# The hyaenodonts (Mammalia) from the French locality of Aumelas (Hérault), with possible new representatives from the late Ypresian

Floréal Solé, Bernard Marandat, Fabrice Lihoreau

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Floréal Solé, Bernard Marandat, Fabrice Lihoreau. The hyaenodonts (Mammalia) from the French locality of Aumelas (Hérault), with possible new representatives from the late Ypresian. Geodiversitas, 2020, 42 (13), pp.185. 10.5252/geodiversitas2020v42a13. hal-04954869

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geodiversitas

2020 • 42 • 13

<figure><img src="image_2.png" /></figure>

The hyaenodonts (Mammalia) from the
French locality of Aumelas (Hérault),
with possible new representatives
from the late Ypresian

Floréal SOLÉ, Bernard MARANDAT & Fabrice LIHOREAU

<figure><img src="image_3.png" /></figure>

art. 42 (13) — Published on 4 June 2020

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# The hyaenodonts (Mammalia) from the French locality of Aumelas (Hérault), with possible new representatives from the late Ypresian

Floréal SOLÉ

D.O. Earth and History of Life, Royal Belgian Institute of Natural Sciences,
rue Vautier 29, B-1000 Brussels (Belgium)

Bernard MARANDAT
Fabrice LIHOREAU

Institut des Sciences de l'Évolution de Montpellier,
Univ. Montpellier CNRS IRD EPHE, Montpellier (France)

Submitted on 31 December 2018 | accepted on 3 September 2019 | published on 4 June 2020

urn:lsid:zoobank.org:pub:52D33BB7-2713-4F9D-AACF-3FAA90FDB31F

Solé F., Marandat B. & Lihoreau F. 2020. — The hyaenodonts (Mammalia) from the French locality of Aumelas (Hérault), with possible new representatives from the late Ypresian, *in* Bonis L. de & Werdelin L. (eds), Memorial to Stéphane Peigné: Carnivores (Hyaenodonta and Carnivora) of the Cenozoic. *Geodiversitas* 42 (13): 185-214. https://doi.org/10.5252/geodiversitas2020v42a13. http://geodiversitas.com/42/13

## ABSTRACT

We describe the carnivorous mammals from the Eocene French locality of Aumelas (Hérault, France). All the carnivores that have been recovered from the fossiliferous locality belong to the Hyaenodonta, a group that was very diverse during the Eocene of Europe. Three new species are described. *Matthodon peignei* n. sp. is intermediate in size between *M. menui* (Rich, 1971) and *M. tritens* Langer-Badré & Haubold, 1990. This is the first known occurrence of this genus in southwestern Europe. *Oxyaenoides aumelasiensis* n. sp. is distinct from other *Oxyaenoides* Matthes, 1967 species by its smaller size (15% smaller than *O. bicuspidens* Matthes, 1967). A third new species is proposed from Aumelas: *Leonhardtina meridianum* n. sp. The hyaenodonts from Aumelas appear to be morphologically similar to hyaenodonts from late Ypresian to early Lutetian of the northwestern Europe (European mammal reference level MP10 and MP11). These new species are incorporated into a phylogenetic analysis and are resolved within the clade Hyaenodontoidea. Our analysis interestingly recovered the Prionogalidae and *Thereutherium* as members of the Hyainailourinae. Based on the carnivorous faunal analysis, Aumelas is likely close to the Ypresian/Lutetian transition. However, because the hyaenodonts display unusual sizes that could result from provincialism, dates for Aumelas based on only these taxa should be treated cautiously, and will require confirmation based on a broader faunal analysis.

## KEY WORDS

Europe,
Carnivores,
Paleogene,
Endemism,
Tethys,
new species.
## RÉSUMÉ

Les hyénodontes (Mammalia) de la localité française d'Aumelas (Hérault), de possibles nouveaux représentants de l'Yprésien tardif.

Nous décrivons les mammifères carnassiers de la localité française d'Aumelas (Hérault, France). Tous les fossiles découverts dans cette localité appartiennent au Hyaenodonta, un groupe de carnassiers très diversifié en Europe durant l'Éocène. Trois nouvelles espèces sont reconnues. *Matthodon peignei* n. sp. est morphologiquement similaire aux deux espèces précédemment référées à ce genre – *M. menui* (Rich, 1971) and *M. tritens* Lange-Badré & Haubold, 1990 – mais elle est intermédiaire en taille. Ceci est la première occurrence de ce genre particulier dans le sud-ouest de l'Europe. *Oxyaenoides aumelasiensis* n. sp. se distingue des autres espèces d'Oxyaenoides Matthes, 1967 par une plus petite taille (15 % par rapport à *O. bicuspidens* Matthes, 1967, la plus petite espèce du genre). La troisième espèce appartient au genre *Leonhardtina* Matthes, 1952: *L. meridianum* n. sp. Les hyénodontes d'Aumelas sont morphologiquement proches des hyénodontes connus à l'Yprésien tardif et au début du Lutétien (niveaux repères MP10 et MP11) dans le nord-ouest de l'Europe. Ces nouvelles espèces ont été incluses au sein d'une analyse phylogénétique des hyaenodontes ; il ressort qu'elles appartiennent au clade des Hyaenodontoidea. Cette analyse phylogénétique se distingue des précédentes en ce qu'elle conduit à envisager les Prionogalidae comme des membres des Hyainailourinae. Sur la base de la faune carnivore, un âge proche de la transition Yprésien/Lutétien pour les fossiles d'Aumelas apparaît possible, contrairement à ce qui avait été proposé jusqu'à présent. Cependant, étant donné que les hyénodontes d'Aumelas présentent des tailles inhabituelles, dues vraisemblablement à du provincialisme, la datation de la localité sur la seule base des hyénodontes demeure incertaine et demande confirmation.

**MOTS CLÉS**
Europe,
carnivores,
Paléogène,
endémisme,
Téthys,
espèces nouvelles.

## INTRODUCTION

The fossil locality of Aumelas was discovered 20 km west of Montpellier (Hérault, France) by Mr. Gabinaud and was first reported by Hartenberger (1963). The first mammals described from this locality were *Propalaeotherium* sp. (cf. *parvulum*) and *Lophiodon isselensis*. These identifications were revised by Sudre (1980) and Rémy *et al.* (2016). Other mammals, notably the adapid primate *Anchomomys* aff. *gaillardi* (Hartenberger *et al.* 1969; later considered as *A. cf. gaillardi* by Godinot 1988) and the artiodactyl *Dichobune robertiana* (Sudre 1988; Sudre & Ginsburg 1993), suggested the locality was slightly older than Lissieu (France), or could be correlated to La Défense or Bouxwiller (France), and thus possibly be close to the MP13 reference-level (late Lutetian) (BiochroM'97 1997). The faunal list of Aumelas is being developed and is currently composed of crocodilians (Martin 2016), chelonians (Hervet 2004) and at least 19 mammalian taxa (marsupials, bats, rodents, lipot-hyphlan, primates, artiodactyls and perissodactyls; see Crochet 1979; Sudre 1980, 1988; Godinot 1988; Escarguel 1999; Mai-tre *et al.* 2006, 2008; Maitre 2014; Rémy *et al.* 2016, 2019).

This study is a contribution to a larger effort to review and revise the Aumelas fauna based on the collections at the Uni-versity of Montpellier and new materials collected over the last decade. Sudre (1980) mentions that the first identification of the carnivorous mammals from Aumelas (Proviverrinae indet) was made by B. Lange-Badré. No carnivoraform has been dis-covered at Aumelas. We here describe three new hyaenodonts based on unpublished fossils from Aumelas. Comparisons with recently published material of Hyaenodonta (Solé *et al.* 2014a,

2015a) and the addition of this material to a recent phylogeny (Borths & Stevens 2019) allows us to assess the previously proposed age of Aumelas. These new materials also permit us to compare the composition of carnivorous faunas at different European Eocene localities

## MATERIAL AND METHODS

### FOSSIL SPECIMENS

The fossils described herein are all from the locality of Aumelas (Hérault, France). Fossil collecting at Aumelas began in 1960 and has continued under the auspices of a team from the Uni-versity of Montpellier. The fossils are removed from the matrix using acid preparation, an effort lead by one of us (B.M.), with additional preparation at the Royal Belgian Institute of Natu-ral Sciences. The fossils are deposited in the collection of the University of Montpellier under the acronym UM-AUMXXX.

### DENTAL NOMENCLATURE AND MEASUREMENTS

The nomenclature of the teeth follows the dental terminology of Szalay (1969). The measurements (length × width in mm) follow Gingerich & Deutsch (1989).

### TAXONOMY

Solé & Mennecart (2019) and Dubied *et al.* (2019) modified the Borths & Stevens (2017c) matrix. Their phylogenetic study refuted the monophyly of “Proviverrinae” sensu Solé (2013), resolving “proviverrines” as part of the hyaenodontine stem lineage. Because their results agree with those of Borths *et al.*
(2016), they proposed to consider the Proviverrinae as a clade that includes the last common ancestor of *Proviverra* and *Parvagula*. They thus applied the name Hyaenodontoidea to the clade that includes the last common ancestor of *Proviverra* and *Hyaenodon*. This resolves Hyaenodontidae and Proviverrinae within Hyaenodontoidea.

However, the combination here presented of the Solé & Mennecart (2019) matrix, with the one published by Borths & Stevens (2019), results in the exclusion of *Parvagula* from the Proviverrinae. Therefore, we here consider that Proviverrinae includes the following hyaenodont genera: *Proviverra*, *Morlodon*, *Allopterodon*, and *Lesmesodon*. This analysis also results in the exclusion of *Parvagula* from the Hyaenodontoidea.

Authorships of taxa are indicated in the Appendix 8.

## PHYLOGENETIC ANALYSIS

We did not build a new character-taxon matrix but we combined the matrices published by Solé & Mennecart (2019), Dubied *et al.* (2019), and Borths & Stevens (2019). However, we have some disagreements with the coding of some European hyaenodonts in the character-taxon matrix created by Borths & Stevens (2017c, 2019). We thus decided to modify this matrix in order to reflect our observations (Appendix 1). It is worth noting that the Borths & Stevens (2019) matrix does not include the following taxa, which were however included in the Borths & Stevens (2017c) matrix: *Thereutherium thylacodes*, *Pakakali rukwaensis*, and the prionogalids *Namasector siaeae* and *Prionogale breviceps*. We added these taxa into our matrix. We also added the new taxa described herein: *Matthodon peignei* n. sp., *Oxyaenoides aumelasiensis* n. sp., and *Leonhardtina meridianum* n. sp. Moreover, as in Solé & Mennecart (2019), three European hyaenodont OTUs (Operational Taxonomic Unit) *Matthodon menui*, *Prodissopsis eocaenicus*, and the genus *Paracynohyaenodon* have been included in the matrix. The resulting character-taxon matrix includes 156 characters (Appendix 2) and 103 OTUs (Appendix 3). We also corrected the numerical ages used for the European hyaenodonts (Appendix 4).

The character-taxon matrix was assembled in Winclada (Nixon 2002). As in Borths & Stevens (2017c, 2019), nineteen multistate characters were treated as ordered, and all characters were treated as equally weighted. Phylogenetic analysis was performed using the Bayesian ‘tip-dating’ phylogenetic methods described by Beck & Lee (2014) and applied to hyaenodonts by Borths *et al.* (2016). This method simultaneously estimates branch length, rate, phylogeny and support for clades recovered in the analysis, a common approach for paleontological systematic analysis (e.g., Beck & Lee 2014; Dembo *et al.* 2015; Borths *et al.* 2016; Gorscak & O’Connor 2016; Lund *et al.* 2016; Turner *et al.* 2017). Bayesian ‘tip-dating’ analyses were performed in MrBayes (Ronquist *et al.* 2012). MrBayes formatted nexus files with all analytical parameters are included in the Appendix 5. Appendix 6 can be used to visualize the results (e.g., age estimates, rate of change) with FigTree.

## BODY MASS

The body mass of each new species is estimated based on the methodology provided by Morlo (1999). The equation is

$$ \text{Log}^{10}(P) = [3.5104 \times \text{Log}^{10}((\Sigma M)/3)] - 2.3469 $$ where P is the estimated body mass (in grams) and ΣM the sum of the length of the three lower molars (in mm). All the values are available in Appendix 7.

