is higher than in *Amphicynodon*; the relatively short m1 has massive cuspirids of the trigonid; the protoconid of m1 is less tall relatively to the height of paraconid and metaconid than in *Amphicynodon*; the postmetaconid cristid is sloping very obliquely toward the pre-entocristid; the paraconid is present or not in m2. The robust P4 has a short metastyle and large shelf-like mesiolingually directed protocone. There is a trend in M1 toward an almost quadratic outline, without paraconule but a more or less developed distolingual metaconule with a large cingulum surrounding the protocone, and the M2 is oval and reduced.

**Remarks:** Schlosser (1887-1889) considered two isolated teeth, P4 and M1, coming from the phosphorites of Quercy as types of a species named *Pachycynodon crassirostris* (op. cit.: pl. IX, figs. 2, 4, 7, 9). The locality is unknown, as is unknown the dating of the specimens, which is the case for all the fossils collected for the old collections of the phosphorites of Quercy. Later, Schlosser considered that these fossils and a new hemimandible from the Quercy (Schlosser, 1899: pl. 13, fig. 1) belong to *Cynodictis crassirostris* Filhol, 1876. However, Filhol's (1976: fig. 67-72) specimens belongs to *Amphicynodon*. The P4 figured by Schlosser was considered as the lectotype of the species (Peigné et al., 2014). The hemimandible figured by Schlosser is really a *Pachycynodon* and could correspond to *P. crassirostris*. Other mandibles were published by Teilhard de Chardin as belonging to *Pachycynodon crassirostris*. They were coming from the old collections of the Quercy, and thus without precise dating. A quite large hemimandible from the museum of Montauban (Teilhard de Chardin, 1915: pl. IV, fig. 11) is figured as *P. crassirostris* and as written by Teilhard de Chardin, keeps in touch with the group of *P. boriei* (Filhol, 1877) which contains the larger species of *Pachycynodon*. On a cast of the Montauban's specimen, we note a robust and high dentary, a large and very high p4, a quite large m1 (L = 9.5 mm) and a quite long m2 (L = 6.2 mm). A small cingulid encircles the mesial face of the p4 which has a well-developed posterior accessory cuspid followed by a small basined talonid. The m1 is characterized by a paraconid and a metaconid joining in most of their height and a deeply basined talonid; m2 displays a narrow mesial fovea with a mesial cristid on which a small but clear paraconid is present; the protoconid and metaconid are in contact and the talonid basin is small. We do not have any specimen like that in the material studied here.

*Pachycynodon filholi* Schlosser, 1887-1889

Fig. 5(A–G, K)

**Holotype:** Hemimandible with p3-m1 (Schlosser, 1887-99: pl. IX, figs. 1, 3, 5).

**Origin:** Phosphorites of Quercy, locality unknown.

**Material:** Roqueprune 2 (UP-ROQ) md p2-m1, 8 m1, 48 p2, p4-m2, 18 m1, 22 m1. Itardies (UP-ITD) 21 md m1-m2, 24 md m1-m3, 76 md p4-, m3, 236 md p2-m2, 237 md c, p2-m2, 379 frg. skull i3-

M1, i3-p2, 395 skull, 501 p2-m2, 495 c-p4, 497 p3-m3, 501 p2-m2, 568 skull, 568 b frg. skull, 570 md p2-m2, 616 md p4-m3, 622 md m1-m2, 782 md d3-d4, 848 mx M1-M2, 1023 md p2-m1, 1070 max P3 cas P4-M1, 1128 md m1-m3, 1144 md c-m3, 1271 md without m3, 1391 max P3-M2, 1593 md p2-m3, 1594 md c-m2, 1595 md p4-m2, ? 1596 P3-M1. Gardiol 3 (UM-GAO) 437 m1, 438 m1, 439 m1, 426 m2, 427 m2, 428 m2, 429 m2, 454 M1, 455 M1. Mounayne (UP-MOU) 2 md m1-m2, 3 m1, 14 m1, 17 m1, 21 M1 d4, 23 m1, 24 m1, 28 P4.

**Age:** Probably early Oligocene.

**Measurements:** see Table 2.

**Emended Diagnosis:** simple p3; p4 as tall as the protoconid of m1 or slightly higher; m1 with large metaconid and long talonid in which the basin is shallow and quite symmetric; m2 with small mesially situated proconid and metaconid, without paraconid and anterior basin, large shallow posterior basin; crown slightly tapering distally.

**Description:** The species was named by Schlosser (1887-1889: pl. IX, figs. 1, 3, 5) for the holotype hemimandible with p3-m1 and for an edentulous piece of maxilla (op. cit.: pl. IX, fig. 19) on material coming from the phosphorites of Quercy. The author's diagnosis was succinct: the new species is very similar to *P. crassirostris* except for a very small diastema between p1 and p2 and its size is smaller, p3 has a basal cingulid, p4 has also a posterior accessory cuspid. The main cuspid of the m1 (Length = 8 mm) is slightly less high than p4, a normal character in the genus. The talonid would be even longer than in *P. crassirostris* itself; it forms almost half of the tooth and represents a wide but very shallow basin; the hypocristid wall is much stronger than the entocristid one, but slopes very gently towards the inside. On a cast of the holotype, the mesial side of p3 is steeply inclined while the distal edge is gently inclined backward and encircles a small talonid to give a small shallow basin; p4 is larger than p3 and is more backwardly elongated, so, the buccal and lingual cingulid encircles distally a larger basin than in p3; there is a posterior accessory cuspid situated buccodistally at mid-height of the distal edge. On m1, the paraconid is lower and smaller than the metaconid, and a cristid runs from its top to the basis; the protoconid is as high as in p4; the talonid is quite long and the cristids fuse distally and are horse-shoe like around a wide and flat basin; a short buccal cingulid is at the basis of the protoconid blade.

The P4, which is not in the figured specimens, would be characterized by the narrowness of its protocone, just like that of *P. crassirostris*; like this, it is also very massively built; there is a well-developed basal cingulum. Based on the alveoli, the M1 must have been big. The P3 is very simple and only would have a weak basal bump.

**Lower dentition.** UP-ROQ48 is a well-preserved left hemimandible slightly larger than the holotype, with p2, p4-m2 preserved and p1, p3 and m3 alveoli. The lower border of the corpus is straight and there is a mental foramen under the posterior root of p3. The p2, whose top is broken off, is asymmetric; there is a minute anterior accessory cuspid, a cingulid encircling the crown and a slightly basined small posterior cingulid shelf; p4 is as tall as the protoconid of m1; it has a small anterior cingulid cuspid, a cingulid around the crown, a large apac and a slightly basined pcsd; m1 is low with robust trigonid cuspirids, particularly a large metaconid as tall as the protoconid. The large talonid displays a large elongate hypoconid followed by an uninterrupted high cristid with small cusculids; this cristid closes the deep talonid basin with a small entoconid separated from the metaconid by a notch; the cristid oblica is straight and parallel to the longitudinal plane of the tooth row to the contact with the protoconid; there is a short cingulid along the paraconid. There is no trace of paraconid on the bean-shaped, relatively small m2; the large and pyramidal pro-
<figure><img src="image_8.png" /><figcaption>**Fig. 5.** **A-G, K. Pachycynodon filholi.** A: right hemimandible with c-m3, ITD1260, in buccal (1) and lingual (2) views; B: left hemimandible with p3-m2 without m3, ITD1271, in buccal (1), lingual (2) and occlusal (3) views; C: fragment of left mandible with m1, ITD1803, in buccal (1) and lingual (2) views; D: P4, MOU28, in buccal (1) and occlusal (2) views; E: skull, ITD568, in dorsal (1), ventral (2), and maxilla with p4-m2 (3) views; F: left hemimandible with d3-d4, ITD782, in buccal (1) and lingual (2) views; G: juvenile maxilla with right D3-D4 and left D3-D4, ITD1062, occlusal view; K: UP-ROQ48, left mandible with p2, p4-m2, lingual view. **H1-H3. Pachycynodon boriei**, right hemimandible with p4-m2, GAO351, in buccal (1), lingual (2) and occlusal (3) views. **H4-H7. Pachycynodon cf. boriei.** H4: M1, GAO373, occlusal view; H5: M1, GAO374, occlusal view; H6: M1, GAO375, occlusal view; H7: M2, GAO385, occlusal view. **J.**?Pachycynodon amphictina, left hemimandible with p2-m3, ITD649, buccal view. Scale bars: 1 cm.</figcaption></figure>
<table><caption>**Table 2**
Measurement table of lower and upper teeth of *Pachycynodon filholi* from Itardies (early Oligocene; 'Phosphorites du Quercy', southwestern France).</caption><thead><tr><th>Pachycynodon filholi</th><th>Lower teeth</th><th>cL</th><th>cw</th><th>p1L</th><th>p1w</th><th>p2L</th><th>p2w</th><th>p3L</th><th>p3w</th><th>p4L</th><th>p4w</th><th>m1L</th><th>m1trg</th><th>m1w</th><th>m2L</th><th>m2w</th><th>m3L</th><th>m3w</th></tr></thead><tbody><tr><td rowspan="10">Itardies</td><td>mean</td><td>3.70</td><td>2.40</td><td>2.57</td><td>1.57</td><td>4.19</td><td>1.98</td><td>4.73</td><td>2.12</td><td>5.76</td><td>2.85</td><td>7.46</td><td>3.86</td><td>3.71</td><td>3.59</td><td>3.24</td><td>2.3</td><td>2.25</td></tr><tr><td>min</td><td>3.4</td><td>2.2</td><td>2.2</td><td>1.4</td><td>3.8</td><td>1.9</td><td>4.2</td><td>2.0</td><td>5.1</td><td>2.4</td><td>7.0</td><td>3.4</td><td>3.1</td><td>3.0</td><td>3.0</td><td>2.0</td><td>1.8</td></tr><tr><td>max</td><td>3.9</td><td>2.6</td><td>2.9</td><td>1.9</td><td>4.6</td><td>2.1</td><td>5.3</td><td>2.4</td><td>6.3</td><td>3.5</td><td>8.2</td><td>5.6</td><td>4.1</td><td>4.2</td><td>3.7</td><td>2.6</td><td>2.6</td></tr><tr><td>n</td><td>4</td><td>4</td><td>6</td><td>6</td><td>10</td><td>10</td><td>13</td><td>13</td><td>15</td><td>15</td><td>20</td><td>19</td><td>19</td><td>19</td><td>19</td><td>16</td><td>16</td></tr><tr><td><strong>Upper teeth</strong></td><td><strong>P4L</strong></td><td><strong>P4w</strong></td><td><strong>M1L</strong></td><td><strong>M1w</strong></td><td><strong>M2L</strong></td><td><strong>M2w</strong></td><td></td><td></td><td></td><td></td><td></td><td></td><td></td><td></td><td></td><td></td><td></td><td></td></tr><tr><td>mean</td><td>6.83</td><td>5.73</td><td>4.94</td><td>7.19</td><td>3.74</td><td>4.35</td><td></td><td></td><td></td><td></td><td></td><td></td><td></td><td></td><td></td><td></td><td></td></tr><tr><td>n</td><td>1</td><td>1</td><td>1</td><td>1</td><td>1</td><td>1</td><td></td><td></td><td></td><td></td><td></td><td></td><td></td><td></td><td></td><td></td><td></td></tr><tr><td><strong>Lower milk teeth</strong></td><td><strong>d2L</strong></td><td><strong>d2w</strong></td><td><strong>d3L</strong></td><td><strong>d3w</strong></td><td><strong>d4L</strong></td><td><strong>d4w</strong></td><td><strong>m1L</strong></td><td><strong>m1w</strong></td><td></td><td></td><td></td><td></td><td></td><td></td><td></td><td></td><td></td></tr><tr><td>mean</td><td>3.60</td><td>1.40</td><td>4.80</td><td>2.00</td><td>5.70</td><td>2.60</td><td>7.90</td><td>3.90</td><td></td><td></td><td></td><td></td><td></td><td></td><td></td><td></td><td></td></tr><tr><td>min</td><td>3.5</td><td>--</td><td>4.4</td><td>1.6</td><td>5.0</td><td>2.2</td><td>7.6</td><td>4.0</td><td></td><td></td><td></td><td></td><td></td><td></td><td></td><td></td><td></td></tr><tr><td>max</td><td>3.7</td><td>--</td><td>5.3</td><td>2.4</td><td>6.2</td><td>2.9</td><td>8.3</td><td>3.7</td><td></td><td></td><td></td><td></td><td></td><td></td><td></td><td></td><td></td></tr><tr><td>n</td><td>2</td><td>1</td><td>7</td><td>7</td><td>13</td><td>13</td><td>2</td><td>2</td><td></td><td></td><td></td><td></td><td></td><td></td><td></td><td></td><td></td></tr><tr><td><strong>Upper milk teeth</strong></td><td><strong>D3L</strong></td><td><strong>D3w</strong></td><td><strong>D4L</strong></td><td><strong>D4w</strong></td><td></td><td></td><td></td><td></td><td></td><td></td><td></td><td></td><td></td><td></td><td></td><td></td><td></td></tr><tr><td>mean</td><td>4.61</td><td>3.29</td><td>4.26</td><td>3.24</td><td></td><td></td><td></td><td></td><td></td><td></td><td></td><td></td><td></td><td></td><td></td><td></td><td></td></tr><tr><td>min</td><td>4.51</td><td>3.16</td><td>4.07</td><td>3.41</td><td></td><td></td><td></td><td></td><td></td><td></td><td></td><td></td><td></td><td></td><td></td><td></td><td></td></tr><tr><td>max</td><td>4.70</td><td>3.41</td><td>4.45</td><td>3.41</td><td></td><td></td><td></td><td></td><td></td><td></td><td></td><td></td><td></td><td></td><td></td><td></td><td></td></tr><tr><td>n</td><td>2</td><td>2</td><td>2</td><td>2</td><td></td><td></td><td></td><td></td><td></td><td></td><td></td><td></td><td></td><td></td><td></td><td></td><td></td></tr></tbody></table>

protoconid is separated by a large notch from the hardly distinct hypoconid which is prolonged by a cristid closing the basin to the hardly distinct entoconid and joins the metaconid; protolophid and metalophid join at the mesial border, underlined by a bit of cingulid; the bottom of the talonid basin is partly constituted by the gentle slope of the hypoconid opposed to the more steeply slope of the entocristid; the tooth being unworn, we see some tiny cusps on the hypocristid; the m3 is small, rounded, with a flat bottom completely encircled by a cristid.

UP-ITD1260 is a well-preserved mandible slightly larger than the holotype. The right side of UP-ITD1260d bears c-m3 (Fig. 5 (A)). The dentary is convex below the m1-m2 limit and going up below the ascending ramus; the latter is quite low when compared to that of *Amphicynodon*; the angular apophysis is long and thin; the condyle is situated slightly higher than the level of tooth row. The canine is strong but not very high. The trigonid is like that of *P. filholi* with a short cingulid at the buccal basis of paraconid and protoconid; on the rectangular m2, there is no paraconid and no mesial basin in front of protoconid; metaconid, hypoconid and entoconid are marked. The tops of the premolar and molar cusps are truncated by an horizontal wear facet while we do not see the common vertical wear facet on the buccal side of protoconid and paraconid.

Teilhard de Chardin (1915) in his monograph on the small Quercy carnivorans noted that the m3 was lacking in some hemimandibles of *Pachycynodon*. The studied material was not dated and it was impossible to know if each specimen in which m3 was lacking belonged to a similar population or was an accident in a population in which m3 was present. Thus, Teilhard de Chardin considered generally these fossils as varieties or subspecies. The new research in the locality of Itardies allows studying a population of a dozen of hemimandibles of *Pachycynodon* in which two of them do not have m3. Except for the number of molars, these mandibles display the size and all the characteristics of *P. filholi* and we think that the specimens without m3 do not belong to another taxon but they are only individual variants (Fig. 5(B)).

There are several described species of *Pachycynodon* (Filhol, 1882; Schlosser, 1887-1889; Teilhard de Chardin, 1915; Peigné et al., 2014; Bonis et al., 2019). Because of the large difference in size we may leave out *P. boriei* (Filhol, 1873), *P. curvirostris* (Filhol, 1873), *P. gryei* (Filhol, 1876), *P. robustus* (Filhol, 1876), and *P. leymeriei* Filhol, 1876 to focus on the smaller species of which m1 length ranges between 7 and 9 mm (Bonis et al., 2019: p. 606).

*Pachycynodon tenuis* Teilhard de Chardin, 1915 differs by a lower p4, low for a *Pachycynodon*, with a thick lingual vertical pillar and longer relative to m1; by m1 with a larger metaconid and a greater height difference between protoconid and paraconid; and m2 with a mesial basin posteriorly limited by a cristid between the protoconid and metaconid.

*Pachycynodon dubius* (Filhol, 1882) differs by its larger p4 with a less steeply inclined mesial border, the lower trigonid of m1 and its less steeply inclined distal face.

*Pachycynodon amphictina* Teilhard de Chardin, 1915 differs by a reduced distal basin of p4, the presence of a paraconid in m2, and the m3 situated in the vertical border of the coronoid process.

*Pachycynodon filholi* Schlosser, 1888 was erected for a mandible from the Quercy (Schlosser, 1888: pl. IX, fig. 1). Teilhard de Chardin (1915: pl. 4, fig. 7), who knew the original specimen, figured another mandible housed in the Catholic University of Louvain-la-Neuve, Belgium; several labelled *P. filholi* hemimandibles are in the MNHN collection, Paris. These specimens display the same features described in UP-ROQ48 and UP-ITD1260. Several hemimandibles or isolated teeth are allocated to the same species (Fig. 5(A)). *P. filholi* is present in all the localities but it is, by the number of specimens, the dominant carnivoran in Mounayne. We note the horizontal wear facets noted by Teilhard de Chardin (1915: p. 39) on some specimens, and, for instance, the tooth ITD1803 from Itardies, displaying a wear pattern unusual for a carnivore (Fig. 5(C)).

Despite some faint variations, all mandibles display the same characters as those described above. The size cannot separate the specimens coming from the different localities, and no character may justify the presence of another species than *Pachycynodon filholi* in this group (Table 2).

**Upper dentition.** Upper tooth remains are rarer than those of the lower ones. We never found associated upper and lower dentitions. Nevertheless, we consider P4s like UP-MOU28 (Fig. 5(D)) which has a very long, flat, broad (almost the whole lingual side) and antero-lingually projecting protocone as belonging to *Pachycynodon*. They differ from that of *Amphicynodon*. The metastyle is relatively short and there is a small parastyle. A robust cingulum encircles the tooth. Similar P4 is also present in a badly preserved skull from Itardies (UP-ITD568) of which anterior and posterior part are broken off but upper P4-M2 are preserved. The zygomatic arches of the skull are thin (Fig. 5(E)). In dorsal view, the faint temporal lines are fused at the level of the mandible articulation; more posteriorly the bone is damaged and we cannot control a putative
sagittal crest. In ventral view, P4, although damaged, displays the same characters that UP-MOU18. In M1, the pyramidal paracone is slightly larger than the conical metacone; a thick cingulum follows the buccal base of the crown; we may observe a part of the trigone "V" of which the base is missing; as in *Amphicynodon*, a crescent-like cingulum extends lingually. M2 is oval and more reduced than that of *Amphicynodon*; small paracone and metacone are close to each other in the narrow buccal portion of the tooth which is bordered by a thick cingulum; the lingual portion is wider than the buccal one with a small protocone (Fig. 5(E3)). We add to *P. filholi* a fragment of maxilla with M1-M2 (UP-ITD848) whose molars have the same proportions as those of UP-ITD568.

Milk dentition. A small milk mandible with d3-d4 (UP-ITD782) cannot belong to *Palaeogale*, *Stenoplesictis* and *Stenogale* of which dentitions are more hypercarnivore (Fig. 5(F)). The d3 is tall, robust and pointed without accd, contrary to the d3 of *Amphicynodon lep-torhynchus*; the mesial face is steeply inclined; there is a large pacd buccally situated at the mid-height of the distal crest; a cingulid rounds the crown and, at the distal base, it closes a large shelf with a shallow basin; d4 displays a quite open trigonid and a wide talon-id of which hypoconid and entoconid have almost the same size, a large hypoconulid closing a deep talonid basin. The deciduous lower teeth of *Amphicynodon* are different: d3 is thinner, more trenchant with a well-developed accd, the trigonid of d4 is less open and, in its talonid, the hypoconid is more developed than the entoconid and the hypoconulid is smaller and distally pointed. Thus, UP-ITD782 is assigned to *Pachycynodon filholi*.

A maxilla fragmented in two parts (UP-ITD1062) corresponds to *P. filholi*. The size is smaller than that of *Amphicynodon lep-torhynchus*. D3 has a paracone with a convex buccal face and a well-defined parastyle. The protocone is more mesially situated than in *P. leptorynchus* and it is relatively longer. D4 has a less expended metastyle and the lingual half of the crown is narrow (Fig. 5(G)).

**Remark:** The horizontal wear facets in the dentition are known in extant and fossil hyenids as well as in the fossil canids of the subfamily Borophaginae. In both cases, this uncommon wear is linked to a bone-breaking adaptation. We may suppose the same for some *Pachycynodon* hunting for rodents and breaking their bones. The presence in the fossil remains of carnivoran teeth which have been digested could indicate cannibalism into the carnivoran community. Another explanation could be a vegetal diet with the consumption of roots, tubers or grains. A study of the micro-striations may help solving this point.

?*Pachycynodon amphictina* Teilhard de Chardin, 1915 = *Pachy-cynodon filholi* var. *amphictina* Teilhard de Chardin, 1915 (p. 38, pl. IV, fig. 8) = *Pachycynodon crassirostris* Schlosser, 1888 (p. 253) (Peigné et al., 2014: p. 33, fig. 20a)

Fig. 5(J)

**Material:** Itardies (UP-ITD) 649 md p1-m3.

**Remarks:** This species was created from what Teilhard de Chardin considered a "variant" (Bonis et al., 2019). One of the characteristics of this species is the place of m3, which laid out in the coronoid process. It seems to be the case of a small hemimandible of Itardies (UP-ITD649) with p1-m3. Unfortu-nately, the specimen bears well-preserved p1-p4 but the m1 and m2 are badly preserved and we cannot be sure of the place of m3 (Fig. 5(J)).

*Pachycynodon boriei* (Filhol, 1873) = *Cynodictis curvirostris* Filhol, 1873.

Fig. 5(H1-H3)

**Holotype by designation:** *Cynodictis boriei* Filhol, 1873, fragment of skull with P4-M1 (Filhol, 1876: pl. 14, fig. 34; pl. 15 fig. 33), and hemi-mandible with c, p2-m1 (Filhol, 1876: pl. 16, figs. 37, 40).

**Material:** *Gardiol 3* (UM-GAO) 351 md p4-m2, 352 md p4, 354 m1, 355 m1, 356 m1, 357 m1, 358 m1, 359 m1 cas, 373 M1, 374 M1, 375 M1, 376 M1, 385 M2.

**Measurements:** see Table 3.

**Diagnosis:** Largest species of *Pachycynodon*; simple P2; P4 with short metastyle, protocone as a plateau but not significantly pro-jected. M1 with symmetric buccal cusps, thick buccal cingulum. Low lower premolars except p4 which is higher than the proto-conid of m1 (character of the genus). No diastema between the premolars (shortly snouted).