## ABBREVIATIONS

<table><tr><td>MNHN.F</td><td>Muséum national d'Histoire naturelle, Paris, Palaeontology collections;</td></tr><tr><td>RZ</td><td>collection from Rouzilhac, Aude, France;</td></tr><tr><td>UM</td><td>Université de Montpellier, France;</td></tr><tr><td>UM-AUM</td><td>Material from locality of Aumelas, Hérault, France deposited in the UM;</td></tr><tr><td>L</td><td>length;</td></tr><tr><td>n</td><td>number of specimens;</td></tr><tr><td>OR</td><td>observed range;</td></tr><tr><td>W</td><td>width.</td></tr></table>

## SYSTEMATICS

Order HYAENODONTA Van Valen, 1967

Family HYAENODONTIDAE Leidy, 1869

Genus *Matthodon* Lange-Badré & Haubold, 1990

TYPE SPECIES. — *Matthodon tritens* Lange-Badré & Haubold, 1990.

OTHER SPECIES. — *Matthodon menui* (Rich, 1971); *Matthodon peignei* n. sp.

*Matthodon peignei* n. sp.
(Fig. 1)

urn:lsid:zoobank.org:act:F79F2003-3B67-466B-AA9E-03F94442912F

HOLOTYPE. — UM-AUM694, left mandible with p2-m3 and the alveoli for the canine and p1.

ETYMOLOGY. — Dedicated to the memory of Dr. S. Peigné (1972-2017), who described numerous carnivorous mammals from the Paleogene and Neogene of Eurasia and Africa.

REFERRED SPECIMENS. — UM-AUM268, left mandible with m2 and m3.

HORIZON AND AGE. — Lacustrine limestone of Montpellier, proposed as Lutetian in age in previous study but here considered possibly late Ypresian/early Lutetian in age

TYPE LOCALITY. — Aumelas, Hérault, France.

DIAGNOSIS. — *Matthodon peignei* n. sp. is intermediate both metrically and morphologically between *Matthodon menui* and *Matthodon tritens*: *M. peignei* n. sp. differs from *M. menui* by having longer premolars and molars and from *M. tritens* by having mesiodistally shorter premolars and molars. *Matthodon peignei* n. sp. differs from *M. tritens* by having less developed paraconids on p2, p3 and p4; the larger and more individualized metaconid on the molars; the longer talonid on the m1 and m2; and by shorter molars (4% for the m1, 8% for the m2 and m3), and longer p2 and p4 (respectively 9% and 5%). *Matthodon peignei* n. sp. differs from *M. menui* and *M. tritens* by the relative larger size of the p2, and narrower premolars.

MEASUREMENTS. — Table 1.
<table><caption>TABLE 1. — Measurements (in mm) of *Matthodon peignei* n. sp. from Aumelas (MP10-MP11?). Body mass (c. 14.95 kg) estimated after Morlo (1999).</caption><thead><tr><th>Locus</th><th></th><th>n</th><th>OR</th><th>Mean</th></tr></thead><tbody><tr><td rowspan="2">c</td><td>L</td><td>1</td><td>14.4</td><td>–</td></tr><tr><td>W</td><td>1</td><td>11.46</td><td>–</td></tr><tr><td rowspan="2">p1</td><td>L</td><td>1</td><td>6.06</td><td>–</td></tr><tr><td>W</td><td>1</td><td>4.76</td><td>–</td></tr><tr><td rowspan="2">p2</td><td>L</td><td>1</td><td>9.32</td><td>–</td></tr><tr><td>W</td><td>1</td><td>5.66</td><td>–</td></tr><tr><td rowspan="2">p3</td><td>L</td><td>1</td><td>9.67</td><td>–</td></tr><tr><td>W</td><td>1</td><td>5.92</td><td>–</td></tr><tr><td rowspan="2">p4</td><td>L</td><td>1</td><td>11.81</td><td>–</td></tr><tr><td>W</td><td>1</td><td>6.85</td><td>–</td></tr><tr><td rowspan="2">m1</td><td>L</td><td>1</td><td>11.59</td><td>–</td></tr><tr><td>W</td><td>1</td><td>6.22</td><td>–</td></tr><tr><td rowspan="2">m2</td><td>L</td><td>2</td><td>12.56-12.89</td><td>12.75</td></tr><tr><td>W</td><td>2</td><td>7.33-8.14</td><td>7.74</td></tr><tr><td rowspan="2">m3</td><td>L</td><td>2</td><td>12.31-13.00</td><td>12.66</td></tr><tr><td>W</td><td>2</td><td>7.88-8.26</td><td>8.07</td></tr><tr><td>MD</td><td></td><td>1</td><td>26.27</td><td>–</td></tr></tbody></table>

## DESCRIPTION

UM-AUM 694 is almost a complete mandible that bears well-preserved teeth, except the i3, canine and p1, which are represented by their alveoli (Fig. 1A-C). The mandible is deep (26.3 mm below the m1). Two mental foramina are present: the first one opens below the posterior root of p2, while the second is present below the anterior root of p4. The masseteric fossa is very deep. A deep fossa for insertion of the temporal muscle is present along the anterior margin of the crest. The angular process is curved sharply dorsally and is relatively wide. The mandibular condyle is robust, and is mediolaterally wide.

The presence of an incisor (possibly the i3) is indicated by its tiny alveolus in front of the canine. The canine is buccolingually wide. The canine, premolars and molars are very close together. The p1 is single-rooted. The teeth display crenulated enamel. The p2 is simple with only the hypoconid present on the talonid. A small fold indicates the position of the paraconid. The p3 is mesiodistally longer and buccolingually wider than the p2. It differs by the presence of a paraconid, and by having three cusps on the talonid: the larger corresponds to the hypoconid, while the other two are situated in the position of the entoconid. The p4 is more symmetrical in lateral view than the p3. The paraconid is much more developed. On the talonid, two cusps are present: the hypoconid and the entoconid, which form a postfossid.

The three molars display similar morphologies. The m3 is slightly mesiodistally shorter than the m2, but this is due to the reduction of the talonid. The paraconid is prominent and located mesially. The anterior keel is well developed. The metaconid is lower than the paraconid and almost completely fused with the protoconid. The latter is distinctly the highest cusp. The talonid is mesiodistally short and narrow. The postfossid is narrow and surrounded by three bulbous cusps: entoconid, hypoconid and hypoconulid; the entoconid being the tallest one. The m3 differs from m1 and m2 in having a much more reduced talonid in which the talonid cusps can-

<table><caption>TABLE 2. — Mean width/length ratio of the p2, p3 and p4 respectively in the *Matthodon* species.</caption><thead><tr><th>Species</th><th>p2</th><th>p3</th><th>p4</th></tr></thead><tbody><tr><td><i>Matthodon menui</i></td><td>0.65<br>(OR: 0.63-0.67, n=2)</td><td>0.66<br>(OR: 0.65-0.67, n=2)</td><td>0.6<br>(OR: 0.57-0.63, n=2)</td></tr><tr><td><i>Matthodon tritens</i></td><td>0.71<br>(OR: 0.69-0.73, n=4)</td><td>0.67<br>(OR: 0.54-0.75, n=5)</td><td>0.65<br>(OR: 0.59-0.68, n=4)</td></tr><tr><td><i>Matthodon peignei</i><br>n. sp.</td><td>0.61<br>(n=1)</td><td>0.61<br>(n=1)</td><td>0.58<br>(n=1)</td></tr></tbody></table>

not be distinguished. The precingulid is short (limited to the paraconid) but is very wide, notably on m3.

## DISCUSSION

The new fossils share with *Matthodon* species the peculiar distal positions of the mental foramina (the first is inferior to the distal margin of p1 and the second is inferior to p4), the presence of a single-rooted p1, a closely packed toothrow, and the poorly developed metaconid on molars. All these features distinguish *Matthodon* from the European Ypresian and Lutetian hyaenodonts. UM-AUM694 is among the best preserved mandibles ever found for *Matthodon*.

The genus *Matthodon* is represented by two species: *Matthodon menui* from Chavot, Monthelon, Mancy, and Grauves (Ypresian, MP10; France) and *Matthodon tritens* from Geiseltal-Untere Unterkohle (Lutetian, MP11; Germany).

The fossils from Aumelas differ from those of the Lutetian species, *Matthodon tritens*, by having less developed paraconids on p2, p3 and p4; the larger and more individualized metaconid on the molars; the longer talonid on the m1 and m2; and by shorter molars (4% for the m1, 8% for the m2 and m3), and longer p2 and p4 (respectively 9% and 5%). These differences are reminiscent of the Ypresian species *Matthodon menui*.

The fossils from Aumelas, are distinct from those referred to *Matthodon menui* by having longer premolars and molars: the molars are 6% longer, the p2 9% longer, and the p3 and p4 4% longer. The mesio-distal diameter of the alveolus of the p1 is also distinctly longer than those measured for *M. menui*. As indicated above, the teeth from Aumelas are morphologically more similar to those of *M. menui* than to those of *M. tritens*.

The mean width/length ratio of the p2, p3 and p4 differen- tiates *Matthodon peignei* n. sp. from *M. menui* and *M. tritens* by having narrower premolars (Table 2).

The fossils from Aumelas appear intermediate both metri- cally and morphologically between Ypresian *M. menui* and Lutetian *M. tritens*. It differs from the two previously described taxa by the relatively large size of the p2. This feature is highly characteristic because the relative size of the p2, which decreases from *M. menui* to *M. tritens*, can be considered as an indication of the reduction of the anterior dentition (p1 and p2). The larger size of the p2 in UM/AUM 694 thus can be regarded as a primitive feature. The same observation can be made for the narrowness of the premolars.
<figure><img src="image_5.png" /><figcaption>FIG. 1. — *Matthodon peignei* n. sp. from Aumelas (MP10-MP11?): **A-C**, UM-AUM694 (Holotype), left mandible bearing the p2-m3 and the alveoli of the canine and p1; **A**, occlusal view; **B**, lingual view; **C**, labial view; **D-F**, UM-AUM268, left mandible bearing the m2 and m3; **D**, occlusal view; **E**, lingual view; **F**, labial view.
Scale bars: 1 cm.</figcaption></figure>
<table><caption>TABLE 3. — Measurements (in mm) of *Oxyaenoides aumelasiensis* n. sp. from Aumelas (MP10-MP11?). *, based on roots and/or alveoli; Body mass (c. 4.24 kg) estimated after Morlo (1999).</caption><thead><tr><th>Locus</th><th></th><th>n</th><th>OR</th><th>Mean</th></tr></thead><tbody><tr><td rowspan="2">P3</td><td>L</td><td>2</td><td>9.88-11.10</td><td>10.49</td></tr><tr><td>W</td><td>2</td><td>6.39-6.78</td><td>6.59</td></tr><tr><td rowspan="2">P4</td><td>L</td><td>2</td><td>11.28-11.55</td><td>11.42</td></tr><tr><td>W</td><td>2</td><td>7.63-8.75</td><td>8.19</td></tr><tr><td rowspan="2">M1</td><td>L</td><td>2</td><td>9.39-9.52</td><td>9.46</td></tr><tr><td>W</td><td>1</td><td>10.54</td><td>–</td></tr><tr><td rowspan="2">M2</td><td>L</td><td>1</td><td>8.32</td><td>–</td></tr><tr><td>W</td><td>1</td><td>10.11</td><td>–</td></tr><tr><td rowspan="2">p2</td><td>L</td><td>1</td><td>5.88</td><td>–</td></tr><tr><td>W</td><td>1</td><td>2.31</td><td>–</td></tr><tr><td rowspan="2">m1</td><td>L</td><td>1</td><td>8.71*</td><td>–</td></tr><tr><td>W</td><td>1</td><td>4.65*</td><td>–</td></tr><tr><td rowspan="2">m2</td><td>L</td><td>1</td><td>7.74*</td><td>–</td></tr><tr><td>W</td><td>1</td><td>5.16*</td><td>–</td></tr><tr><td rowspan="2">m3</td><td>L</td><td>1</td><td>9.24</td><td>–</td></tr><tr><td>W</td><td>1</td><td>5.37</td><td>–</td></tr><tr><td>MD</td><td></td><td>1</td><td>16.68</td><td>–</td></tr></tbody></table>

Based on these comparisons the Aumelas specimen represents a new species of *Matthodon*, the first occurrence of the genus in the southwestern part of Europe.