**Description:** Until now, nobody firmly recorded the species in the new collections of the Quercy (Bonis et al., 2019) but the local-ity Gardiol 3 allows recording some remains considered here as *P. boriei*. A mandible fragment with p4-m2 (GAO351) shows the char-acters of this species (Fig. 5(H1-H3); Table 3). The p4 is tall (taller than the protoconid of m1) and pointed with a light mesial and buccal cingulid which is prolonged distally, closing a small basin. There is a developed accessory cuspid at mid-height of the distal cristid. The carnassial is low with a grouped trigonid in which para-conid and metaconid have almost the same size, the latter being not distally shifted; there is a slight wear facet on the buccal face of the protoconid-paraconid shearing blade but not on the top of the three cuspid; a wide talonid is circled by the thick hypoconid and entoconid joining distally, the latter has a lingual wear facet; both have a flat horizontal wear facet both are joining distally, clos-ing a flat talonid basin in which there is another round wear facet in which the dentine appears in the distal part of this basin. The index 100 × width/length of m1 is generally higher than 50. The quadrangular m2 is long and slightly tapering distally; in the mesi-olingual corner, there is a small conical paraconid displaying a small hollowed-out wear facet on the top; protoconid, metaconid, the latter higher than the former, close a small mesial basin, while thick hypoconid and entoconid close a deep talonid basin. A p4 with a partly broken-off p3 (UM-GAO352) in a piece of mandible is not well preserved but seems similar to that of UM-GAO351; there is no diastema between p4 and the broken p3. Two isolated m1 (UM-GAO358 and UM-GAO359) are similar to the UM-GAO351 m1 and are referred to *P. boriei*. Isolated lower m3s, small and rounded, would be compatible with this species but also with other taxa.

**Remarks:** Specimens from Gardiol 3 clearly match large species of the genus *Pachycynodon* (m1 length from 10.8 to 13.4 mm). *Filhol* (1873) cited, without figure, under the generic name *Cynod-ictis*, *Pachycynodon boriei* and *P. curvirostris*. The former was erected for a piece of skull with unworn P4-M2 coming from the phospho-rites of Quercy and later figured (Filhol, 1876: figs. 34-36) together; he wrote "two years ago, I described an upper maxilla of one of the larger carnivores ever found in the Quercy" (authors' translation) and an almost complete lower maxilla (op. cit.: figs. 37-40) "was linked to the skull, it is the same individual" (Authors' translation). These specimens constitute the holotype of *Pachycyn-odon boriei* (Filhol, 1873). In the same paper, Filhol figured (op. cit.: figs. 46-48) as the same species another piece of mandible, slightly smaller and he described, always under the generic name *Cynodic- tis*, *P. robustus* for a piece of hemi-mandible (op. cit.: figs. 30-32), *P. gryei* for a skull (op. cit.: figs. 58-60), *P. curvirostris* and *P. leymeriei* for two hemi-mandibles (op. cit.: figs. 52-54 and 55-57, respec-tively), and a piece of maxilla with P2-M2 (op. cit.: figs. 35-36).

So, following Filhol, the mandible with c, p2-m1, of which the lower border is broken off between p3 and the alveolus of m3 (op. cit.: fig. 40), is part of the type together with the maxilla.
<caption>**Table 3**</caption>

Measurement table of lower and upper teeth of *Pachycynodon boriei* from Gardiol 3 and *P. amphictina* from Itardies (early Oligocene; 'Phosphorites du Quercy', southwestern France).

<table><thead><tr><th>Pachycynodon boriei</th><th>Lower teeth</th><th>p3L</th><th>p3w</th><th>p4L</th><th>p4w</th><th>m1L</th><th>m1Ltr</th><th>m1wtr</th><th>m1wtl</th><th>m2L</th><th>m2wtr</th><th>m2wtl</th></tr></thead><tbody><tr><td rowspan="12">Gardiol 3</td><td>mean</td><td>7.43</td><td>4.34</td><td>9.92</td><td>5.44</td><td>11.39</td><td>7.10</td><td>5.64</td><td>5.77</td><td>6.30</td><td>4.65</td><td>4.55</td></tr><tr><td>min</td><td>—</td><td>—</td><td>9.70</td><td>5.02</td><td>10.36</td><td>6.10</td><td>5.14</td><td>5.11</td><td>5.78</td><td>4.35</td><td>3.97</td></tr><tr><td>max</td><td>—</td><td>—</td><td>10.06</td><td>5.85</td><td>13.40</td><td>8.01</td><td>6.58</td><td>7.14</td><td>7.04</td><td>5.46</td><td>5.42</td></tr><tr><td>n</td><td>1</td><td>1</td><td>2</td><td>2</td><td>9</td><td>9</td><td>9</td><td>8</td><td>8</td><td>8</td><td>7</td></tr><tr><td><strong>Upper teeth</strong></td><td><strong>M1L</strong></td><td><strong>M1w</strong></td><td><strong>M2L</strong></td><td><strong>M2w</strong></td><td><strong>M3L</strong></td><td><strong>M3w</strong></td><td></td><td></td><td></td><td></td><td></td></tr><tr><td>mean</td><td>9.60</td><td>11.30</td><td>6.45</td><td>8.26</td><td>2.15</td><td>2.50</td><td></td><td></td><td></td><td></td><td></td></tr><tr><td>min</td><td>8.83</td><td>9.40</td><td>5.59</td><td>6.34</td><td>1.96</td><td>1.45</td><td></td><td></td><td></td><td></td><td></td></tr><tr><td>max</td><td>9.60</td><td>12.10</td><td>7.13</td><td>9.23</td><td>2.35</td><td>3.29</td><td></td><td></td><td></td><td></td><td></td></tr><tr><td>n</td><td>5</td><td>4</td><td>3</td><td>3</td><td>2</td><td>2</td><td></td><td></td><td></td><td></td><td></td></tr><tr><td><strong>Upper milk teeth</strong></td><td><strong>D3L</strong></td><td><strong>D3w</strong></td><td><strong>D4L</strong></td><td><strong>D4w</strong></td><td></td><td></td><td></td><td></td><td></td><td></td><td></td></tr><tr><td>mean</td><td>5.50</td><td>3.40</td><td>4.84</td><td>5.69</td><td></td><td></td><td></td><td></td><td></td><td></td><td></td></tr><tr><td>min</td><td>4.50</td><td>2.05</td><td>4.00</td><td>4.29</td><td></td><td></td><td></td><td></td><td></td><td></td><td></td></tr><tr><td>max</td><td>6.60</td><td>4.70</td><td>6.09</td><td>7.58</td><td></td><td></td><td></td><td></td><td></td><td></td><td></td></tr><tr><td>n</td><td>14</td><td>14</td><td>15</td><td>15</td><td></td><td></td><td></td><td></td><td></td><td></td><td></td></tr><tr><td rowspan="2">Pachycynodon amphictina?</td><td>Lower teeth</td><td>p1L</td><td>p1w</td><td>p2L</td><td>p2w</td><td>p3L</td><td>p3w</td><td>p4L</td><td>p4w</td><td>m1L</td><td>m1trg</td><td>m1w</td><td>m2L</td><td>m2w</td><td>m3L</td><td>m3w</td></tr><tr><td>mean</td><td>3.12</td><td>2.12</td><td>4.98</td><td>2.22</td><td>5.52</td><td>2.70</td><td>5.82</td><td>3.28</td><td>8.14</td><td>5.20</td><td>4.25</td><td>—</td><td>—</td><td>1.64</td><td>1.65</td></tr><tr><td>Itardies</td><td>n</td><td>1</td><td>1</td><td>1</td><td>1</td><td>1</td><td>1</td><td>1</td><td>1</td><td>1</td><td>1</td><td>1</td><td>—</td><td>—</td><td>1</td><td>1</td></tr></tbody></table>

The premolars join along the jaw and are not separated by significant diastemas. Part of the skull (MNHN QU9214) is preserved in the Paris collections but we did not find the hemimandible in these collections. It seems that there was an accident in which specimens were broken off between the deposit of the Filhol collection and the study of Teilhard de Chardin. The mandible figured by Teilhard de Chardin (1915: pl. IV, fig. 14) as *P. boriei* does not correspond to the figure of the type of Filhol by its scattered premolars.

In the holotype of *P. boriei* (MNHN-QU9214), the upper dentition is characterized by low and elongated P2 without accessory cusp, P4 is short relative to the molars with a large width, a relatively robust paracone, a short and thick metastyle. The protocone is a large mesio-lingual plateau with a small cusp in the centre but it is not significantly projecting lingually and forwardly. A cingulum surrounds the crown. The triangular M1 has symmetric buccal cusps – paracone and metacone – of similar size. The protocone is not clearly distinct from the two crests of the trigone encircling a large and deep trigone basin. A thick lingual cingulum is prolonged and vanishes along the crests of the trigone. There is no trace of clear metaconule. Filhol described a canine, a P1 and a P3 which are absent on the specimen.

The type mandible, on Filhol's drawing, bears a not very tall but robust canine and p2-m1. The premolars, without any diastema, are slightly elongated with a buccal cingulid, p2 and p3 are low and p4 is as tall as the protoconid of m1, a character of the genus, and displays a very small dacid and a small dcsd; m1 is short and robust. The figure is from the buccal face and we cannot estimate the morphology of the buccal trigonid and the features of the talonid. Nevertheless, we shall give some details below from the specimens we refer to *P. boriei*.

In the collections of the MNHN, another mandible has the same characteristics and a compatible size and allows describing other details. MNHN-QU9208 is the type mandible of "Cynodictis" curvirostris Filhol, 1873, figured in Filhol (1876: pl. 19, figs. 52–54). The figured mandible bears alveoli of incisors, canine, p1 and crowns of p3-m2. The characters used by Filhol to create the new species are founded on details of the relative size of the teeth and the deepness of the mandibular corpus. We think that none may be significant

enough to separate the specimen from *P. boriei*. The type specimen is short-snouted (no diastema between the premolars) and the trigonid of m1 is grouped, with no shifted metaconid, similar to that of *P. boriei*. MNHN-QU9216 is a hemimandible with c, alveolus of p1, p2-m1 (Filhol, 1876: figs. 46–48) and is considered as a slightly small *P. boriei*. The low and elongated premolars without diastema and the morphology of the carnassial confirm the determination. Another hemimandible (MNHN-QU9210) displays the same characters and belongs to the same taxon.

Filhol (1876: pl. 16, figs. 35, 36) figured as *P. boriei* a quite large maxilla with P1-M1 which corresponds by its scattered premolars to *P. robustus* and not to the mandible of *P. boriei* figured in the same plate.

Recently, *P. boriei* has been recorded (Rabi et al., 2018) "from the late Oligocene (Egerian) of Hungary", so a level younger than MP23. *Pachycynodon* belongs to a carnivoran fauna which contains also a m1 allocated to cf. *Amphictis* Pomel, 1853 or *Franconictis* Wolsan, 1993. If *Amphictis* is known in the late Oligocene of the Quercy (Bonis, 1976; Cirot and Bonis, 1993), *Franconictis* comes from the early Miocene of Wintershof-west in Germany. A third Hungarian taxon is *Potamotherium valletoni* Geoffroy Saint-Hilaire, 1833, also found in the early Miocene localities of Saint-Gérand le Puy, France (Viret, 1929). This questions the current biochronological correlations between the mammalian scales in Western and Central Europe.

*Pachycynodon robustus* (Filhol, 1876)

**Holotype:** hemimandible with p2-m2, MNHN-QU9217 (Filhol, 1876: pl.13, figs. 30–32).

**Origin:** Phosphorites of Quercy, unknown age.

**Diagnosis:** Large species of *Pachycynodon*; elongated lower premolars separated by diastemas; large distal accessory cuspid in p4.