### Genus *Oxyaenoides* Matthes, 1967

**TYPE SPECIES.** — *Oxyaenoides bicuspidens* Matthes, 1967.

**OTHER SPECIES.** — *Oxyaenoides schlosseri* (Rütimeyer, 1891); *Oxyaenoides lindgreni* (Rich, 1971); *Oxyaenoides aumelasiensis* n. sp.

### *Oxyaenoides aumelasiensis* n. sp.
(Fig. 2)

urn:lsid:zoobank.org:act:AFFF7C27-FB65-49C0-A788-96352B214851

**HOLOTYPE.** — UM-AUM693, left mandible bearing m3, the anterior root and talonid of the m1, the posterior alveolus of the p4, and the alveoli of m2.

**ETYMOLOGY.** — Latin, from Aumelas (type locality).

**REFERRED SPECIMENS.** — UM-AUM686, right M2; UM-AUM692, combination of a left maxillary fragment bearing the P2-M1, a right maxillary fragment bearing the P3-P4, the protocone of the M1 and the alveoli of the P2; UM-AUM695, labial part of a M1.

**HORIZON AND AGE.** — Lacustrine limestone of Montpellier, proposed as Lutetian in age in previous study but here considered possibly late Ypresian/early Lutetian in age

**TYPE LOCALITY.** — Aumelas, Hérault, France.

**DIAGNOSIS.** — The species is 15% smaller than *O. bicuspidens* and 25% smaller than *O. bicuspidens* and *O. schlosseri*. It differs from *O. lindgreni* and *O. bicuspidens* by a more buccally aligned metastyle on molars. It differs from *O. bicuspidens* and *O. schlosseri* by a less retroflexed protoconid.

## MEASUREMENTS. — Table 3.

### DESCRIPTION

The P2 has a very small lingual fold that corresponds to an incipient parastyle. The metastyle is short and low (Fig. 2D, E). There is a small diastema between the P2 and P3 (Fig. 2F, G). The P3 and P4 are mesiodistally elongated. The P3 is characterized by the presence of a distinct parastyle and a strong metastyle. There is a large lingual bulge but no protocone. The P4 also displays a very large parastyle and metastyle; the latter is very tall. The protocone projects lingually and is mesiodistally narrow (Fig. 2D-G).

The two known molars have a very similar morphology (Fig. 2D, E, H, I). On M1, the parastyle is present and bulbous but is mesiodistally short and narrow. The metastyle is mesiodistally elongated. The paracone and metacone are partially fused; only the apexes are fully separated. The protocone is low. The protofossa is very mesiodistally short and narrow. The paraconule is present. The metaconule is absent. The M1 is morphologically identical to the M2 except that the parastyle is wider and the metastyle is less distally located.

The mandible is narrow. The masseteric fossa is deep. As in *Matthodon*, a deep fossa for insertion of the temporal muscle is present along the anterior margin of the coronoid crest. The latter is distally inclined and the angle between the coronoid crest and the body of the dentary is close to 120°.

UM/AUM693 is a fragment of a left mandible that bears the talonid of m1 and a complete m3 (Fig. 2A-C). The talonid of the m1 bears three distinct cusps (entoconid, hypoconid, and hypoconulid). The entoconid is lower than the hypoconulid, which is distally located, and lower than the hypoconid. Contrary to the m1, the m3 is complete. The paraconid is tall, long and well mesially located. There is no metaconid. The m3 is thus characterized by a cutting trigonid. The talonid is very mesiodistally short, low, and narrow with only one cusp (hypoconulid?) present. The precingulid is developed.

### DISCUSSION

The fossils from Aumelas are characterized by a very cutting morphology (e.g., long metastyle on P4 and molars, absence of metaconid, long paracristid on m3). Only one hyaenodont genus known from the Ypresian and Lutetian of Europe has molars that are characterized by the absence of a metaconid, a hypercarnivorous feature: *Oxyaenoides* (MP10-?MP13) (Matthes 1967; Lange-Badré & Haubold 1990; Solé *et al.* 2014a). The teeth found at Aumelas share with *Oxyaenoides* mesiodistally short and narrow talonids bearing crestiform entoconids on molars. This genus is known both in northwestern (Matthes 1967; Lange-Badré & Haubold 1990; Solé *et al.* 2014a, 2015a) and southwestern Europe (Crochet *et al.* 1976).

Three species of *Oxyaenoides* are presently known: *Oxyaenoides lindgreni* from MP10 (Rich 1971; Solé *et al.* 2014a), *Oxyaenoides bicuspidens* from MP11 (Matthes
<figure><img src="image_6.png" /></figure>

<figure><img src="image_7.png" /><figcaption>FIG. 2. — *Oxyaenoides aumelasiensis* n. sp. from Aumelas (MP10-MP11?: **A-C**, UM-AUM693 (Holotype), left mandible bearing m3, the anterior root and talonid of the m1, and the posterior alveolus of the p4, and the alveoli of m2; **A**, occlusal view; **B**, lingual view; **C**, labial view; **D-G**, UM-AUM692, combination of a left maxillary fragment bearing the P2-M1, a right maxillary fragment bearing the P3-P4, the protocone of the M1 and the alveoli of the P2; **D**, occlusal view of left maxillary; **E**, labial view of the left maxillary; **F**, occlusal view of the right maxillary; **G**, labial view of the right maxillary (reversed view); **H-I**, UM-AUM686, right M2; **H**, occlusal view; **I**, labial view. Scale bars: A-G, 1 cm; H, I, 50 mm.</figcaption></figure>
<table><caption>TABLE 4. — Measurements (in mm) of *Leonhardtina meridianum* n. sp. from Aumelas and Rouzilhac (MP10-MP11?). Body mass (c. 843.29 g) of *L. meridianum* n. sp. from Aumelas estimated after Morlo (1999).</caption><thead><tr><th rowspan="2">Locus</th><th rowspan="2"></th><th colspan="2">Leonhardtina meridianum n. sp.</th></tr><tr><th>from Aumelas</th><th>from Rouzilhac</th></tr><tr><th></th><th></th><th>n</th><th>OR</th><th>n</th><th>OR</th></tr></thead><tbody><tr><td rowspan="2">M2</td><td>L</td><td>1</td><td>4.96</td><td>0</td><td>–</td></tr><tr><td>W</td><td>1</td><td>6.07</td><td>0</td><td>–</td></tr><tr><td rowspan="2">M3</td><td>L</td><td>1</td><td>2.91</td><td>0</td><td>–</td></tr><tr><td>W</td><td>1</td><td>5.49</td><td>0</td><td>–</td></tr><tr><td rowspan="2">p2</td><td>L</td><td>0</td><td>–</td><td>1</td><td>6</td></tr><tr><td>W</td><td>0</td><td>–</td><td>1</td><td>2.7</td></tr><tr><td rowspan="2">p3</td><td>L</td><td>1</td><td>6.5</td><td>1</td><td>7</td></tr><tr><td>W</td><td>1</td><td>3.19</td><td>1</td><td>3.2</td></tr><tr><td rowspan="2">p4</td><td>L</td><td>1</td><td>5.39</td><td>1</td><td>5.4</td></tr><tr><td>W</td><td>1</td><td>2.51</td><td>1</td><td>2.7</td></tr><tr><td rowspan="2">m1</td><td>L</td><td>1</td><td>4.95</td><td>0</td><td>–</td></tr><tr><td>W</td><td>1</td><td>2.94</td><td>0</td><td>–</td></tr><tr><td rowspan="2">m2</td><td>L</td><td>1</td><td>5.49</td><td>0</td><td>–</td></tr><tr><td>W</td><td>1</td><td>3.21</td><td>0</td><td>–</td></tr><tr><td rowspan="2">m3</td><td>L</td><td>1</td><td>5.78</td><td>1</td><td>5.9</td></tr><tr><td>W</td><td>1</td><td>3.67</td><td>1</td><td>4</td></tr><tr><td>MD</td><td></td><td>1</td><td>10.48</td><td>1</td><td>10.68</td></tr></tbody></table>

1967; Crochet *et al.* 1976; Lange-Badré & Haubold 1990) and *Oxyaenoides schlosseri* from ?MP13 (Rütimeyer 1891; Lange-Badré 1972; Solé *et al.* 2015a). The syntypes of *O. schlosseri* are from Egerkingen γ (Lange-Badré 1972), but the MP reference-level of this site (possibly MP13) is uncertain (BiochroM’97 1997), as is Aigues-Vives 2 where a mandible of *O. schlosseri* was discovered (Solé *et al.* 2015a).

The most striking feature of the fossils from Aumelas is their small size compared to other *Oxyaenoides* material. The teeth are 15% smaller than those of *O. lindgreni* and 25% smaller than those of *O. bicuspidens* and *O. schlosseri*. The size of the new taxon is actually almost equivalent to that of *Preregidens langebadrae* (Saint-Papoul; c. MP8+9).

*Oxyaenoides aumelasiensis* n. sp. differs from those of *O. lindgreni* and *O. bicuspidens* by a more buccally aligned metastyle on upper molars. It also differs from *O. bicuspidens* and *O. schlosseri* by a mesially oriented protoconid – this feature accommodates an elongate preprotocristid, a structure found on cutting molars. *Oxyaenoides aumelasiensis* n. sp. appears more primitive than northwestern species by having a buccally aligned metastyle on upper molars and the less retroflexed protoconid on molars.

To conclude, the fossils of *Oxyaenoides aumelasiensis* n. sp. share with the other *Oxyaenoides* species several features including: a reduced metaconid and reduced talonid morphology. However, based on the small size and distinctive features, we distinguish *O. aumelasiensis* n. sp. from previously described species of *Oxyaenoides*.

The fossils from Aumelas are important because they are the first to provide so much information on the morphology of the upper premolars of the genus *Oxyaenoides*. The fossils from Aumelas represent the smallest hypercarnivorous hyae-nodont ever discovered in the Ypresian and Lutetian.

## Genus *Leonhardtina* Matthes, 1952

**TYPE SPECIES.** — *Leonhardtina gracilis* Matthes, 1952.

**OTHER SPECIES.** — *Leonhardtina godinoti* Solé, Falconnet & Laurent, 2014; *Leonhardtina meridianum* n. sp.

### *Leonhardtina meridianum* n. sp.
(Fig. 3)

urn:lsid:zoobank.org:act:DA2667CF-BFAD-4CFD-AFE0-E9B1416140D2

**HOLOTYPE.** — UM-AUM54, right mandible bearing p3 to m3.

**ETYMOLOGY.** — *Meridianum* (Latin) means from the Midi, the south. In reference to the fact that the species is recorded only in the southern part of France.

**REFERRED SPECIMENS FROM AUMELAS.** — UM-AUM530, right maxillary fragment bearing M2 and M3.

**OTHER REFERRED SPECIMENS.** — MNHN.F.RZ142, right mandible bearing p2-p4 and m3.

**HORIZON AND AGE.** — Lacustrine limestone of Montpellier, proposed as Lutetian in age in previous study but here considered possibly late Ypresian/early Lutetian in age

**TYPE LOCALITY.** — Aumelas, Hérault, France.

**OTHER LOCALITY.** — Rouzilhac, Aude, France (Ypresian-Lutetian; MP10b, Godinot *et al.* 2018)

**DIAGNOSIS.** — The new species is smaller than *Leonhardtina godinoti* (5% based on the length of the molars) and *Leonhardtina gracilis* (20%). It also differs from these species by a narrower postfossil on molars, and by relatively larger p3 relative to p4.

**MEASUREMENTS.** — Table 4.

## DESCRIPTION

The upper teeth are represented by the M2 and M3 (Fig. 3A, B). The M2 is characterized by the separation of the paracone and metacone. These cusps appear to be of similar height. The parastyle is well-developed. The metastyle is long and mainly buccally aligned. The protocone is extended buccolingually and mesiodistally short. The protofossa is worn. The M3 is buccolingually elongated. The parastyle is long. Only the paracone is visible; the metacone-metaconule area is broken. The protocone is almost as tall as the paracone. A paraconule is discernable.

The mandible is deep (Fig. 3C, D). The coronoid crest is slightly distally inclined; the angle between the coronoid crest and the body of the dentary is close to 100°. As in the two other hyaenodonts from Aumelas, a deep fossa for the insertion of the temporal muscle is present along the anterior margin of the coronoid crest.