**Remarks:** In the type mandible MNHN-QU9217, the premolars are separated by diastemas; p2 and p3 are elongated but the crowns are partially broken off; p4 is large, slightly shorter than m1 and displays a large distal accessory cuspid and a short distal shelf. This large distal accessory cuspid is absent on *P. boriei*. The trigonid of m1 is more open than in *P. boriei*, with a slightly distally
shifted metaconid which appears clearly in a buccal view of the mandible; the talonid has a robust hypoconid and a low entoconid. The m2 is quite long (index m2/m1 > 0.5). The trigonid is short with advanced protoconid and metaconid on the same line and almost the same size, limiting a small anterior basin closed by a low cristid without any paraconid. A large posterior talonid basin is closed by a low cristid. The deepness of the corpus is a little exaggerated by a transversal compression. Another mandible housed in the MNHN (QU9233) displays also a large size, scattered premolars and more open trigonid of m1, an elongate m2 and fits the type mandible.

Pachycynodon cf. *boriei*
Fig. 5(H4-H7)

**Description (upper dentition):** There is no direct link between the specimens of Gardiol 3 described above and any upper dentition. Nevertheless, upper molars from Gardiol 3 are compatible with a large form like *P. boriei*. UM-GAO373 and UM-GAO374 (Fig. 5(H4, H5)) are two similar unworn right and left M1s whose size is close to that of the teeth of *P. boriei* (Table 3). A small cingulum runs on the buccal face. The paracone is slightly taller and larger, by its buccomesial part, than the metacone, giving a small buccomesial protrusion; both are almost conical. The mesial and distal crests are in a mesio-distal line except the preparacrista directed toward the mesio-buccal corner; on the mesio-lingual side of the paracone. A line of tiny tubercles is going down to contact the preprotocone crista. The protocone is low, the preprotocrista merges with a thin cingulum at the level of the paracone; the short postprotocrista stops at a notch before a low but quite wide metaconule while the hypometacrista runs to the distal border and turns buccally to join the distal part of the metacone. A large cingulum extends lingually and frames the protocone.

**Remarks:** The pattern of these molars is similar to that of the *P. boriei* molar (Filhol, 1876: fig. 34) but, as far as it is possible to judge from Filhol's figure, similar also to the M1 of the maxilla (Filhol, 1876: pl. 16, figs. 35, 36) with scattered premolars that we refer to *P. robustus*. Two unworn other upper molars (Fig. 5 (H6)) belong also to a large *Pachycynodon* (UM-GAO375 and UM-GAO376). They are slightly smaller than UM-GAO373 and UM-GAO374, their lingual half being slightly narrower and the metaconule position being a little more buccal. These teeth could correspond to m1s that we may refer to *P. robustus*, or to smaller individuals of *P. boriei*. In doubt, we prefer to choose *P. boriei* for all the material. A M2 (Fig. 5(H7); Table 3) also belongs to the same group. Oval, it displays small, bunodont paracone and metacone close each other and, lingually, two small cusps probably corresponding to a protocone and a metaconule bordered by a large cingulum. Some very small rounded, one-rooted teeth are certainly m3 or M3 but they cannot be identified.

Ursidae Batsch, 1788
Hemicyoninae Frick, 1926
Cephalogalini Bonis, 2013

**Remarks:** The Cephalogalini are present in the new collections of the Quercy but their remains are scarce in the early Oligocene levels. Only a piece of mandible was described so far in the localities of MP22 (Bonis et al., 2019) and there are only two upper teeth and a p1 that we can refer to this tribe in Pech Crabit and Itardies.

Cephalogalini indet. 1
Fig. 6(A)

**Material:** Pech-Crabit (UP-PC) 34, M1.

**Description:** PC34 is a left M1 of a large-sized Cephalogalini (L = 11.6 mm, w1 = 15.6 mm, w2 = 13.9 mm). The tooth is encircled by

a cingulum thicker lingually and, to a lesser extent, buccally as a small parastyle and modest pad at the level of the metacone. The paracone is larger than the metacone, the former being pyramidal and the latter conical. There is a low crescent-shaped protocone with protocrista and postprotocrista finishing fused to the cingulum, mesial and distal, the latter bearing a very small metaconule. There are three roots, two cylindrical buccal, the distal one partly fused to the third which is lingual, larger and slightly compressed mesiodistally. The measurements of PC34 are compatible with those of several species of large Cephalogalini such as *Cyonarctos dessei*, *Phoberogale depereti*, *P. bonali*, *Cephalogale geoffroyi*, *C. ginesticus* and the morphology of M1 is almost the same. Thus, PC34 is considered as Cephalogalini indet. 1.

Cephalogalini indet. 2
Fig. 6(B)

**Material:** *Itardies* (UP-ITD) 1814, M1.

**Description:** Another M1 (UP-ITD3939) is smaller than the precedent. Its measurements (Length = 8.7 mm, width = 11.9 mm) correspond to those of a middle-sized Cephalogalini. Its overall morphology is quite the same as that of PC34 and we are unable to give it a species name – but it comes from a species different than of UP-PC34, and we name it Cephalogalini indet. 2.

Cephalogalini sp.
Fig. 6(C)

**Material:** *Itardies* (UP-ITD) 1814, p1.

**Description:** It is quite the same for an isolated p1 (UP-ITD1814) whose size is that of an Irish setter hound (Length = 4.6 mm; breadth = 2.3 mm; total height = 13.3 mm; crown height = 6 mm). The root is relatively short and robust. The crown is unworn; it is much asymmetric, the mesial part being far shorter than the distal one. The top is rounded; the mesial face is steeply inclined while the distal one is elongated and gently inclined with a distal plateau. A light bead encircles the base. The size and the morphology fit those of some Oligocene species and UP-ITD1814 is a Cephalogalini indet. The p1 (ITD1814) could belong to a quite large or middle-sized Cephalogalini whatever the species, maybe one described above, and we name it Cephalogalini sp.

?Amphicyonidae Haeckel, 1866

**Remarks:** The family is known in Eurasia from the Eocene to the late Miocene (Viranta, 1996). Its systematic position has been long time discussed. The Amphicyonidae were considered as Canidae, Ursidae, and now as an independent family. Several characters are shared with Ursidae and could be considered as entering the Arctoidea (Wang et al., 2023). For example, the ‘ursid loop’ into the basioccipital (Bonis, 1966: fig. 10), the osteology close to that of primitive Arctoidea, or some ursid characters (Hunt, 1998).

?Amphicyonidae indet.
Fig. 6(D)

**Material:** Roqueprune 2 (UP-ROQ) 33, P1.

**Description and remarks:** An isolated one-rooted first premolar of Carnivora (ROQ33) whose size is that of a P1 of a setter hound (Length = 4.6 mm; breadth = 2.3 mm; total height = 13.3 mm; crown height = 6 mm) comes from Roqueprune 2 (Fig. 6(D)). The crown is relatively tall for a first premolar and is pointed and slightly back curved; the buccal face is regularly convex while the lingual one is slightly concave in its distal portion. The root is long and slightly hooked at its extremity although a small piece of the top is broken off. The size would correspond to a nimravid, an ursid (Cephalogalini) or an amphicyonid. Into the nimravids,
<figure><img src="image_9.png" /><figcaption>**Fig. 6.** **A.** Cephalogalini indet. 1, left M1, PC34, occlusal view. **B.** Cephalogalini indet. 2, left M1, ITD3939, occlusal view. **C.** Cephalogalini sp., left p1, ITD1814, in buccal (1) and lingual (2) views. **D.** Amphicyonidae indet., left P1, ROQ33, in buccal (1) and lingual (2) views. **E, M, N.** *Huntictis minima* nov. gen., nov sp. E: left m1, GAO415 (holotype), in buccal (1), lingual (2) and occlusal (3) views; M: right d4, UM-GAO416a, buccal view; N: right m1, UM-GAO416a, occlusal view. **F.** Musteloidea gen. et sp. indet., left m1, GAO416, in buccal (1), lingual (2) and occlusal (3) views. **G, H, J.** *Mustelictis olivieri*. G: right hemimandible with p2-m2, ITD1063, in buccal (1) and lingual (2) views; H: right hemimandible with p2-m1, ITD1818, in buccal (1), lingual (2) and occlusal stereo (3) views; J: P4, ITD1809, in buccal (1) and occlusal (2) views. **K.** *Mustelictis* sp., left hemimandible with p4-m2, GAO408, in buccal (1), lingual (2) and occlusal (3) views. **L.** *Peignictis pseudamphictis*, part of left hemimandible with m2, GAO407, in buccal (1) and lingual (2) views. **P.** Musteloidea indet., right P4, GAO-413, buccal view. Scale bar: 1 cm.</figcaption></figure>

the first premolars of *Dinailurictis* are absent, like those of *Eusmilus*; those of *Eofelis* are much smaller, like those of *Nimravus* (Peigné, 2000: tables 1, 3). The lower first premolars of the amphicyonids and the Cephalogalini are low and are slightly elongated (Helbing, 1928: table 2, figs. 7–9; Bonis, 1973: pl. 1, fig. 5, pl. 2, fig. 5; Springhorn, 1977: figs. 45, 54; Ginsburg, 1966: fig. 8, 1977: fig. 6; Peigné and Heizmann, 2003: pl. 3, fig. 6).

Nevertheless, the amphicyonid P1 is taller and more pointed (Kuss, 1965: fig. 17; Bonis, 1973: pl. 6, fig. 3; Springhorn, 1977: fig. 36; Peigné and Heizmann, 2003: fig. 5) and closer to the morphology of UP-ROQ33. Thus, we assign UP-ROQ33 to the family Amphicyonidae.

Musteloidea Fischer, 1917

**Remarks:** The superfamily Musteloidea contains presently the families Mustelidae, Procyonidae, Ailuridae, and Mephitidae. The Mustelidae family exhibits a large diversity among its genera: in terms of size from the 30 g least weasel to the ~45 kg seaOtter, of ecology with generalists, fossorial, climbing and aquatic species, of diet from hyper-carnivorous to vegetarian, and body proportions from massive to elongated bodies. The origins and classification of Musteloidea were discussed for a long time. Nevertheless, in the last 30 years, with the development of cladistics and molecular phylogeny, many attempts were published on these topics (Bryant et al., 1993; Wolsan, 1993, 1999; Dragoo and Honeycut,
1997; Sato et al., 2004, 2009; Flynn et al., 2005; Fulton and Strobeck, 2007; Koepfli et al., 2007; Finarelli, 2008; Schnitzler et al., 2017; Law et al., 2018). These studies, particularly the molecular ones, solved the problem of relationships between the different extant families but that of the fossil remains is yet unclear despite an anatomical study of the skull of extant and extinct taxa (Schmidt-Kittler, 1981).

In Europe, the first wave of Musteloidea arrived with the large faunal immigration of the 'Grande Coupure', together with the amphicynodontids (see above), the nimravids, the ursids, some amphicyonids and some other mammal orders.

One of the most significant characters to separate the families of musteloids relates to the basis of the skull, particularly a depression in the roof of the auditory meatus called the supra-meatal fossa. It is shallow but present in the basal arctoid *Amphicynodon* with the most primitive pattern, and it displays differences in size and morphology among the musteloids. The extant ailurid (lesser panda) retains the same pattern. Another step is the hollow depression found in procyonids and some mustelids. The latter have more complicated patterns with a lateral extension into the bone, and, in some species, the loss of the depression. The mephiids are distinguished by an enlargement of the recessus epitympanicus in a huge cavity including probably the supra-meatus fossa (Schmidt-Kittler, 1981). So far the oldest known procyonid comes from the Phosphorites du Quercy with the skull of *Pseudobassaris riggi* Pohle, 1917 and another skull described later (Wolsan and Lange-Badré, 1996). Both specimens come from the old collections of the Quercy and thus they cannot be dated precisely, even if they are probably coming from Oligocene localities. The oldest known mephitid, *Palaeomephitis steinheimensis* Jäger, 1839, is also European, and was found in the middle Miocene of Steinheim, Germany, and re-studied more recently (Wolsan, 1999). Although primitive by some characters, the ailurids are not recorded before the Miocene with the European genera *Simoeycyon* Wagner, 1858 and *Magerictis* Ginsburg, Morales, Soria and Herráez, 1997.