The p3 is elongated mesiodistally and is longer than the p4 (Table 4). The tooth is too damaged to describe its morphology (Fig. 3F-H). The p4 probably had a paracoid, but the anterior part of the tooth is too damaged to be sure. The protoconid is robust. The talonid is short in
<figure><img src="image_8.png" /><figcaption>FIG. 3.—*Leonhardtina meridianum* n. sp. from Aumelas (MP10-MP11?): **A**, **B**, UM-AUM530, right maxillary fragment bearing M2 and M3; **A**, occlusal view; **B**, labial view; **C-H**, UM-AUM54 (Holotype), right mandible bearing p3 to m3; **C**, lingual view; **D**, labial view; **E**, occlusal view; **F**, occlusal close-up; **G**, lingual close-up; **H**, labial close-up. Abbreviations: **me**, metacone; **pa**, paracone; **pr**, protocone. Scale bars: 1 cm.</figcaption></figure>
length and bears two cusps: a hypoconid and a lingually located entoconid. The molars are similar in morphology except the m3 has a narrower and more mesiodistally elongated talonid than m1 and m2 (Fig. 3F-H). The three teeth have a sharply pointed and mesiodistally compressed trigonid. The paraconid is lingually located relative to the protoconid. The metaconid is buccolingually aligned with the protoconid. The apexes of the metaconid and paraconid are worn preventing comparison of cusp height; the paraconid was however probably shorter in height than the metaconid. The talonid is almost as wide as the trigonid. The postfossid is narrow. The three talonid cusps are well individualized and bulbous. The talonid is labially oriented relative to the horizontal ramus as in primitive hyaenodonts (e.g., *Prototomus, Eoproviverra*). The hypoconulid is slightly taller than the two other talonid cusps. The hypoconulid and entoconid are close together. On the molars, the three talonid cusps are well-defined. The precingulid seems to have been present as well as an incipient postcingulid, but this is uncertain due to taphonomic processes. No ectocingulid is present.

## DISCUSSION

The fossils of *Leonhardtina meridianum* n. sp. described differ from those of *Matthodon* and *Oxyaenoides* by their smaller size; the p3 longer than the p4; the wide talonid on the molars; and the sharply pointed and mesiodistally compressed trigonid of the molars.

According to our phylogenetic analysis (see below), *Leonhardtina* is closely related to *Preregidens langebadrae*. This latter species is only represented by one mandible discovered at Saint-Papoul (MP8+9; France) (Solé *et al.* 2015a). *Preregidens* shares with *Leonhardtina* the presence of mesiodistally short trigonid and buccolingually narrow postfossid on molars. However, *Preregidens* differs from *L. gracilis* and *L. meridianum* n. sp. by having buccolingually narrower talonid on molars, the absence of postcingulid on molars, shorter talonid on m3, and possibly the p3 shorter than the p4 (the p3 is only represented by its alveoli on the sole known fossil of *Preregidens*). *Preregidens* also differs from *L. gracilis* by having a higher and sharper p4.

The p3 longer than the p4, the wide talonid on the molars, and the sharply pointed and mesiodistally compressed trigonid of the molars are features that characterize the small hyaenodonts *Leonhardtina*, *Proviverra*, *Lesmesodon*, and *Allopteroodon*, which form a clade in Solé *et al.* (2014a, 2015a). This clade is not recovered in the recent analyses (e.g., Borths & Stevens 2017c, 2019) nor in the phylogenetic analysis we performed (Fig. 4).

*Leonhardtina meridianum* n. sp. differs from *Proviverra*, *Lesmesodon* and *Allopteroodon* by having weakly developed labial cingulids on the p4 and the molars; the latter are indeed well-developed in the three European hyaenodontoid genera, notably *Proviverra* (Lange-Badré 1981; Morlo & Habersetzer 1999). *Leonhardtina* further differs from these genera by having more oblique cristid obliqua, and narrower talonids and postfossids on the molars.

Some species of the genera *Cynohyaenodon*, *Paracynohyaenodon*, and *Quercytherium* are also close in size to *Leonhardtina*. However, these genera clearly differ from *Leonhardtina* by the p3 shorter than the p4 and a mesiodistally elongated trigonid on molars. These features also distinguish *Leonhardtina* from *Boritia*, a monospecific genus phylogenetically close to *Cynohyaenodon*, *Paracynohyaenodon*, and *Quercytherium*.

The weakly developed labial cingulids on the molars are also found in late Ypresian and early Lutetian genus *Leonhardtina*. This genus is represented by two species: *L. godinoti* from Grauves (Ypresian, MP10; Solé *et al.* 2014a) and *L. gracilis* from Geiseltal-Untere Mittelkohle and Geiseltal-Obere Mittelkohle (Lutetian, MP12-13; Matthes 1952; Lange-Badré & Haubold 1990). The fossils from Aumelas are smaller than other *Leonhardtina* species: 5% smaller than *L. godinoti* and 20% smaller than *L. gracilis*. The newly described fossils are 15-20% larger than those of *Proviverra typica*, which is found with *L. gracilis* in Geiseltal (Lange-Badré & Haubold 1990).

The new fossils also differ from the two species of *Leonhardtina* by having a relatively larger p3. The ratio of p3 length to p4 length equals 1.21 for *L. meridianum* n. sp. The ratio equals 1.04 in *L. godinoti* and 1.08 in *L. gracilis*. However, the ratio in *L. meridianum* n. sp. may be overestimated because the p3 and p4 are crushed, which probably results in lengthening the p3 measurement.

Godinot *et al.* (2018) described a mandible discovered in the locality of Rouzilhac (Aude, France; MP10b) that they referred to *Leonhardtina cf. gracilis*. The teeth preserved on the mandible are very close in size to those preserved on UM-AUM54 (Table 4). Moreover, the p4 and m3 are morphologically identical (e.g., talonid mesiodistally compressed on m3). Because of these similarities, we propose that MNHN.F.RZ142 and UM-AUM54 belong to the same taxon.

The p3 and p4 are better preserved on the fossil from Rouzilhac: the ratio p3 length/p4 length equals 1.3 in this specimen. This is consistent with *L. meridianum* n. sp. having a relatively large p3 compared to *L. godinoti* and in *L. gracilis*.

*L. meridianum* n. sp. differs from the other *Leonhardtina* species by having narrower talonids and postfossids and a more oblique cristid obliqua on the molars. The morphology of the talonids of the molars on UM-AUM54 are similar to the earliest European hyaenodonts such as *Eoproviverra* (Rians; Ypresian, *c.* MP7) and *Parvagula* ( Palette, Fournes, Fordones; Ypresian, *c.* MP7) rather than *Leonhardtina*, *Allopteroodon* or *Proviverra*.

The upper teeth (M2 and M3) are poorly preserved and not very diagnostic. However, they are morphologically similar to those of *Leonhardtina gracilis* in having a separated paracone and metacone; a buccally aligned metastyle; and a lingually elongated protocone.

To conclude, the *Leonhardtina* species found at Aumelas is smaller than the two *Leonhardtina* species known in north-western Europe, and has the potentially primitive feature of a narrower talonid and postfossid on molars. However, *L. meridianum* n. sp. has the potentially derived feature of a relatively long p3.
<figure><img src="image_9.png" /></figure>

<figure><img src="image_10.png" /><figcaption>FIG. 4. — Results of the phylogenetic analysis of Hyaenodonta character-taxon matrix. Results are visualized as an “all compact” (majority rule plus compatible groups) consensus tree. Major, named clades recovered or discussed in this analysis and recovered in other analyses are illustrated. In bold: new hyaenodonts species described herein.</figcaption></figure>
## PHYLOGENETIC ANALYSIS

The character-taxon matrix that forms the basis of the present analysis has been used – and improved – in several studies of hyaenodonts systematics (Borths et al. 2016; Borths & Seiffert 2017; Borths & Stevens 2017a, b, c, 2019; Solé & Mennecart 2019; Dubied et al. 2019). The analysis (“all compact” consensus tree; Fig. 4; Appendix 6) recovered all the clades found in the previous phylogenetic analyses.

The “all compact” consensus tree shows the monophyly of the Hyainailouroidea (Posterior Probability = 44%), the Hyainailouridae (PP = 46%), as well as that of the Teratodontinae (PP = 30%), Hyainailourinae (PP = 33%), and Apterodontinae (PP = 99%), three subfamilies of the Hyainailouroidea. These values are close to those found by Borths & Stevens (2019). The relationships among Hyainailouroidea have been extensively discussed by Borths et al. (2016), Borths & Seiffert (2017), and Borths & Stevens (2017a, b, 2019). The clades established by our analysis are similar to those found by Borths & Stevens (2019). However, numerous polytomies within the Hyainailourinae are present in the resulted tree of our analysis (Fig. 4), which was not the case in the previous published phylogenetic analyses.

The most dramatic difference with the previous phylogenetic studies is the presence of the enigmatic *Thereutherium* and of the prionogalids *Namasector* and *Prionogale* within the Hyainailourinae – the clade Prionogalidae+*Thereutherium* is supported by a PP of 57%, while the Prionogalidae is supported by a PP of 100%. This position differs from the results of Solé & Mennecart (2019): indeed, they were resolved as sister taxa of Hyaenodontinae by this analysis. A close relationship with Hyaenodontinae was also found by Borths & Stevens (2017c) when the characters were ordered. When the characters were unordered, the Prionogalidae (i.e., *Namasector* and *Prionogale*) were located within the Hyainailourinae, but *Thereutherium* was still closely related to the Hyaenodontinae (Borths & Stevens 2017c). Therefore the presence of both Prionogalidae and *Thereutherium* among the Hyainailourinae was unexpected.

This result questions the relationships of the Prionogalidae and *Thereutherium*, which highlights in addition the difficulty to understand the origin of these enigmatic hyaenodonts. Because the prionogalids have been only found in Africa (Schmidt-Kittler & Heizmann 1991; Morales et al. 2008; Borths & Stevens 2019), their presence among Hyainailourinae, which in all probability originated in Africa (Solé et al. 2014b, 2015b; Borths & Stevens 2019), is biogeographically logical. The case of *Thereutherium* is more complex, this small-bodied hypercarnivorous hyaenodont being only known in the Oligocene of Europe (Lange-Badré 1979). Based on the present topology, one can imagine that this hyaenodont dispersed from Africa into Europe in the earliest Oligocene. Such dispersal (i.e., from Africa to Europe in the early Oligocene) of a hyaenodont has been already proposed for *Apterodon* (Simons & Gingerich 1976; Lange-Badré & Böhme 2005; Grohé et al. 2012).

Some polytomies are also present at the nodes located closely to the Hyainailouroidea. A polytomy is also present close to the base of the Hyaenodonta: this node actually groups the Hyainailouridae and related taxa (e.g., *Galecyon* clade), the Hyaenodontoidea+*Parvagula*, and the Limnocyoninae and related taxa (*Arfia*, *Prototomus*, and *Sinopa*).

Besides the position of *Thereutherium* and Prionogalidae within the Hyainailourinae, the second striking difference with the previous phylogenetic studies is the very basal position of the Moroccan hyaenodonts *Lahimia* and *Boualitomus*; only the European hyaenodont *Eoproviverra* is more basally located than these hyaenodonts. This position is surprising because these carnivorous mammals were generally found to be close to the Hyainailouroidea (Borths et al. 2016, Borths & Seiffert 2017; Borths & Stevens 2017a, b, c). However, one can remember that *Lahimia* and *Boualitomus* were more basally located in Borths & Stevens (2019) (i.e., before the split between the Hyae-nodontidae and the Hyainailouroidea). The basal position of these two hyaenodont genera is interesting because the presence of *Lahimia* in the Paleocene of Morocco, has been notably used as an argument for an African origin of the Hyaenodonta by Solé *et al.* (2009) and Solé (2013); Gheerbrant (1995) and Gheerbrant *et al.* (2006) also pro-posed this hypothesis but before the discovery of *Lahimia*. This hypothesis has been notably criticized by Borths *et al.* (2016) and Borths & Stevens (2019). However, the issue of the geographic origin of the hyaenodonts is beyond the present research and should be analyzed only based on biogeographic analysis such as those performed by Borths *et al.* (2016) and Borths & Stevens (2019).

Contrary to the previous phylogenetic analyses (Borths & Seiffert 2017; Borths & Stevens 2017a,b,c), but similarly to Solé & Mennecart (2019), some of the “proviverrines” sensu Solé (2013) (i.e., *Proviverra*, *Lesmesodon*) are not located at the base of the hyaenodont clade, but are more deeply nested in the present phylogenetic tree. Therefore, our analysis is significant in that it recovers almost all the “proviverrine” taxa sensu Solé (2013) within the same clade except *Eoproviverra* (Fig. 4). In the present analysis *Eoproviverra* is the earliest-diverging hyaenodont (Fig. 4).

In the previous analyses (Borths et al. 2016; Borths & Seiffert 2017; Borths & Stevens 2017a, c), the Proviverrinae is a clade located at the base of the Hyaenodonta and generally included the European genera *Proviverra*, *Eoproviverra*, *Lesmesodon*, *Parvagula*, and *Morlodon*. This group also included *Allopterodon* in Borths et al. (2016), but this genus is closer to the Hyaenodontinae in the other analyses.

Comparably to the results of Solé & Mennecart (2019) and Dubied et al. (2019), we found a clade (PP = 55%) that gathers the European hyaenodont genera *Proviverra*, *Allopterodon*, *Lesmesodon*, and *Morlodon*. As discussed in Solé & Mennecart (2019), this clade partially corresponds to the concept of Proviverrinae of Borths et al. (2016). Borths et al. (2016) and Solé & Mennecart (2019) considered that the Proviverrinae is a clade that includes the common ancestor
of *Proviverra* and *Parvagula*. However, in the present analysis, *Parvagula* is outside this clade and is the sister taxon to a clade that groups the *Proviverrinae* and *Hyaenodontidae* (Fig. 4). Therefore, we here propose that the *Proviverrinae* is a clade that includes the common ancestor of *Proviverra* and *Morlodon*. From the recent phylogenetic analyses, it seems that the content of the *Proviverrinae* is susceptible to vary in the future, but we think that this concept is useful and should be conserved.

As in Solé & Mennecart (2019) and Dubied *et al.* (2019), the clade *Proviverrinae* is not located at the base of the *Hyaenodonta* but is the sister clade of the *Hyaenodontidae*. This result supports the concept of *Hyaenodontoidea* proposed by Solé & Mennecart (2019) to designate the clade formed by the common ancestor of *Proviverra* and *Hyaenodon* (Fig. 4).

Among the *Proviverrinae*, the *Allopterodon* clade (*Allopterodon*, *Proviverra*, *Lesmesodon*) of Solé *et al.* 2014a is partially recovered (PP = 71%). Contrary to Solé & Mennecart (2019), the genera *Lesmesodon* and *Proviverra* are not sister taxa; the proviverrines *Allopterodon* and *Proviverra* are indeed more closely related (PP = 83%).

All the other “proviverrine” taxa *sensu* Solé (2013) are located among the *Hyaenodontidae* (PP = 41%), as most of the previous phylogenetic analyses (e.g., Borths *et al.* 2016; Borths & Stevens 2019; Solé & Mennecart 2019). In this clade, only the *hyaenodontines* (i.e., *Propterodon* and *Hyae-nodon*) are not “proviverrine” taxa *sensu* Solé (2013). The *Hyaenodontinae*, which contains the genera *Propterodon* and *Hyaenodon*, is supported by a PP of 90% (Fig. 4). The genera *Cynohyaenodon*, *Paracynohyaenodon*, and *Quercytherium* form the *Cynohyaenodon* clade (according to Solé *et al.* 2014a) (PP = 95%); the genus *Cynohyaenodon* is not monophyletic (Fig. 4). This clade is closely related to *Leonhardtina* (PP = 30%). This – poorly supported – relationship was not found neither in Solé & Mennecart (2019) nor in Dubied *et al.* (2019). The new *Leonhardtina* species described herein is closely related to *Leonhardtina gracilis* (PP = 80%).

There is a clade (PP = 59%) that contains the *Hyaenodontinae* and the *hyaenodontids* *Eurotherium*, *Prodissopsalis*, *Matthodon*, and *Oxyaenoides*. *Eurotherium* and *Prodissopsalis* are closely related (PP = 100%) and constitute the most basal clade. One can note that the genus *Eurotherium* is not monophyletic: *E. theriodis* is closely related to *Prodissopsalis* (PP = 84%) (Fig. 4).

The other *hyaenodontids* are in a clade that has a PP of 89%. The next clade to diverge is *Matthodon* (PP = 79%) (Fig. 4). The new species, *M. peignei* n. sp., is closely related to *Matthodon menui* (PP = 94%). This supports the generic referral of the new *hyaenodont* from Aumelas to this genus. *Oxyaenoides* (PP = 85%) is the *hyaenodontid* genus most closely related to *Hyaenodontinae*. The new species, *O. aumelasiensis* n. sp. is closely related to *O. bicuspidens* (PP = 46%) (Fig. 4). As for *Leonhardtina* and *Matthodon*, the phylogenetic analyses confirm our observations based on dental material discovered at Aumelas. It is worth noting that the *hyaenodontines*, *Matthodon*, and *Oxyaenoides* dis-

play hypercarnivorous dentition characterized by the loss of the metaconid and reduction of the talonid, an adaptation that appeared several times among *Hyaenodontinae* (Borths *et al.* 2016).

Finally, one can note the presence of a clade (PP = 26%) that gathers the *Limnocyoninae* (PP = 99%), and the genera *Arfia*, *Prototomus*, and *Sinopa*. This clade is commonly recovered in the phylogenetic analyses that deal with the relationships within *Hyaenodonta*.

## DISCUSSION

### IMPLICATIONS OF THE HYAENODONT FAUNA CONCERNING THE AGE OF AUMELAS

The carnivorous mammals from Aumelas are represented by three new species, representing three previously described genera – *Oxyaenoides*, *Leonhardtina*, and *Matthodon*. It is important to note that no carnivoran is known from this locality. This is not surprising because in localities close to the MP10, MP11 (except Messel, Germany, where two carnivoran are known; Springhorn 1980, 2000), MP12, MP13, and MP14 reference-levels, carnivoran are rarer and less diverse than the *hyaenodonts*.

If the number of carnivorous taxa is low, the morphological disparity shown by the fossils is high: *Matthodon* and *Oxyaenoides* represent large (*Matthodon*: 15 kg) and medium-sized (*Oxyaenoides*: 4 kg) hypercarnivorous predators, while *Leonhardtina* represents a small – probably insectivorous – predator (843 g).

Specimens referred to *Oxyaenoides* have been discovered at several localities in southwestern Europe: Palassou (Crochet *et al.* 1976), Aigues-Vives 2 (Solé *et al.* 2015a), and Rouzilhac (Godinot *et al.* 2018). *Leonhardtina* was only recently discovered in the south of France (*Leonhardtina* cf. *gracilis* in Godinot *et al.* 2018 but *Leonhardtina meridianum* n. sp. herein). This is the first occurrence of *Matthodon* from the southwestern Europe, a genus that was previously only known from the northwest (Lange-Badré & Haubold 1990; Solé *et al.* 2014a). *Oxyaenoides* and *Leonhardtina* have been discovered together in the locality of Rouzilhac. The *Leonhardtina* species at Rouzilhac is similar to *L. meridianum* n. sp., but the *Oxyaenoides* species (*O. bicuspidens*) is larger than *Oxyaenoides aumelasiensis* n. sp. found at Aumelas and is closer in size to the *Oxyaenoides* species known from the northwestern Europe (Godinot *et al.* 2018).

The occurrence of these three *hyaenodonts* at Aumelas has interesting implications for relative age of the site. *Oxyaenoides* and *Leonhardtina* have long stratigraphic ranges: the former is known from localities close to the MP10, MP11 and ?MP13 reference-levels (Lange-Badré & Haubold 1990; Solé *et al.* 2014a, 2015a), while the latter is known from localities close to the MP10, MP12 and MP13 reference-levels (Lange-Badré & Haubold 1990; Solé *et al.* 2014a). The stratigraphic record of *Matthodon* is shorter, having only been reported from localities close to the MP10 and MP11 reference-levels (Lange-Badré & Haubold 1990; Solé *et al.* 2014a). Its particular ecology as a
<table><caption>TABLE 5. — Hypothetical correlations between the two mammalian successions of the Northwestern and Southwestern European provinces during the Ypresian and earliest Lutetian, with indication of the hyaenodonts recorded for the Northwestern and Southwestern provinces localities. Correlations based on BiochroM'97 (1997) and Marandat *et al.* (2012). PE biozones are from Hooker (1996, 1998). In bold: localities and fossils described and discussed in the present article: *, taxa not recorded in the type-locality of the reference-level. Note: the age of Aumelas and Rouzilhac is uncertain as the two localities are considered to be close to MP10 and MP11 reference-levels with uncertainty (see text); they are here grouped in a MP10b level, intermediate in age between MP 10 and MP 11, as proposed by Godinot *et al.* (2018) for Rouzilhac. Authorships of taxa are indicated in the Appendix 8.</caption><thead><tr><th>ELMA</th><th>Ma</th><th>Biozone</th><th>MP</th><th>Reference-locality</th><th>Northwestern Province</th><th>Southwestern Province</th><th>Hyaenodonts from Northwestern Province</th><th>Hyaenodonts from Southwestern Province</th></tr></thead><tbody><tr><td>Robiacian</td><td>–</td><td>–</td><td>14</td><td>Egerkingen α+β</td><td>Le Guépelle Geiseltal Oberkohle</td><td>Lissieu Issel</td><td>Prodissopsalis eocaenicus; Eurotherium theriodis; Allopterodon torvidus; Proviverra typica; Cynohyaenodon trux; C. ruetimeyeri</td><td>Prodissopsalis eocaenicus; Eurotherium theriodis</td></tr><tr><td rowspan="3">Geiseltalian</td><td rowspan="2">44</td><td rowspan="2">–</td><td rowspan="2">13</td><td rowspan="2">Geiseltal-Obere Mittelkohle</td><td rowspan="2">?Egerkingen y La Défense Bouxwiller</td><td rowspan="2">?Aigues-Vives 2</td><td rowspan="2">Prodissopsalis eocaenicus; Leonhardtina gracilis; Proviverra typica; Cynohyaenodon trux; Allopterodon torvidus*; Oxyaenoides schlosseri*; Eurotherium theriodis*; Alienetherium buxwilleri*; Praecodens acutus*; Cartierodon egerkingensis*</td><td rowspan="2">cf. Oxyaenoides schlosseri*; Proviverra typica*.*; Oxyaenoides schlosseri*; Eurotherium theriodis*</td></tr><tr><td>–</td><td>–</td></tr><tr><td>47-48</td><td>–</td><td>12</td><td>Geiseltal-Untere Mittelkohle</td><td>–</td><td>–</td><td>–</td><td>Prodissopsalis eocaenicus; Leonhardtina gracilis; Proviverra typica; Cynohyaenodon trux</td><td>–</td></tr><tr><td rowspan="3">Grauvian</td><td rowspan="2">–</td><td rowspan="2">–</td><td rowspan="2">11</td><td rowspan="2">Geiseltal-Untere Unterkohle</td><td rowspan="2">Messel</td><td rowspan="2">Palassou</td><td rowspan="2">Oxyaenoides bicuspidens; Matthodon tritens; Eurotherium matthesi; Lesmesodon edingeri*; L. behnkeae*</td><td rowspan="2">Oxyaenoides bicuspidens; Oxyaenoides aumelasiensis n. sp.*; Leonhardtina meridianum n. sp.*; Matthodon peignei n. sp.*</td></tr><tr><td>–</td><td>–</td></tr><tr><td>–</td><td>–</td><td>10b</td><td>–</td><td>–</td><td>–</td><td>–</td><td>–</td><td>Oxyaenoides bicuspidens; Oxyaenoides aumelasiensis n. sp.*; Leonhardtina meridianum n. sp.*; Matthodon peignei n. sp.*</td></tr><tr><td rowspan="3">Neustrian</td><td rowspan="2">–</td><td rowspan="2">PE V</td><td rowspan="2">8+9</td><td rowspan="2">Avenay</td><td rowspan="2">Grauves</td><td rowspan="2">Cuis, Chavot, Mancy, Monthelon</td><td rowspan="2">–</td><td rowspan="2">Oxyaenoides lindgreni*; Matthodon menui; Protoproviverra palaeonictides*; Leonhardtina godinoti</td><td rowspan="2">–</td></tr><tr><td>–</td><td>–</td></tr><tr><td>–</td><td>PE IV</td><td>8+9</td><td>–</td><td>Mutigny</td><td>–</td><td>–</td><td>Minimovellentodon russelli* –</td><td>–</td></tr><tr><td rowspan="3">–</td><td rowspan="3">55.12</td><td rowspan="3">PE III</td><td rowspan="3">8+9</td><td rowspan="3">–</td><td rowspan="3">Abbey Wood</td><td rowspan="3">–</td><td rowspan="3">–</td><td rowspan="3">Prototomus minimus; Prototomus girardoti;</td><td rowspan="3">–</td></tr><tr><td>–</td><td>–</td></tr><tr><td>–</td><td>–</td></tr><tr><td>–</td><td>PE II</td><td>–</td><td>–</td><td>–</td><td>Soissons</td><td>Rians, Palette, Fournes, Fordones</td><td>Prototomus girardoti</td><td>Eoproviverra eisenmanni*; Parvagula palulae*; Galecyon gallus*</td></tr><tr><td>–</td><td>55.8</td><td>PE I</td><td>7</td><td>Dormaal</td><td>Le Quesnoy</td><td>–</td><td>Arfia gingerichi; Prototomus minimus; Prototomus girardoti; Galecyon morloi</td><td>–</td></tr></tbody></table>

possible bone-cracking predator (Solé *et al.* 2014a) might be related to its short stratigraphic distribution, but only further studies on the ecology of the Eocene carnivorous mammals could validate this hypothesis.