Normally, the P4 (upper carnassial) of the Carnivora wears a notch between the paracone and the metastyle but in mustelids, the shearing blade may be continuous without solution of continuity, a derived character of the recent members of the group. Mustelids show a large panel of adaptations and are unique in the musteloids to contain hypercarnivore genera. The presence in the fossil record of specimens of Musteloidea with an evolutionary trend toward hypercarnivory is one of the signatures of the family.

**Genus *Huntictis*** nov. gen.

**Derivation of the name:** from ictis = weasel (Greek), and dedicated to R.M. Hunt Jr. for his works on the Quercy carnivorans.

**Type species:** *Huntictis minima* nov. gen., nov. sp.

**Age:** MP23, early Oligocene, ca. 30.5 Ma.

**Diagnosis:** small Oligocene musteloid (size close to that of the Feliformia *Palaeogale*); m1 primitive by its high trigonid with a

very well-developed metaconid as high as the paraconid and slightly distally and buccally inclined; narrow but basined talonid.

*Huntictis minima* nov. gen., nov. sp.

Fig. 6(E, M, N)

**Derivation of the name:** from minima = very small (Latin).

**Holotype:** UM-GAO415 (University of Montpellier), left m1.

**Locality:** Gardiol 3.

**Material:** Gardiol 3 (UM-GAO) 415, m1 (holotype).

**Age:** MP23, early Oligocene, ca. 30.5 Ma.

**Measurements:** see Table 4.

**Diagnosis:** as for the genus, by monotypy.

**Description:** The measurements of the m1 holotype (L = 5.3 mm, w = 2.7 mm) are close to those of *Palaeogale*, but its morphology is very different (Fig. 6(E)). The mesial root of the m1 of *H. minima* nov. gen., nov. sp., is partially broken off but the distal one is complete; both are robust and do not correspond to decidual tooth roots. The trigonid is high and pointed. The protoconid has a triangular section; its distal face is flat and almost vertical; the blade portion is relatively short. The top of the paraconid is steeply inclined and oblique relative to the row. There is a clear wear facet in the buccal surface of the blade made by paraconid and protoconid. No trace of wear on the top of the cuspids. The conical metaconid is large and as high as the paraconid; its top is slightly distally shifted and visible if buccal view. Both paraconid and metaconid are slightly puffy at their bases. The talonid is short and narrow but basined. There is a short hypocristid finishing by a conical hypoconid buccally convex but with a flat and steeped lingually face. There is no clear entoconid but a relatively thin and trenchant entocristid. The narrow talonid basin is closed by a hypoconulid separated by narrow notches from hypoconid and entocristid. This talonid differs from those of the other described adult musteloids from the "Phosphorites of Quercy".

Another tooth (UM-GAO508) could be a damaged germ of another m1. The enamel of the crown and the dentine of the roots are thin. The trigonid is complete with a well-developed metaconid higher than the paraconid. The talonid displays a large metaconid a low entocristid and a small hypoconulid. This cuspid is present in Eocene carnivoramorphans and its presence in the teeth of *Huntictis* nov. gen. could be a remembrance of an ancestral feature. A third m1, partially broken off (UM-GAO417), displays also the same characters as UM-GAO415 and belongs to the same taxon.

**Remarks:** *Huntictis* nov. gen. differs from most other Oligocene musteloids by its small size, the relatively shorter cutting carnassial blade and by its large and well-separated metaconid of m1. The morphology of m1 is close to that of the Feliformia *Anictis simplicidens* (Schlosser, 1887). *Anictis* Kretzoi, 1945 is considered to retain "a moderately plesiomorphic dentition" (Hunt, 1998) in the hypercarnivorous Oligocene aeluroids. But the type species, *A. simplicidens*, differs from *H. minima* nov. gen., nov. sp. by an almost two times larger size and a more reduced metaconid (Schlosser, 1887-1889: pl. IX, figs. 10, 34; Hunt, 1998: fig. 3).

The size and morphology of *Huntictis minima* nov. gen., nov. sp. m1 seem to be close to those of a m1 from the IVPP locality 9309, Daochazigou, China, (Wang et al., 2005: fig. 8) which is dated to early Miocene despite the presence of *Hyaenodon* in the locality. The description of the tooth is very close to that of *Huntictis* nov. gen., nov. sp., except for the small size of the entoconid cristid and the absence of hypoconulid: "This isolated carnassial, 4.6 mm long and 2.1 mm wide, is smaller than any known early Miocene carnivoran in Asia. The metaconid is well developed, slightly offset (located slightly posterior to) from the protoconid in occlusal view, and of approximately the same height as the paraconid. The talonid appears to be narrow, but this is partly due to

<table><caption>**Table 4**
Measurement table of lower teeth of *Huntictis minima* from Gardiol 3 (early Oligocene; 'Phosphorites du Quercy', southwestern France).</caption><thead><tr><th><i>Huntictis minima</i></th><th>m1</th><th>m1L</th><th>m1w</th></tr></thead><tbody><tr><td rowspan="5">Gardiol 3</td><td>mean</td><td>4.90</td><td>2.23</td></tr><tr><td>min</td><td>3.96</td><td>1.79</td></tr><tr><td>max</td><td>5.83</td><td>2.7</td></tr><tr><td>n</td><td>3</td><td>3</td></tr></tbody></table>
damage to its labial portion. The talonid is dominated by a centrally located hypoconid (only the lingual half is preserved), and the entoconid is poorly differentiated". These teeth perhaps indicate the presence in Eurasia of an unknown group of musteloids.

Musteloidea gen. et sp. indet.
Fig. 6(F)

**Material:** Gardiol 3 (UM-GAO) 416, m1.

**Description:** A lower carnassial (UM-GAO416) displaying the same length as that of *Huntictis minima* nov. gen., nov. sp. (5.2 x 2. 2 mm) differs by different proportions (Fig. 6(F)). The cuspid of the trigonid are lower; more massive and their heights are close to each other. The talonid is relatively short with two thick hypo- and ento-cristids limiting a wide and flat basin. It represents a more hypocarnivore adaptation. Nevertheless, its place among the carnivorans is difficult to establish precisely.

Mustelidae Fischer von Waldheim, 1817
Genus *Mustelictis* Lange, 1969.

**Type species:** ?Plesictis robustus Filhol in Teilhard de Chardin (1915: p. 58, pl. VII, fig. 13), by original designation.

**Remarks:** The genus was established on a skull from the old collections of the Quercy, thus without any precise dating (Lange, 1969, 1970). The skull belongs probably to the same genus with a part of several mandibles of the old collections of the Quercy described with the generic name "Plesictis". The latter genus was erected for fossils of the late Oligocene and early Miocene and its skull differs clearly from that of early and 'middle' Oligocene musteloids. Later, another species was included in *Mustelictis: M. olivieri* Bonis, 1997.

*Mustelictis olivieri* Bonis, 1997.
Fig. 6(G, H, J)

**Holotype:** UP-MGB60, skull (Bonis, 1997).

**Paratypes:** two hemimandibles, UP-MGB7 and UP-MGB60.

**Material:** Itardies (UP-ITD) 1063, md p2, 1818, md p2-m1, 1809, P4.

**Diagnosis:** small sized *Mustelictis*. Differs from *M. piveteaui* by a more flat promontorium rotunda fenestra visible in ventral view, fossa muscularis major less wide, shallower epitympanic recessus, narrow fossa jugularis, smaller infra-orbital foramen higher situated. Relatively shorter P4 with a less extended protocone and a less developed parastyle; narrower M1 in its middle part, relatively less reduced M2; lower carnassial with a well-developed metaconid and basined talonid with jutting out hypoconid; m2 with almost flat paraconid and talonid more developed than the trigonid.

**Description:** Two hemimandibles from the locality of Itardies and less complete specimens from Gardiol 3 (Fig. 6(G, H)) display the characters of *Mustelictis* (Bonis, 1997). The right hemimandible UP-ITD1063 (Fig. 6(G)) bears p2-m2 and oval alveolus of the one-rooted p1. The corpus is low (4.2 mm at the level of p2-p3, 5.4 mm for p4-m1, and 5.3 mm for m1-m2), and the lower border is slightly convex. The coronoid process is broken off at mid-height, the masseteric fossa is not very deep and it is limited anteriorly by a robust crest; a part of the condyle is preserved but the angular process is broken off. There is no diastema between the teeth; the premolars are thin and pointed with a faint basal cingulid; p2 is simple and slightly asymmetrical, the mesial part being smaller with a steeply anterior border; the mesial sagittal cristid turns lingually at the base to a very tiny mccd and there is a small dcsd; p3 is slightly larger and more asymmetrical than p2, and as tall as the paraconid of m1; there is a small mccd and a dcsd more devel-

oped than that of p2 with a tiny basin; p4 is taller than the paraconid of m1, it bears a mccd larger than that of p3, a well-developed pacd lowly situated on the sagittal ridge and followed by a dcsd with a small basin. The paraconid-protoconid shearing blade is well developed and crosses the sagittal plane with an acute angle, the protoconid being clearly taller than the other cuspid; the metaconid is moderately developed, conic and clearly separated from the protoconid; the hypoconid is high, relatively trenchant and the cristid oblica is directed to the lingual part of the distal wall of the protoconid, leaving a small basined place on the buccal side; the lingual face of the hypoconid descends gently to a low entocristid which continues to close the shallow talonid basin. The trigonid of m2 is complete, the paraconid is reduced but present in the mesiolingual corner of the tooth; protoconid and metaconid are separated, the almost pyramidal protoconid being taller than the conic metaconid – the last character frankly differs from what is seen in amphicynodontids –; the talonid is narrow, tapering distally and similar to that of m1.

The right hemimandible UP-ITD1818 bears p2-m1 and the alveoli of c, p1 and m2 (Fig. 6(H)). It is close to UP-ITD1063 although showing a slightly shallower corpus. The tooth measurements are close to each other and we think that both specimens belong to the same species. In Gardiol 3 *M. olivieri* is present with two fragments of mandible. UM-GAO405 has a gracile dentary with p3 and m2; p3 is asymmetric with a mesio-distal cutting edge, a mesial bit of cingulid and a small distal shelf; there is no paraconid in m2, the protoconid is taller and larger than the metaconid, a thin mesial cristid closes a small mesial basin, and the cuspidated talonid is distally tapering. UM-GAO406 is less gracile than UM-GAO405 and bears the roots of p2 and p3-p4; p3, asymmetric, is pointed and relatively thick, a cingulid circles the crown with a small mesial bump and closes a small distal basin; p4 is almost symmetric, taller than p3, it has a large cingulid giving a small mesial cuspid and a distal basin; the distal cutting edge bears a distal accessory cuspid. All these features are, at different degrees, present in *M. olivieri*.

The holotype of *M. olivieri* comes from Mas de Got, a MP22 locality, and the fossils from Itardies and Gardiol 3 fit very well those of Mas de Got. In addition, we consider that the specimens described from the localities La Plante 2 and Valbro, identified as *M. aff. olivieri* (Bonis et al., 2019) enter the variation of a single species with those of Itardies, Gardiol 3 and Mas de Got (Table 5). Taking into account the time (~1 myr) separating the two groups of localities, *M. olivieri* does not show significant morphological variations during this time interval in the Quercy, indicating the preservation of its ecological niche in this area. Other small carnivorans that did exist with *M. olivieri* are stenoplesictids which are more hypercanivores with certainly different ecological niches, and *Peignictis pseudamphictis* which differs from *M. olivieri* by m1 with a less open trigonid and an enlarged talonid of which the cristid oblica is parallel to the sagittal plane.

An isolated P4 from Itardies (UP-ITD1809) also belongs to the genus and fits *M. olivieri* (Fig. 6(J)).

?*Mustelictis* sp.
Fig. 6(K)

**Material:** Gardiol 3 (UM-GAO) 408, p4-m2.

**Measurements:** Table 5.

**Description and remarks:** UM-GAO408 is a hemimandible bearing p4-m2. Unfortunately, the m1 is badly preserved, weathered with several fissures, but enough preserved to see the main characters. Despite the fissures, we can see that p4 is relatively low, not surpassing the top of the paraconid of m1, and displays a slight mesial cingulid and a distal cristid closing a small basin; the top of the protoconid is broken off, the paraconid is obliquely
<caption>**Table 5**</caption>

Measurement table of lower and upper teeth of *Mustelictis olivieri* from Itardies and ?*Mustelictis* sp. from Gardiol 3 (early Oligocene; 'Phosphorites du Quercy', southwestern France).

<table><thead><tr><th></th><th></th><th></th><th></th><th></th><th></th><th></th><th></th><th></th><th></th><th></th><th></th><th></th><th></th><th></th></tr></thead><tbody><tr><td rowspan="4">Mustelictis olivieri</td><td>Lower teeth</td><td>p2L</td><td>p2w</td><td>p3L</td><td>p3w</td><td>p4L</td><td>p4w</td><td>m1L</td><td>m1trL</td><td>m1trw</td><td>m1tlw</td><td>m2L</td><td>m2w</td></tr><tr><td>Itardies</td><td>mean</td><td>3.00</td><td>1.48</td><td>3.87</td><td>2.01</td><td>4.33</td><td>1.65</td><td>5.23</td><td>3.27</td><td>2.65</td><td>2.05</td><td>3.03</td><td>2.00</td></tr><tr><td></td><td>min</td><td>2.89</td><td>1.45</td><td>4.04</td><td>2.61</td><td>4.31</td><td>1.74</td><td>5.15</td><td>3.17</td><td>2.50</td><td>2.10</td><td>—</td><td>—</td></tr><tr><td></td><td>max</td><td>3.11</td><td>1.51</td><td>3.70</td><td>1.41</td><td>4.34</td><td>1.55</td><td>5.30</td><td>3.37</td><td>2.79</td><td>1.99</td><td>—</td><td>—</td></tr><tr><td></td><td>n</td><td>2</td><td>2</td><td>2</td><td>2</td><td>2</td><td>2</td><td>2</td><td>2</td><td>2</td><td>2</td><td>1</td><td>1</td></tr><tr><td rowspan="4">?Mustelictis sp.<br>Gardiol 3</td><td>Lower teeth</td><td>p4L</td><td>p4w</td><td>m1L</td><td>m1trL</td><td>m1trw</td><td>m1tlw</td><td>m2L</td><td>m2w</td><td></td><td></td><td></td><td></td></tr><tr><td>mean</td><td>3.77</td><td>2.41</td><td>5.17</td><td>3.55</td><td>3.33</td><td>2.60</td><td>2.34</td><td>1.73</td><td></td><td></td><td></td><td></td></tr><tr><td>n</td><td>1</td><td>1</td><td>1</td><td>1</td><td>1</td><td>1</td><td>1</td><td>1</td><td></td><td></td><td></td><td></td></tr><tr><td>Upper teeth</td><td>P4L</td><td>P4w</td><td>M1L</td><td>M1w1</td><td>M1w2</td><td>M2L</td><td>M2w</td><td></td><td></td><td></td><td></td><td></td></tr><tr><td></td><td>mean</td><td>4.76</td><td>2.60</td><td>3.75</td><td>5.30</td><td>4.20</td><td>3.30</td><td>1.55</td><td></td><td></td><td></td><td></td><td></td></tr><tr><td></td><td>min</td><td>—</td><td>—</td><td>3.52</td><td>4.9</td><td>4.1</td><td>—</td><td>—</td><td></td><td></td><td></td><td></td><td></td></tr><tr><td></td><td>max</td><td>—</td><td>—</td><td>4.12</td><td>5.53</td><td>5</td><td>—</td><td>—</td><td></td><td></td><td></td><td></td><td></td></tr><tr><td></td><td>n</td><td>1</td><td>1</td><td>4</td><td>4</td><td>4</td><td>1</td><td>1</td><td></td><td></td><td></td><td></td><td></td></tr></tbody></table>

situated and the trigonid valley is largely open. The most striking feature is the shape of the talonid with a high and trenchant hypoconid similar to that of *Stenogale* but situated along the buccal side, the lingual part gently going down to a faint entocristid; m2 has a complete trigonid in which the vestigial paraconid is a small cristid separated from the protoconid by a notch; the metaconid is as tall as the protoconid and the talonid is a small copy of that of m1. These characters remain those of "Plesictis" *stenogalinus* Teilhard de Chardin, 1915. Nevertheless, the latter is larger and its metaconid of m1 is more distally shifted with a more open trigonid of m1. The size of this mandible is close to that of *Mustelictis olivieri* (Table 5) but it differs by the higher hypoconid of m1, relatively shorter p4, buccaly shifted talonid of m1, relatively higher hypoconid of m1, shorter m2, and relatively shorter talonid of m2. Thus, for the moment, UM-GAO408 could belong to *Mustelictis* but cannot be precisely identified.

Some upper teeth of Gardiol 3 match this mandible (Table 5). P4 is elongated with a high and slightly buccolingually compressed paracone. A parastyle is forwardly projected with a tiny cusp in the middle. There is a quite long metastyle blade. The protocone is mesio-lingually projected and slightly hollowed, while a cingulum runs along the lingual border. M1 is an irregular triangle with a mesial border longer than the distal one; the three cusps of the trigone are conic, the paracone being larger than the metacone. There is a globulous parastyle bearing a tiny cusp and a distally-directed metastyle. The protocone is encircled by a cingulum, while the paracone and metacone are separated from a buccal crest by a flat and slightly depressed area. A tiny cusp between the paracone and protocone could be a vestigial paraconule. The small M2 is extended transversally but is mesio-distally very narrow.

Genus *Peignictis* Bonis, Gardin and Blondel, 2019

**Emended diagnosis:** Small Mustelidae. Shallow corpus mandibula, deep masseteric fossa distally to the level of m2 but shallower in its upper part and gently diminishing; broad triangular coronoid process at its base but the distal border is oblique forward and narrows up to the condyle. Condyle highly situated; small, medially advanced and slightly down-directed angular process. High trigonid of m1 with protoconid far higher than the other cusps and with an almost vertical distal face; the protoconid and paraconid provide a clear buccal shearing surface; metaconid pointed and higher than the paraconid but not especially reduced or distally displaced. Complete trigonid of m2; large, almost flat trigonid basin with a small paraconid pointing in the mesiolingual corner; conical protoconid and metaconid, the former being slightly taller than the latter; high hypoconid and short entocristid; the talonid is not tapering distally as it is the case in *Mustelictis*.

*Peignictis pseudamphictis* Bonis, Gardin and Blondel, 2019
Fig. 6(L)

**Material:** Gardiol 3 (UM-GAO) 407, m2.

**Description:** A dentary from Gardiol 3 with a relatively shallow corpus, a large part of a triangular ascending process with a relatively deep masseteric fossa, and bearing m2 (GAO407) differs from the specimens studied above. The corpus is moderately robust with an ascending process low and triangular with a pointed top. The other known ascending processes of *Mustelictis* are tall with an axe-shaped or rounded top. The m2 is larger than that of *Mustelictis olivieri* but smaller than those of other species of *Mustelictis* (note here that the measurements given in Bonis et al., 2019: table 3 for M. cf. major are erroneous and come from a mistake in constructing the table – the described specimen does not bear any m2). There is a large, almost flat basin with a small paraconid pointing in the mesio-lingual corner; the protoconid and metaconid are conical, the former being slightly taller than the latter; there is a large notch between the protoconid and a high hypoconid while there is a small notch between the metaconid and a short entocristid which runs distally to close a narrow transversal basin; the distal part of the talonid is not tapering distally as it is the case in *Mustelictis*.

Another dentary with a high and probably triangular ascending ramus was described from the locality of Valbro (MP22) under the name *Peignictis pseudamphictis* (Bonis et al., 2019). The quite long m2, estimated from the alveoli, justified the specific assignment. The dentary UM-GAO407 fits the characters of this species. The slightly larger size (measurements of m2 "L = 3.1/w = 1.9" and L = 2.5 mm for *P. pseudamphictis*) could correspond to a measurement taken in alveoli with another one taken directly on the tooth or to the younger age of Gardiol 3 (MP23). The presence of m2 allows for completing the generic diagnosis.

## 4. Discussion

The MP23 (early Oligocene) Quercynian fossil localities allow the recording of about a dozen of species of Caniformia carnivorous. There are different types of karstic MP23 localities leaving carnivorous. Pech Crabit and Roqueprune are large and deep open cavities (emptied by the phosphorite workers by the end of the 19th century), the first one with a cliff of 15–20 m of clay with scattered bones into the clay, and the second with four places in the bottom containing remains of fossiliferous clay with scattered fossils. Other sites are smaller in surface and volume of fossiliferous sediments, such as Itardies which is located at the base of a lateral fissure of a large cavity containing a highly fossiliferous level of ca. 50–60 cm of height and a surface less than 2 m². Itardies is a major MP23 site in the Quercy. Nevertheless, large mammals such as herbivores and carnivores are very rare in this locality – there are some large bones or jaws of cadavers probably carried into the cavity such as small fragments of maxilla and some carpal
or tarsal bones of rhinocerotid. The bulk of the carnivoran fauna consists of the remains of small carnivorans in which *Amphicynodon leptorhynchus* is by far the dominant taxon, together with rodents and bats. The abundance of that species, the presence of anatomically associated limb bones, skull and mandibles, the absence of weathering and marks of carriage and the relative abundance of milk dentitions lead us to consider that the locality had been for a long time a den for these small carnivorans. The other innumerable remains of mammals are from bats and rodents, the latter being probably the preys of *Amphicynodon* and other small carnivorans. This concentration of remains attributed to *A. leptorhynchus* may indicate a gregarious behaviour. Currently, the coatis are the only living small scansorial Caniformia to live in groups, so the hypothesis of such a lifestyle for *A. leptorhynchus* is possible, but it could more likely be a succession of occupations by families. In the richest MP23 locality, Itardies, the carnivoran fauna is therefore slightly different from those of the other MP23 localities, likely because of the relative dispersion of the sites and local vegetation, small differences in time, size of the fissures and chance in fossilisation.

We note that Hyaenodonta were prosperous during the Eocene but their decline seems to have begun with the Oligocene (Solé et al., 2022). They are present in the Quercynian MP23 localities by few remains in Itardies (*Hyaenodon dubius* Filhol, 1873, and *Thereutherium thylacodes* Filhol, 1876). The Quercynian Hyaenodonta species have a low biostratigraphic significance such as *Hyaenodon gervaisi* Martin, 1906 (MP19 to MP29), *Hyaenodon dubius* Filhol, 1873 (MP21 to MP23), or *Hyaenodon leptorhynchus* Laizer and Parrieu, 1838 (MP22 to MP29). Many of them have no precise biochronological references (Lange-Badré, 1979). In Europe, the Oligocene is the time of the beginning of their end and conversely, the time of diversification of the Carnivora (Solé et al., 2022).