The locality of Aumelas has generally been considered close to the MP13 reference-level. This hypothesis is consistent with the range of *Oxyaenoides* and *Leonhardtina*, but conflicts with the range of *Matthodon* (Table 5). The
<figure><img src="image_11.png" /><figcaption>FIG. 5.— Values of the Ln (body mass) of mesonychids, oxyaenids, and hyaenodonts (“Proviverrinae” sensu Solé (2013), “Sinopinae”, “Arfiinae”, Hyaenodontinae, and Hyainailourinae) from MP7 to MP19 with particular attention on hyaenodonts from Aumelas. Aumelas is here represented to be close to the MP11 reference-level. Abbreviations: **ELMA**, European Land Mammal Ages; **EMP**, Mammal Palaeogene. Values are available at Appendix 6.</figcaption></figure>

only reference-levels that could include all three genera are the MP10 and MP11. The genera are known from Paris Basin localities close to the MP10 and around MP11 in the Geiseltal area. *Oxyaenoides* and *Matthodon* are both recorded in Geiseltal-Untere Unterkohle (MP11) (Table 5), but not *Leonhardtina*, which is only known in Geiseltal-Untere Mittelkohle (MP12) and Geiseltal-Obere Mittelkohle (MP11) in the Geiseltal area. Because this genus is known in MP10 localities, it may have been present around MP11 in Geiseltal. It is interesting that only large and middle-sized hyaenodonts are known from Geiseltal-Untere Unterkohle.

The specimens referred to *Oxyaenoides*, *Matthodon*, and *Leonhardtina* from Aumelas are distinctive enough from specimens discovered in the northwestern Europe that we designated three new species at Aumelas. These species each possess relatively primitive characters with respect to other younger species in the genus; however, they present also probable derived features.

The presence of a species of *Matthodon* as well as the primitive features of the new *Matthodon*, *Oxyaenoides*, and *Leonhardtina* species at Aumelas suggests an age close to the MP10 and MP11 reference-levels (maybe closer to MP10 reference-level) rather than the MP13 age previously proposed based on other mammalian groups (Bio-chroM'97 1997).

## THE EVOLUTION OF EUROPEAN HYAENODONTS

The new topology of the phylogenetic tree (close relationship between the Proviverrinae and Hyaenodontidae) supports the hypothesis of Solé *et al.* (2014a) that the hyaenodonts had a radiation in Europe around the Ypresian/Lutetian transition (see Solé *et al.* 2014a for an extensive description of this radiation during the Eocene of Europe). Therefore, it is interesting to look at the evolution of the ecology of the European hyaenodonts.

The body mass of extinct carnivorous mammals is important for understanding their ecological “guard” (*sensu* Root 1967). Reconstructing the body mass of extinct carnivores also facilitates examination of larger evolutionary trends. Following Solé *et al.* (2014a, 2015a), we estimated the body mass of the new hyaenodonts found at Aumelas and compared them to those of other European hyaenodonts in a chronostratigraphic context (Fig. 5). The body masses of the new hyaenodonts fit with the general pattern observed in European hyaenodonts, which shows a body mass increase through the Ypresian and Lutetian (see also Solé & Mennecart 2019).

We specially focused on the hyaenodonts recorded from the MP10, MP11 and MP12 reference-levels (Fig. 6). The body mass of *Oxyaenoides aumelasiensis* n. sp. (4.2 kg) is smaller than its relatives found in the northwestern Europe, as discussed above: 9.7 kg for *O. lindgreni*; 14.2 kg for *O. bicuspidens* and 16.9 kg for *O. schlosseri*. The body mass of *O. bicuspidens* recorded
<figure><img src="image_12.png" /><figcaption>FIG. 6. — Comparison of the range of body masses (in LN) of the hyaenodonts from Aumelas with those recorded from the MP10 (Cuis, Grauves, Monthelon, Mancy), MP11 (Geiseltal-Untere Unterohle) and MP12 (Geiseltal-Untere Mittelkohle) reference-levels in a chronostratigraphic context. Two hypotheses are shown: **Hypothesis 1**, Aumelas is close to MP10 reference-level (Ypresian); **Hypothesis 2**, Aumelas is close to MP11 reference-level (Lutetian). The hypothesis 1 is based on the primitive features displayed by the fossils from Aumelas; the hypothesis 2 is based on the intermediate size of *Matthodon peignei* n. sp. between *Matthodon menui* (MP10) and *Matthodon tritens* (MP11). Note: the range of body masses provided for the MP10 level combines the data available from Grauves (reference-locality), Cuis, Chavot, Mancy, and Monthelon. Authorships of taxa are indicated in the Appendix 8.</figcaption></figure>

at Rouzilhac is 14.1 kg. Because the three other *Oxyaenoides* species have been reconstructed as hypercarnivorous predators (Morlo 1999; Solé *et al.* 2014a, 2015a), and *O. aumelasiensis* n. sp. is part of this genus, it likely was also a hypercarnivorous mammal. However, because of the difference in body mass, *O. aumelasiensis* n. sp. likely had a different ecological role than the three other *Oxyaenoides* species. Compared to the faunas recorded in MP10-MP12 localities, *O. aumelasiensis* n. sp. is close in size to the hyaenodont species *Eurotherium matthesi* recorded in Geiseltal-Untere Unterkohle (MP11) (Fig. 6). *E. matthesi* is reconstructed as a hypercarnivorous mammal (Morlo 1999) with an estimated body mass close to 6.5 kg. Therefore, it seems possible that in southwestern Europe, the genus *Oxyaenoides* filled two distinct ecological niches: the one it occupied in northwestern Europe (i.e., as a large hypercarnivorous predator as evidenced at Rouzilhac), but also the one occupied by *Eurotherium* (i.e., as a fox-sized meat-eating predator).

*Matthodon peignei* n. sp. has a body mass (c. 15 kg) close to that of the MP10 and MP 11 *Matthodon* species recorded in northwestern Europe. *Matthodon* may have been a durophagous hyaenodont, based on the size of its premolars (see Solé *et al.* 2014a, but see Morlo 1999 for a different opinion). Because the new species is close in size and morphology to the northwestern species, these species likely belonged to the same guild: large meat/bone eater – it is important to note that ‘large’ herein is relative to the overall body mass of the carnivorous mammals from the Ypresian and Lutetian and does not correspond to the concept of ‘large’ carnivores in the physiological sense of Carbone *et al.* (2007). Finally, the *Leonhardtina* species (c. 850 g) filled the same ecological niche in northwestern and southwestern Europe (i.e., small meat/ non-vertebrate eater). Therefore, only *O. aumelasiensis* n. sp. is occupying unexpected niche-space among the hyaenodonts from the southwestern and northwestern Europe.

The hyaenodont specimens from Aumelas suggest mammalian predator faunas in northwestern and southwestern

Europe were homogeneous at the genus level around the Ypresian/Lutetian transition. This differs from the biogeographic generic diversity of faunas during the early Ypresian described by Marandat (1997) and Marandat *et al.* (2012) (see Solé *et al.* 2014a, 2015a for a discussion concerning the case of Hyaenodonta).

The overall bodymass disparity of the hyaenodonts is similar between the two European areas (Fig. 6); in contrast, the mesonychids have different body masses between the two European areas (the southern mesonychids have twice the body mass of the northern mesonychids; see Solé *et al.* 2018). Furthermore, the case of *Oxyaenoides* shows that the ecological niches occupied by this genus were probably different between the northwestern and southwestern Europe.

These two observations indicate that the European carnivorous faunas were probably not ecologically identical despite an homogeneity at the genus level. However, it is worth keeping in mind that the evolution of carnivorous communities during the Paleogene of Europe has not been comprehensively analyzed.

## CONCLUSION

Three new carnivorous mammals referred to Hyaenodonta are now known from Aumelas. They provide information on the radiation of the hyaenodonts in Europe around the Ypresian/ Lutetian transition, when specialized ecomorphotypes – like hypercarnivorous *Oxyaenoides* and the durophageous *Matthodon* (Solé *et al.* 2014a) – appeared.

This new material demonstrates that the northwestern and southwestern European faunas were homogenous (i.e., presence of the same genera in the two areas), in contrast to the faunas in the early Ypresian (Marandat 1997; Marandat *et al.* 2012). However, minor differences in the composition of the carnivore fauna (i.e., ecological niches occupied by *Oxyaenoides*) are intriguing and must be investigated in the future.
The presence of these hyaenodonts, especially *Matthodon*, supports a reclassification of Aumelas as close to Ypresian/Lutetian boundary rather than close to the MP13 reference-level (middle Lutetian). This stratigraphic hypothesis requires further research on the rest of the fauna preserved at Aumelas.

## Acknowledgements

We kindly thank L. de Bonis and L. Werdelin for initiating this memorial in tribute to S. Peigné. We would also like to express our warmest regards for Stéphane and his family. We are grateful to our retired colleagues of the ISEM lab that have explored the locality of Aumelas since the 1960s (Hartenberger J.-L., Sudre J., Sigé B.†, Vianey-Liaud M., Crochet J.-Y.) and to our close collaborator Rémy J. We also thank Adnet S., Jiquel S., Charruault A.-L. and the promotion of master 2 of paleontology of the University of Montpellier for the more recent excavation (2008-2009). We thank N. Vallée Gillette (RBINS) for the pictures and preparation of the fossils, and J. Cilis (RBINS) for the SEM pictures. We thank L. Werdelin (Editor), M. Borths, and one anonymous reviewer for their constructive comments that improved this article, as well as the quality of the English of the manuscript. The MNHN gives access to the collections in the framework of the RECOLNAT national Research Infrastructure. This work was supported by project Brain Pionnier BR/175/PI/CARNAGES of the Federal Science Policy Office of Belgium and is part of an ISEM-UM structured program for the revision of mammalian fauna from the Paleogene of Montpellier led by Lihoreau F. and Tabuce R.

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Submitted on 31 December 2018;
accepted on 3 September 2019;
published on 4 June 2020.
Allopterodon torvidus

14: 1 → 0

16: 0 → 1

25: 0 → 1

64: ? → 0

Boritia duffaudi

15: 0 → 1

21: 0 → 1

31: 1 → 0

36: 0 → 1

Cynohyaenodon cayluxi

26: 0 → 1

84: 0 → 1

87: 1 → 2

105: 1 → 0

Cynohyaenodon trux

87: 1 → 2

105: 1 → 0

Eurotherium matthesi

31: 1 → 0

57: 1 → 0

58: 1 → 0

93: 1 → 0

94: 1 → 0

Eurotherium theriodis

7: ? → 1

18: 1 → 0

31: 1 → 0

57: 1 → 0

58: 1 → 0

93: 1 → 0

94: 1 → 0

Galecyon morloi

Characters 19 to 29 are coded '?' because no P₄ is known for this species (see Zack 2011)

Leonhardtina gracilis

21: 0 → 1

43: 1 → 0

Lesmesodon spp.

42: 0 → 1

Matthodon tritens

57: 1 → 0

60: 1 → 0

Molodon vellerei

11: 0 → 1

16: 0 → 1

21: 0 → 1

26: 0 → 1

42: 0 → 1

45: 0 → 1

78: 0 → ?

79: 0 → ?

84: 0 → 1

93: 1 → 0

Oxyaenoides bicuspidens

10: 1 → 0

60: 1 → 0

Oxyaenoides lindgreni

36: 0 → 1

100: X → ?

104: X → ?

107: X → ?

Parvagula palulae

42: ? → 0

Preregidens langebadrae

11: 0 → 1

28: 0 → 2

31: 1 → 0

36: 0 → 1

Proviverra typica

11: 0 → 1

16: 0 → 1

21: 0 → 1

26: 0 → 1

44: 0 → 1

102: 0 → 2

105: 1 → 0

Quercytherium simplicidens

36: 0 → 1

39: 0 → 2

60: 1 → 0

93: 1 → 0

94: 1 → 0

114: 0 → 1

Quercytherium tenebrosum

36: 0 → 1

93: 1 → 0

94: 1 → 0
APPENDIX 2. — Descriptions of characters and character states used in the phylogenetic analysis.

The characters are from Borths & Stevens (2019) without
modification.

LOWER DENTITION

*Deciduous Lower Dentition*

1. dP3 paraconid height: paraconid indistinct (0); paraconid present lower than talonid (1); paraconid present and taller than talonid (2).

2. dP3 protostylid between protoconid and talonid: present (0); absent (1). (New Character relative to Borths & Stevens 2017c).

3. dP4 paraconid height: lower than half protoconid height (0); half protoconid height or taller (1).

4. dP4 metaconid height: lower than half paraconid height (0); half paraconid height or taller (1); metaconid absent (2). (compare with Bastl *et al.* 2014: character 12).

5. dP4 talonid basin cusps: Entoconid and hypoconid present (0); Only hypoconid present (1). (compare with Bastl *et al.* 2014: character 13).

6. dP4 talonid length proportion: More than 1/3 length of entire tooth (0); less than 1/3 length of entire tooth (1).

*Adult Lower Dentition*

7. First mental foramen position: inferior to P1 (0); inferior to P2 (1). (Solé *et al.* 2014b: character 0).

8. Second mental foramen position: inferior to P3 (0); inferior to P4 (1). (Solé *et al.* 2014b: character 1).

9. P1: present (0); absent (1). (Solé *et al.* 2014b: character 2).

10. P1 root number: two roots (0); one root (1). (Polly 1996: character 13; Zack 2011: character 2; Solé *et al.* 2014b: character 2).

11. P2 talonid mesiodistal length: absent to short (0); elongate with distinct inflection separating postprotocristid from talonid (1). (modified Polly 1996: character 3; Egi *et al.* 2005: character 34; Solé *et al.* 2014b: character 5).

12. P2 to P3 relative mesiodistal length: P2 shorter than P3 (0); P2 as long or longer than P3 (1). (Egi *et al.* 2005: character 29; Solé *et al.* 2014b: character 6).

13. P3 inclination: perpendicular to horizontal ramus, tooth forms isosceles triangle in buccal view (0); tooth inclines distally, preprotocristid mesially convex (1).

14. P3 buccolingual width relative to mesiodistal length: width 33% of length (0); width 50% of length (1); width more than 50% of length (2).

15. P3 paraconid morphology: absent or small (0); developed with distinct postparacristid (1). (modified Polly 1996: character 6; Egi *et al.* 2005: character 35; Solé *et al.* 2014b: character 7).

16. P3 talonid mesiodistal length: short, cusp-like (0); long, distinct inflection separating postprotocristid from talonid (1). (modified Polly 1996: character 4; Egi *et al.* 2005: character 37; Solé *et al.* 2014b: character 8).

17. P3 entoconid: absent (0); present (1). (Solé *et al.* 2014b: character 9).

18. P3 to P4 relative mesiodistal length: P3 shorter than P4 (0); P3 as long or longer than P4 (1). (Egi *et al.* 2005: character 31; Solé *et al.* 2014b: character 10).

19. P4 inclination: perpendicular to horizontal ramus, tooth forms isosceles triangle in buccal view (0); tooth inclines distally, preprotocristid mesially convex (1).

20. P4 orientation: talonid and protoconid aligned with horizontal ramus (0); talonid and protoconid at oblique angle to horizontal ramus (1). (New Character relative to Borths & Stevens 2017c).

21. P4 paraconid morphology: present but poorly developed (0); paraconid well-developed (1); paraconid indistinct to absent (2). (modified Polly 1996: character 7; Solé *et al.* 2014b: character 11).

22. P4 metaconid: absent (0); present, usually weakly developed or ridge-like (1). (modified Solé *et al.* 2014b: character 12).

23. P4 entoconid: absent (0); present (1). (Solé *et al.* 2014b: character 14).

24. P4 hypoconulid: present and distinct (0); absent or indistinct from entoconid (1). (New Character relative to Borths & Stevens 2017c).

25. P4 hypoconid height: short, less than 33% of protoco- nid height (0); tall, more than 33% protoconid height (1). (modified Solé *et al.* 2014b: character 16).

26. P4 talonid basin: buccolingually compressed and shallow (0); buccolingually wide and deep (1); absent (2).

27. P4 precingulid and postcingulid: absent (0); present (1). (modified Solé *et al.* 2014b: character 18).

28. P4 relative height: mesiodistally longer than height (0); mesiodistally shorter than height (1); mesiodistal length and height subequal (2).

29. P4 height relative to molars: shorter than all molars (0); taller than M1 only (1); taller than M2 (2).

30. P5 presence: present (0); absent (1).

31. M1 and M2 entoconid morphology: well developed or bulbous (0); crestiform with visible apex (1); undiffer- entiated entocristid (2). (modified Zack 2011: character 23; Solé *et al.* 2014b: character 27).

32. M1 and M2 talonid depth: deep (0); shallow (1). (modified Zack 2011: character 25; Solé *et al.* 2014b: character 29).

33. M2 entocristid in lingual view: parallels hypocristid (0); present, stops before metaconid (lower than hypocristid) (1); weak ridge or absent (2).

34. M3 entocristid: parallels hypocristid (0); present, stops before metaconid (lower than hypocristid) (1); weak ridge or absent (2).

35. M1 and M2 talonid buccolingual width: narrow, less than 80% width of trigonid (0); wide, greater than 80% trigonid (1). (modified Solé *et al.* 2014b: character 28).

36. M1 mesiodistal length relative to M2: M1 length subequal or longer than M2 (0); M1 length less than M2 (1). (modified Zack 2011: character 26; Solé *et al.* 2014b: character 31).
37. M1 relative to M2 mesiodistal length: M1 greater than or equal to M2 trigonid length (0); M1 greater than M2 paraconid length (1); M1 equal to or shorter than M2 paraconid length (2). (New Character relative to Borths & Stevens 2017c).

38. M1–M3 trigonid height relative to talonid: trigonid tall on all molars, talonid less than 50% of trigonid height (0); trigonid low on all molars, talonid more than 50% of trigonid height (1); trigonid low on M1 and M2 (2). (modified Solé *et al.* 2014b: character 32).

39. M3 postprotocristid distal trend in buccal view: slopes mesial to distal (0); perpendicular to alveolus (1); slopes distal to mesial (overhangs talonid) (2).

40. M2 cristid obliqua orientation relative to mesiodistal axis: lingual to buccal trend (0); parallel to mesiodistal axis (1); buccal to lingual trend (2). (compare to Zack 2011: character 21).

41. M2 and M3 paraconid position relative to protoconid, angle defined relative to mesiodistal axis of mandible: directly mesial to protoconid, 15 degrees (0); slightly lingual paraconid, 15.1 to 45 degree angle (1); strong lingual position, 45.1 to 60 degrees (2). *Ordered.*

42. M3 postparacristid mesial to distal trend: steep slope to preprotocristid ("V" shaped acute angle) (0); shallow slope to preprotocristid (forms right angle with preprotocristid) (1); forms obtuse angle with preprotocristid (2). *Ordered.*

43. M2 and M3 paraconid height relative to protoconid: paraconid significantly shorter than protoconid (0); paraconid slightly shorter than protoconid (1); paraconid and protoconid subequal in height (2).

44. M3 talonid buccolingual width in lingual view: postparacristid shorter than premetacristid (0); postparacristid subequal to premetacristid (1); postparacristid longer than premetacristid (2).

45. M3 postparacristid length to preprotocristid in buccal view (carnassial blade proportions): postparacristid much shorter than preprotocristid (30%) (0); postparacristid half length of preprotocristid (1); postparacristid more than half preprotocristid length (2); subequal lengths (3).

46. M2 and M3 metaconid expression: connate and connects to paraconid base (0); connate, separated from paraconid (1); fold or ridge (2); absent (3). *Ordered.*

47. M1 metaconid: taller than paraconid (0); subequal to paraconid (1); shorter than paraconid or absent (2). *Ordered* (compare to Polly 1996: character 18).

48. M2 metaconid: taller than paraconid (0); subequal to paraconid (1); shorter than paraconid or absent (2). *Ordered* (compare to Polly 1996: character 18).

49. M3 metaconid: taller than paraconid (0); subequal to paraconid (1); shorter than paraconid or absent (2). *Ordered* (compare to Polly 1996: character 19).

50. M2 mesiodistal length to M3 length: M2 shorter than M3 (0); M2 subequal to M3 (1); M2 longer than M3

(2); M3 absent (3). *Ordered* (compare to Zack 2011: character 30).

51. M2 talonid mesiodistal length (% of total mesiodistal length): >40% (0); 40% to 30% (1); 29% to 21% (2); <20% (3). *Ordered.*

52. M3 talonid mesiodistal length (% of total mesiodistal length): >40% (0); 40% to 30% (1); 29% to 21% (2); <20% (3). *Ordered.*

53. M3 talonid: present, bears hypoconid and hypoconulid (0); present, only one distinct cusp (1); absent (2). *Ordered.*

54. M2 buccal talonid margin: steep slope distal to mesial (0); shallow slope distal to mesial (1); parallel to alveolus (2); slopes mesial to distal (3). *Ordered.*

55. M3 buccal talonid margin angle from highest point to lowest: steep slope distal to mesial (0); shallow slope distal to mesial (1); parallel to alveolus (2); slopes mesial to distal (3). *Ordered.*

56. M1–M3 ectocingulid: weakly expressed to absent (0); distinct (1). (modified Solé *et al.* 2014b: character 34)

57. M1–M3 postcingulid: absent (0); present (1). (modified Solé *et al.* 2014b: character 35).

58. M1–M3 ectocingulid to postcingulid connection: separated (0); fused (1). (modified Solé *et al.* 2014b: character 36).

59. M3 talonid buccolingual width relative to M2 talonid width: equal (0); narrower (1).

60. Mandible inflection anterior to angular process (Solé *et al.* 2015b): present (0); absent (1).

61. Angular process morphology: distinct process with medial inflection (0); gently curved process in line with mandibular corpus (1); ventral inflection (2).

62. Mandibular condyle position: superior to M3 alveolus (0); directly distal to M3 alveolus (1); inferior to M3 alveolus (2).

63. Coronoid process shape: tall, anterior and posterior slopes similar (0); tall, posterior slope concave (1); low, rounded (2).

64. Anterior coronoid angle relative to horizontal ramus: near vertical, 90 to 100 degrees (0); slight posterior inclination, 100 to 110 degrees (1); strong posterior inclination, greater than 110 degrees (2).

65. Masseteric fossa depth: deeply excavated with strong anterior angle, inferior margin welldefined (0); rounded anterior margin, little inferior definition (1); deep fossa but poorly defined inferior margin (2).

## UPPER DENTITION

### *Deciduous Upper Dentition*

66. dP3 parastyle mesiodistal length: more than half meta-style length (0); less than half metastyle length (1).

67. dP3 metacone-paracone fusion: metacone distinct cusp (premetacrista slopes to metacone apex; See *Pterodon dasyuroides*) (0); metacone fused to paracone (premetacrista subhorizontal; See *Apterodon*) (1).
68. dP3 paracone morphology: pre- and postparacrista similar in slope (apex isosceles triangle in buccal view) (0); preparacrista distally inclined (apex closer to right triangle in buccal view) (1).

69. dP3 metastyle notch: Small inflection between postmetacrista and metastyle (0); deep notch between postmetacrista and metastyle (1).

70. dP3 Protocone prominence: mesiodistal length shorter than buccolingual width (narrow) (0); mesiodistal length equal to or longer than buccolingual width (wide) (1). (modified from Bastl *et al.* 2014: character 2).