Nowadays, terrestrial carnivorans are widespread in many different environments ranging from boreal forests (taiga to temperate forests), tropical forests, steppes, grasslands and savannas, each environment having its own carnivoran fauna. Even if human activities have deeply modified the faunas and the environments over the last decades to millennia, remaining extant animals may give a quite good idea of the original faunas. The onset of the Oligocene witnessed significant environmental changes that had a profound impact on both fauna and flora (Legendre, 1989; Escarguel et al., 2008; Weppe et al., 2023). There was a notable decline in annual temperatures, an increase in rainfall seasonality, and a shift towards more open and arid environments. As a result, the once prevalent humid evergreen tropical forests in Central and Western Europe gradually gave way to warm temperate deciduous forests during the late Eocene, followed by more open wooded savannahs landscape in the early Oligocene (Knobloch et al., 1993; Hutchinson et al., 2021; Suc et al., 2025). Despite these changes, MP23 sites in southeastern France and Germany still revealed some remnants of flora and fauna characteristic of humid tropical zones (Knobloch et al., 1993; Duhamel and Louchard, 2019–2021). For instance, the latter authors describe a quite tropical bird fauna in Oligocene layers close to the Mount Luberon (Southeastern France), ca. 500 km from the Quercy. The Itardies site, with its diverse small carnivores and five genera of glirid rodents, suggests a relatively closed and wooded environment. This aligns with the climbing adaptations described for *A. leptorhynchus*, probably linked to an arboreal lifestyle. The Oligocene Caniformia may have reached quite large sizes with some Cephalogalini being close to a wolf size and some bear-like amphicyonids. Nevertheless, they are extremely rare in the study localities and only with few isolated teeth. This is not due to the different sizes of the fossil-bearing fissures but probably to the environment

which did not favour large hunter such as Cephalogalini, amphicyonids or large nimravids. The members of the bulk of the Quercynian small Caniformia were mesocarnivores or even hypocarnivores with some ecological niches that do not exist nowadays.

The amphicynodontids are principally known in French early Oligocene localities, from Ronzon (MP21) in the Massif Central to the Quercy (MP23) that contains the richest localities. They are also present in some localities in Hungary, Germany and Spain. *Amphicynodon* was also recorded, with few specimens, in North America (Wang et al., 2022) and in Asia (Matthew and Granger, 1924; Wang and Qiu, 2003; Wang et al., 2005), but when compared to southwestern France, these faunas are limited concerning the number of *Amphicynodon* species and individuals.

### CRediT authorship contribution statement

**Louis de Bonis:** Writing – original draft, Writing – review & editing, Data curation. **Axelle Gardin:** Writing – original draft. **Gilles Escarguel:** Writing – review & editing, Formal analysis.

### Data availability

Data will be made available on request.

### Declaration of competing interest

The authors declare that they have no known competing financial interests or personal relationships that could have appeared to influence the work reported in this paper.

### Acknowledgements

We are grateful to T. Pélissié (Association des Phosphatières du Quercy) for his generous and continuous help in the field; to the team of the Reserve Naturelle Nationale d'Intérêt Géologique du Lot and Parc Naturel Régional/Géoparc Mondial UNESCO des Causses du Quercy, as well as all the members of the Quercy research team (Institut des Sciences de l'Evolution de Montpellier, Montpellier; Laboratoire d'Ecologie des Hydrosystèmes Naturels et Anthropisés, Lyon; Muséum national d'Histoire naturelle, Paris; Paléontologie Evolution Paléoécosystèmes Paléoprimatologie, Poitiers) for their help and work in the field and in the laboratories; to the curators of the MNHN-Paris – Guillaume Billet, Claire Sagne, and Christine Argot; to Sabine Riffaut who helped for the photos; and to Edith Cirot for her past work on the studied material. G.E. acknowledges fundings by the French Agence Nationale de la Recherche ENLIVEN program (ANR-22-CE2-0014-01).

### References

* Aymard, A., 1846. Essai monographique sur un nouveau genre de mammifère fossile trouvé dans la Haute Loire, et nommé *Entelodon*. Annales de la Société d'Agriculture, Sciences, Arts et Commerce du Puy 12, 227-267.

* 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.

* BiochroM'97, 1997. Synthèses et tableaux de corrélations. In: Aguilar, J.P., Legendre, S., Michaux, J. (Eds.), Actes du Congrès BiochroM'97. Mémoires et Travaux de l'E. P.H.E., Institut de Montpellier 21, pp. 769-805.

* Bonis, L.de, 1966. Arrières-Crânes et moulages endocraniens de Carnivores fossiles. Annales de Paléontologie 52, 143-162.

* Bonis, L.D., 1973. Contribution à l'étude des mammifères de l'Aquitanien de l'Agenais: rongeurs, carnivores, périsnodactyles. Mémoires du Muséum National d'Histoire Naturelle, Paris, n.s., C, Science de la Terre 28, 1-192.

* Bonis, L.D., 1976. Découverte d'un crâne d'*Amphictis* (Mammalia, Carnivora) dans l'Oligocène supérieur des Phosphorites du Quercy. Comptes Rendus de l'Académie des Sciences de Paris, D 283, 327-330.
L. de Bonis, A. Gardin and G. Escarguel

Geobios xxx (xxxx) xxx

Bonis, L.D., 1997. Précisions sur l'âge géologique et les relations phylétiques de *Mustelictis olivieri* n. sp. (Carnivora, Mustelidae), carnassier de l'Oligocène inférieur (MP 22) des phosphorites du Quercy (France). Geobios MS 20, 55-60.

Bonis, L.D., Gardin, A., Blondel, C., 2019. Carnivora from theearly Oligocene of the 'Phosphorites du Quercy' in southwestern France. In: de Bonis, L., Werdelin, L. (Eds.), Memorial to Stéphane Peigné: Carnivores (Hyaenodonta and Carnivora) of the Cenozoic. Geodiversitas, pp. 601-621.

Bryant, H.N., Russell, A.P., Fitch, W., 1993. Phylogenetic relationships within the extant Mustelidae (Carnivora): appraisal of cladistic status of the Simpsonian subfamilies. Zoological Journal of the Linnean Society 108, 301-334.

Cirot, E., 1992. Etude phylogénétique de quelques genres d'Arctoidea de l'Oligocène eurasiatique, comparaisons des données morphologiques et moléculaires. Ph.D. Thesis, Université de Poitiers, France (unpubl.).

Cirot, E., Bonis, L.D., 1992. Révision du genre *Amphicynodon*, carnivore de l'Oligocène. Palaeontographica 220, 103-130.

Cirot, E., Bonis, L.D., 1993. Le crâne d'Amphicnis ambiguus (Carnivora, Mammalia): son importance pour la compréhension de la phylogénie des mustéloïdes. Comptes Rendus de l'Académie des Sciences de Paris, II 316, 1327-1333.

Clark, J., Guensburg, T.E., 1972. Arctoid genetic characters as related to the genus *Paricis*. Fieldiana Geology 26, 1-71.

Daubrée, M., 1871. Gisements dans lesquels la chaux phosphatée a été récemment découverte dans les départements du Tarn-et-garnie et du Lot. Comptes rendus de l'Académie des Sciences de Paris 73, 1028-1036.

Delfortrie, M., 1872. Les gites de chaux phosphatée dans le département du Lot: leur faune, le mode et l'époque probable de leur formation. Actes de la Société linnéenne de Bordeaux, 505-518.

Dragoo, J.W., Honeycut, R.L., 1997. Systematics of mustelid-like carnivores. Journal of Mammalogy 78, 426-443.

Duhamel, A., Louchard, A., 2019-2021. Regard sur l'avifaune fossile de l'Oligocène du Parc du Luberon et ses implications paléobiologiques, incluant de nouvelles données fossiles. Courrier scientifique du Parc naturel régional du Lubéron et de la réserve de biosphère Luberon-Lure 2021, 16-2019-2020, 98-117.

Durand-Delga, M., 2006. De la découverte des phosphorites du Quercy au renouveau de leur étude avec Bernard Gêze. Strata, Sér 1 (13), 25-36.

Escarguel, G., Legendre, S., 2006. New methods for analysing deep-time meta-
community dynamics and their application to the paleogene mammals from
the Quercy and Limagne area (Massif Central, France). Strata, Sér 1 (13), 243-
273.

Escarguel, G., Legendre, S., Sigé, B., 2008. Unearthing deep-time biodiversity changes: The Palaeogene mammalian metacommunity of the Quercy and Limagne area (Massif Central, France). Comptes Rendus Geoscience 340, 602-614.

Escarguel, G., Marandat, B., Legendre, S., 1997. Sur l'âge numérique des faunes de mammifères du Paléogène d'Europe occidentale, en particulier celles de l'Eocène inférieur et moyen. In: Aguilar, J.P., Legendre, S., Michaux, J. (Eds.), Montpellier, pp. 443-460.

Filhol, H., 1873. Sur les vertébrés fossiles trouvés dans les dépôts de phosphate de chaux du Quercy. Bulletin de la Société Philomathique de Paris 10, 85-89.

Filhol, H., 1882. Mémoires sur quelques mammifères fossiles des phosphorites du Quercy. Imp. Vialelle, Toulouse.

Filhol, 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 15, 1-220 (première partie); 16, 1-338 (deuxième partie).

Finarelli, J.A., 2008. A total evidence phylogeny of the arctoidea (Carnivora: Mammalia): relationships among basal taxa. Journal of Mammalian Evolution 15, 231-259.

Flynn, J.J., Finarelli, J.A., Zehr, S., Hsu, J., Nedball, M.A., 2005. Molecular phylogeny of the Carnivora (Mammalia): assessing the impact of increased sampling on resolving enigmatic relationships. Systematic Biology 54, 317-337.

Francheschi, D.D., Le Gall, C., Escarguel, G., Hugueney, M., Legendre, S., Simon-Coincon, R., Pélissié, T., Sigé, B., 2006. Une paléoflore des phosphatières du Quercy (Sud Ouest France): première découverte, résultats et perspective. Strata, Sér 1 (13), 97-101.

Fulton, T.L., Strobeck, C., 2007. Novel phylogeny of the raccoon family (Procyonidae: Carnivora) based on nuclear and mitochondrial DNA evidence. Molecular Biology Evolution 43, 1171-1177.

Gardin, A., Salesa, M.J., Siliceo, G., Anton, M., Pastor, F., Bonis, L.D., 2021. Climbing Adaptations of an Enigmatic Early Arctoid Carnivoran: the Functional Anatomy of the Forelimb of Amphicynodon leptorhynchus From the Lower Oligocene of the Quercy Phosphorites (France). Journal of Mammalian Evolution 28, 785-811.

Gardin, A., Salesa, M.J., Siliceo, G., Anton, M., Pastor, J.F., Bonis, L.D., 2022. The hindlimb of Amphicynodon leptorhynchus from the lower Oligocene of the Quercy Phosphorites (France): highlight of new climbing adaptations of this early arctoid. Journal of Mammalian Evolution 29, 815-843.

Gervais, P., 1872. Sur les mammifères dont les ossements accompagnent les dépôts à chaux phosphatée du Tarn-et-Garonne et du Lot. Comptes rendus de l'Académie des Sciences de Paris 74, 1367-1371.

Ginsburg, L., 1966. Les Amphicyons des Phosphorites du Quercy. Annales de Paléontologie (Vertébrés) 52, 23-44.

Helbing, H., 1928. Carnivore des oberen Stampien. Abhandlungen der
schweizerischen, paläontologischen Gesellschaft 47, 1-83.

Hunt Jr., R.M., 1974. The auditory bulla in carnivora: an anatomical basis for reappraisal of carnivore evolution. Journal of Morphology 143, 21-76.

Hunt Jr., R.M., 1998. Ursidae. In: Janis, C.M., Scott, K., Jacobs, L. (Eds.), Evolution of Tertiary Mammals of North America. Cambridge University Press, pp. 174-189.

Hutchinson, D.K., Coxall, H.K., Lunt, D.J., Steinthorsdottir, M., De Boer, A.M., Baatsen, M., Von Der Heydt, A., Huber, M., Kennedy-Asser, A.T., Kunzmann, L., Ladant, J.B., 2021. The Eocene-Oligocene transition: a review of marine and terrestrial proxy data, models and model-data comparisons. Climate of the Past 17, 269-315.

Knobloch, E., Kvaček, Z., Bůžek, C., Mai, D.H., Batten, D.J., 1993. Evolutionary significance of floristic changes in the Northern Hemisphere during the Late Cretaceous and Palaeogene, with particular reference to Central Europe. Review of Palaeobotanic and Palynology 78, 41-54.

Koeppli, K.P., Matthew, E., Gompper, M.E., Eizirik, E., Cheuk-Chung Ho, C.-C., Linden, L., Maldonado, J.E., Wayne, R.K., 2007. Phylogeny of the Procyonidae (Mammalia: Carnivora): Molecules, morphology and the Great American Interchange. Molecular Phylogenetics and Evolution 43, 1076-1095.

Kuss, S.E., 1965. Revision der Europäischen Amphicyoninae (Canidae, Carnivora, Mamm.) ausschliesslich der voroberstampischen Formen. In: Sitzungsberichte der Heidelberger Akademie der Wissenschaften - Mathematisch-naturwissenschaftliche Klasse, pp. 5-168.

Lange, B., 1969. Un nouveau mustéliné des Phosphorites du Quercy, *Mustelictis pivetaui*. Comptes rendus hebdomadaires des séances de l'Académie des sciences, Série D, Sciences naturelles 268, 2870-2872.

Lange, B., 1970. *Mustelictis pivetaui*, mustélidé nouveau des Phosphorites du Quercy. Annales de Paléontologie 56, 3-16.

Lange-Badré, B., 1979. Les Créodontes (Mammalia) d'Europe occidentale de l'Eocène supérieur à l'Oligocène supérieur. Mémoires du Muséum National d'Histoire Naturelle de Paris 42, 1-249.

Law, C.J., Slater, G.J., Mehtai, R.S., 2018. Lineage diversity and size disparity in musteloidae: testing patterns of adaptive radiation using molecular and fossil based methods. Systematic Biology 67, 127-144.

Legendre, S., 1989. Les communautés de mammifères du Paléogène (Éocène supérieur et Oligocène) d'Europe occidentale: structures, milieux et évolution. Münchener Geowissenschaftliche Abhandlungen A 16, 1-110.

Legendre, S., Bachelet, B., 1993. The numerical ages: A new method of datation applied to Paleogene mammalian localities from Southern France. Newsletters on Stratigraphy 29, 137-158.

Legendre, S., Marandat, B., Sigé, B., Crochet, J.-Y., Godinot, M., Hartenberger, J.L., Sudre, J., Vianey-Liaud, M., Muratet, B., Astruc, J.-G., 1992. La faune de mammifères de Vielase (phosphorites du Quercy, Sud de la France): preuve paléontologique d'une karstification du Quercy dès l'Eocène inférieur. Neues Jahrbuch für Geologie und Paläontologie - Monatshefte 1992 (7), 414-428.

Legendre, S., Sigé, B., Astruc, J.G., Bonis, L.D., Crochet, J.Y., Denys, C., Godinot, M., Hartenberger, J.-L., Lévêque, F., Marandat, B., Moure-Chauviré, C., Rage, J.-C., Remy, J.A., Sudre, J., Vianey-Liaud, M., 1997. Les Phosphorites du Quercy, 30 ans de recherches. Bilan et perspectives. Geobios 20, 331-345.

Matthew, W.D., Granger, W., 1924. New Carnivora from the Tertiary of Mongolia. American Museum Novitates 104, 1-9.

Mazières, B., 1956. L'interprétation des poches à phosphorites du Quercy. Revue géographique des Pyrénées et du Sud-Ouest 27, 105-121.

Merriam, C.W., 1930. *Allocyon*, a new canid genus from the John Day beds of Oregon. University of California Publications, Bulletin of the Department of Geological Sciences 19, 229-244.

Peigné, S., 2000. A new species of *Eofelis* (Carnivora, Nimravidae) from the Phosphorites of Quercy, France. Comptes Rendus de l'Académie des Sciences de Paris 330, 653-658.

Peigné, S., Heizmann, E.P.J., 2003. The Amphicyonidae (Mammalia, Carnivora) from Ulm-Westtangente (MN 2, Early Miocene), Baden-Württemberg, Germany - Systematics and Ecomorphology. Stuttgarter Beiträge zur Naturkunde (Geologie und Paläontologie) 343, 1-133.

Peigné, S., Vianey-Liaud, M., Pélissié, T., Sigé, B., 2014. Valbro : un nouveau site à vertébrés de l'Oligocène inférieur (MP22) de France (Quercy). Contexte géologique; Mammalia: Rodentia, Hyaenodontida, Carnivora. Annales de Paléontologie 100, 1-45.

Pélissié, T., Orliac, M.J., Antoine, P.-O., Biot, V., Escarguel, G., 2021. Beyond Eocene and oligocene epochs: the causes of the quercy geopark and the grande coupure. Geoconservation Research 4, 573-585.

Pélissié, T., Sigé, B., 2006. 30 millions d'années de biodiversité dynamique dans le paléokarst du Quercy. Journées Bernard Gêze. Strata sér 1 (13), 284.

Prothero, D.R., Emry, R.J., 1996. In: The Terrestrial Eocene-Oligocene Transition in North America. Cambridge University Press, p. 688.

Rabi, M., Bastl, K., Botfalvai, G., Evancis, Z., Peigné, S., 2018. A new carnivoran fauna from the late Oligocene of Hungary. Senckenberg Palaeobio Palaeoenvironment 98, 509-521.

Rémy, J.A., Crochet, J.-Y., Sigé, B., Sudre, J., Bonis, L.D., Vianey-Liaud, 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.

Sato, J.J., Hosoda, T., Wolsan, M., Suzuki, H., 2004. Molecular phylogeny of arctoids (Mammalia: Carnivora) with emphasis on phylogenetic and taxonomic position of the geotrr-ratgers and skunks. Zoological Science 21, 111-118.

Sato, J.J., Wolsan, M., Minami, S., Hosoda, S., Sinaga, M.H., Hiyama, K., Yamaguchi, Y., Suzuki, H., 2009. Deciphering and dating the red panda's ancestry and early adaptive radiation of Musteloidea. Molecular Phylogenetics and Evolution 53, 907-922.
L. de Bonis, A. Gardin and G. Escarguel

Geobios xxx (xxxx) xxx

Schlosser, M., 1899. Über die Bären und bärenähnlichen Formen des europäischen Tertiärs. Palaeontographica 46, 95–151.

Schlosser, M., 1887–1890. Die Affen, Lemuren, Chiropteren, Insectivoren, Marsupialier, Creodonten und Carnivoren des Europäischen Tertiars. Beiträge zur Palaontologishe Osterreich-Ungarns, Pt. I (Vol. 6: 1-224, 1887), Pt.II (Vol. 7: 225-386, 1888), Pt. III (Vol. 8, 387-492, 1890).

Schmidt-Kittler, N., 1981. Zur Stammesgechichte der marderverwandten Raubtiergruppen (Musteloidea, Carnivora). Eclogae Geologicae Helvetiae 74, 753–801.

Schnitzler, J., Theis, C., Polly, P.D., Eronen, J.T., 2017. Fossils matter – understanding modes and rates of trait evolution in Musteloidea (Carnivora). Evolutionary Ecology Resarch 18, 187–200.

Scott, W.B., 1993. The mammals of the Deep River beds. American Naturalist 27, 658–659.

Sigé, B., Aguilar, J.-P., Marandat, B., 1991. Extension au Miocène inférieur des remplissages phosphatés du Quercy, la faune de vertébrés de Crémat (Lot, France). Geobios 23, 497–502.

Simpson, G.G., 1945. The principles of classification and a classification of mammals. Bulletin of the American Museum Natural History 85, 1–350.

Solé, F., Fischer, V., Le Verger, K., Mennecart, B., Speijer, R.P., Peigné, S., Smith, T., 2022. Evolution of European carnivorous mammal assemblages through the Paleogene. Biological Journal of the Linnean Society 135, 734–753.

Springhorn, R., 1977. Revision der alttertiären europäischen Amphicyonidae (Carnivora, Mammalia). Palaeontographica 158, 26–113.

Stirton, R.A., 1960. A marine carnivore from the Miocene Clallam Formation, Washington: its correlation with nonmarine faunas. University of California Publications in Geological Sciences 36, 345–368.

Suc, J.-P., Fauquette, S., Popescu, S.M., Melinte-Dobrinescu, M.C., Huet, B., Sorrel, P., Semmani, M., Fournier, F., Bessedik, M., Zheng, Z., Truc, G., Ferry, S., Rubino, J.-L., Girard, V., Séranne, M., 2025. Late Eocene to Early Miocene environments, vegetation and climate in Southeastern France documented by palynology. Palaeogeography, Palaeoclimatology, Palaeoecology 659, 112625.

Teilhard de Chardin, P., 1915. Les carnassiers des phosphorites du Quercy. Annales de Paléontologie 9, 103–190.

Trutat, E., 1871. Sur les gisements de chaux phosphatée des environs de Saint-Antonin et de Caylus (Tarn-et-Garonne). Comptes rendus de l'Académie des Sciences de Paris 73, 1007-1010.

Vianey-Liaud, M., Legendre, S., 1986. Les faunes des phosphorites du Quercy: principes méthodologiques en paléontologie des mammifères; homogénéité chronologique des gisements de mammifères fossile. Eclogae Geologicae Helvetiae 79, 917–944.

Viranta, S., 1996. European miocene amphicyonidae – taxonomy, systematics and ecology. Acta Zoologica Fennica 204, 1–61.

Viret, J., 1929. Les faunes de mammifères de l'Oligocène supérieur de la Limagne bourbonnaise. Annales de l'Université de Lyon N.S. Science-Médecine 47, 1–327.

Wang, X.M., Emry, R.J., Boyd, C.A., Person, J.J., White, S.C., Tedford, R.H., 2023. An exquisitely preserved skeleton of *Eoarctos vorax* (nov. gen. et sp.) from Fitterer Ranch, North Dakota (early Oligocene) and systematics and phylogeny of North American early arctoids (Carnivora, Caniformia). Journal of Vertebrate Paleontology 42 (2), 1–123.

Wang, X.M., McKenna, M.C., Dashzeveg, D., 2005. *Amphicticeps* and *Amphicynodon* (Arctoidea, Carnivora) from hsanda gol formation, central mongolia and phylogeny of basal arctoids with comments on zoogeography. American Museum Novitates 3483, 1–57.

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.

Weppe, R., Condamine, F.L., Guinot, G., Maugoust, J., Orliac, M.J., 2023. Drivers of the artiodactyl turnover in insular western Europe at the Eocene-Oligocene transition. Proceedings of the National Academy of Sciences, USA 120, e2309945120.

Wolsan, M., 1993. Phylogeny and classification of early European Mustelida Mammalia: Carnivora). Acta Theriologica 38, 345–384.

Wolsan, M., 1999. Oldest mephitine cranium and its implications for the origin of skunks. Acta Palaeontologica Polonica 44, 223–230.

Wolsan, M., Lange-Badré, B., 1996. An arctomorph carnivoran skull from the Phosphorites du Quercy and the origin of Procyonids. Acta Palaeontologica Polonica 41, 277–298.