71. dP3 lingual cingulum: present (distinct lingual connection between parastyle and protocone) (0); absent (faint or no connection between parastyle and protocone) (1). (modified from Bastl *et al.* 2014: character 4).

72. dP4 main cusp height: paracone taller than metacone (0); Paracone subequal to metacone (1); Paracone shorter than metacone (2). *Ordered* (Modified from Bastl *et al.* 2014: character 6).

73. dP4 protocone orientation: Protocone projects mesially to parastyle margin (0); Protocone projects lingually and does not align with parastyle (1).

74. dP4 ectoflexus: Deep and distinct inflection between metacone and metastyle (0); Indistinct or shallow inflection between metacone and metastyle (1).

*Adult Upper Dentition*

75. Upper incisor count: 3 or more (0); 2 or fewer (1).

76. Lateral-most upper incisor: incisiform, similar to mesial incisor (0); caniniform (1).

77. Upper canine alveolar cross-section: distally tapers with mesiodistal axis more than 20% buccolingual axis (0); rounded with mesiodistal axis less than 20% buccolingual axis (1). (New Character relative to Borths & Stevens 2017c).

78. P3 lobe of the protocone or protocone alveolus: absent (0); present but small and alveolus merged lingual alveoli (1); protocone well-developed, individuated and alveolus distinct from lingual alveoli (2). (modified Polly 1996: character 9; Egi *et al.* 2005: character 4; Solé *et al.* 2014b: character 39; modified relative to Borths & Stevens 2017c).

79. P3 root number: two roots (0); three roots (1). (Solé *et al.* 2014b: character 40).

80. P3 contact with P4 parastyle: P3 contacts or aligned with P4 parastyle (0); P3 framed by P4 parastyle (1).

81. P4 parastyle: distinct (0); very reduced to absent (1). (modified Egi *et al.* 2005: character 8; Zack 2011: character 34; Solé *et al.* 2014b: character 41).

82. P4 protocone alignment: transversely aligned with paracone (0); mesially shifted relative to paracone (1). (Zack 2011: character 36; Solé *et al.* 2014b: character 42).

83. P4 protocone morphology: bulbous and distinct from paracone (0); weak separation from paracone, shelf to cingulum-like (1). (modified Polly 1996: character 10; Egi *et al.* 2005: character 6/7; Solé *et al.* 2014b: character 43).

84. P4 metastylar blade (=postmetacrista): short (0); elongate (1). (modified Egi *et al.* 2005: character 9; Zack 2011: character 35; Solé *et al.* 2014b: character 45).

85. P4 metastyle contact with M1: P4 metastyle braced bucally by M1 parastyle (0); P4 metastyle contacts mesial aspect of M1 parastyle (1).

86. M1 and M2 metastyle blade curvature: straight with carnassial notch (0); postmetacrista arcuate, no carnassial notch (1). (Zack 2011: character 42; Solé *et al.* 2014b: character 46).

87. M1 and M2 metastyle blade length: short, carnassial blade shorter than postmetacrista (0); intermediate, subequal to slightly longer than postmetacrista (1); elongate, greater than 1.5xlength of postmetacrista (2). *Ordered* (Egi *et al.* 2005: character 21; Solé *et al.* 2014b: character 47).

88. M1 and M2 metastyle robusticity (length divided by width): robust or 1.0 or greater (0); moderate or between 0.71 and 0.99 (1); gracile or 0.70 or less (2). (New Character relative to Borths & Stevens 2017c).

89. M1 postmetacrista to metastyle angle: Angle less than 130 degrees (0); Angle between 131 and 144 degrees (1); Angle greater than 145 degrees (2).

90. M1 mesiodistal length relative to M2: M1 subequal or longer than M2 (0); M1 shorter than M2 (1). (modified Solé *et al.* 2014b: character 50).

91. M1 and M2 premetaconule crista: present (0); absent (1). (modified Solé *et al.* 2014b: character 51).

92. M1 and M2 conules: metaconule and paraconule present (0); only paraconule present (1); metaconule and paraconule absent (2) (modified Solé *et al.* 2014b: character 53).

93. M1 and M2 precingulum and postcingulum: absent (0); present (1). (modified from Polly 1996: character 20; Egi *et al.* 2005: character 26; Solé *et al.* 2014b: character 54).

94. M1 and M2 precingulum and postcingulum connection: separated (0); fused lingually (1). (Solé *et al.* 2014b: character 55).

95. M1 and M2 protocone morphology: Triangular, mesial and distal margins angled (0); parallel mesial and distal margins (1). (modified Egi *et al.* 2005: character 14).

96. M1 protocone position relative to paracone and metacone: centered (0); directly lingual to the parastyle (1); mesial to the parastyle (2). *Ordered* (modified Egi *et al.* 2005: character 15; Solé *et al.* 2014b: character 56; State 2 added to Borths & Stevens 2017c).

97. M1 protocone lingual projection, measured as the ratio of the protocone buccolingual length divided by the length of the bases of the paracone and metacone: long, ratio greater than 0.9 (0); intermediate, ratio between 0.9 and 0.61 (1); short, ratio less than 0.6 (2).

98. M1 and M2 metacone mesiodistal length relative to paracone: shorter (0); subequal (1); longer (2). (modified Egi *et al.* 2005: character 12).

99. M1 and M2 metacone coronal cross section: circular (0); ovoid, slightly compressed buccolingually (1); ellipsoid, strong buccolingual compression (2).
100. M2 paracone compression: conical, rounded base (0); ovoid (slight buccolingual compression) (1); ellipsoid (strong buccolingual compression) (2).

101. M1 and M2 paracone and metacone separation: separated to base (0); fused between base and half of height (1); almost completely fused (2). *Ordered* (modified Polly 1996: character 28; Egi *et al.* 2005: character 10; Zack 2011: character 40).

102. M1 and M2 paracone height: paracone taller than metacone (0); paracone as tall as metacone (1); paracone shorter than metacone (2). *Ordered* (modified Egi *et al.* 2005: character 11; Zack 2011: character 41; Solé *et al.* 2014b: character 48).

103. M1 and M2 protocone height: shorter than paracone/ metacone separation (0); same height as paracone/meta- cone separation (1); subequal to paracone/metacone apices (2). (compare to Zack 2011: character 49).

104. M2 parastyle: absent (0); present and shelf-like (1); pre- sent, single distinct cusp (2); present multiple distinct cusps (3). (modified from Egi *et al.* 2005: character 19 and Borths & Stevens 2017c).

105. M1 parastyle compared to M2: M1 parastyle relatively shorter than M2 parastyle (0); parastyle on M1 and M2 similar (1).

106. M1 and M2 buccal cingulum: absent (0); weak ridge along metastyle base (1); prominent shelf forms shallow basin between metastyle and cingulum (2). (modified Egi *et al.* 2005: characters 16/17).

107. M2 ectoflexus: strong, strong indentation (0); weak, slight curve (1); absent, straight (2). (modified Egi *et al.* 2005: character 18).

108. Ectoflexus depth on M1 and M2: M1 ectoflexus shal- lower than M2 (0); same relative depth between M1 and M2 (1); M2 lacks metastyle (no ectoflexus) (2). *Ordered* (compare to Zack 2011: character 52).

109. M3 metacone: present (0); absent (1). (modified Polly 1996: character 15; Zack 2011: character 53; Solé *et al.* 2014b: character 58).

*Cranial characters*

110. Nuchal crest morphology: medial to lateral trend from apex to mastoid (0); dorsolateral margin tapers medially with thin connection to exoccipital (1); lateral margins trend medially, very weak ridge connects to exoccipital (2). *Ordered* (modified Polly 1996: character 35).

111. Facial wing of the lacrimal: extensive (larger than orbit diameter) (0); moderate (slightly longer than orbit diameter) (1); reduced (shorter than orbit diameter) (2). (modified Polly 1996: character 36).

112. Foramen rotundum size: slightly larger than foramen ovale (0); much larger than foramen ovale (1). (Polly 1996: character 39).

113. Palatal rugosity or torus at distal margin of palate: well- expressed (0); smooth (1).

114. Zygomatic arch contact: short contact between zygo- matic and squamosal (0); extensive contact between zygomatic and squamosal (1).

115. Superior squamosal morphology: superior and inferior margins parallel (0); torsion along superior margin (1).

116. Foramen ovale orientation: anterior orientation (0); palatal orientation (1).

117. Exoccipital condyle position: tall, lateral placement around foramen magnum (0); ventral placement around foramen magnum (1).

118. Notch between occipital condyles: ring-like with no rostral excavation (0); rounded indentation with condyles meeting medially below foramen magnum (1); deep excavation with occipital "processes" following notch (2).

119. Postmandibular process: vertical orientation (0); strong anterior curvature (1).

120. Posterior orbital process: present, strong expression (0); present, weak with frontal "peaked" (1); absent (2).

121. Frontal furrow: absent or indistinct (0); present and well-defined (1).

122. Palatine and pterygoid medial contact: parallel posterior to palatine torus (0); palatines trend medially or partially fuse (1); fused entire extent of palatines (2).

123. Pterygoid size: broad, ventral projection anteriorly extensive (0); short anterior extent, trends medially (1).

124. Frontoparietal suture in dorsal view: steep constriction (0); gentle curvature (1).

125. Lateral expansion of the mastoid process (Solé *et al.* 2015b): projects to midpoint of mandibular fossa (0); projects beyond mandibular fossa (1).

126. Mastoid/paroccipital process: short tubercle (0); well-defined, prong-like process (1).

127. Process at maxilla/jugal suture (Solé *et al.* 2015b): present (0); absent (1).

128. Squamosal constriction in dorsal view: present, squamosal does not extend laterally (0); absent, squamosal expanded laterally (1).

129. Squamosal ventral projection: same transverse plane as petrosal (0); ventral to petrosal (1).

130. Posterior braincase: broad lateral expansion (0); narrow (1).

131. Subarcuate fossa morphology (Polly 1996): cup-shaped (see *Pterodon*) (0); shallow and horseshoe-shaped (see *Hyaenodon*) (1).

132. Bridge of the stylomastoid foramen primitivum: absent or slender (0); robust (1); roofed over with secondary stylomastoid foramen (2). (Polly 1996: character 40)

133. Mastoid sinus lateral to foramen stylomastoid primitivum: absent (0); present (1). (Polly 1996: character 41).

134. Ridge of bone dividing posterior petrosal sinus from foramen stylomastoid primitivum (Polly 1996: present (0); reduced to absent (1). (Polly 1996: character 42)

135. Posterior petrosal sinus: absent (0); small (1); greatly inflated (2). (Polly 1996; character 43).
## POSTCRANIA

### *Humerus*

136. Humerus cross-section above brachial flange: triangular (0); rounded (1). (Polly 1996: character 44).

137. Brachial flange expression: medium (0); enlarged (1); reduced (2). (Polly 1996: character 47).

138. Entepicondylar foramen: present, rounded (0); present, elongate (1); absent (2).

139. Medial epicondyle: large (bulbous) (0); reduced (elongate) (1).

140. Capitulum morphology: rounded, clearly separated (0); cylindrical (1).

141. Greater tubercle of the humerus: prominent, taller than humeral head (0); subequal to height of humeral head (1).

### *Ulna*

142. Proximal trochlear notch orientation: lateral position on shaft (0); medial position on shaft (1).

143. Radial notch orientation: curved and faces laterally (0); flattened and faces anteriorly (1). (Polly 1996: character 47).

144. Olecranon process length: longer than trochlear notch (0); subequal or shorter than trochlear notch (1).

145. Olecranon process orientation: projects medially (0); projects ventrally (1).

### *Femur*

146. Third trochanter of femur: large (0); small (1). (modified Polly 1996: character 50).

### *Astragalus*

147. Astragalar foramen: large (0); reduced (0). (Polly 1996: character 51).

148. Astragalar condyles divided by: shallow depression (0); well-defined fossa or groove (1).

149. Astragalar condyle orientation: oblique relative to astragalar neck (0); parallel to astragalar neck (1).

150. Astragalar head relative to condyles: horizontal orientation (0); slight vertical orientation (1). (Polly 1996: character 53).

151. Sustentacular facet connection to astraglar head: clearly separated from astragalar head (0); grades into articulation of astragalar head (1).

152. Sustentacular facet position on astragalar neck: plantar astragalar neck (0); medial astragalar neck (1).

### *Calcaneum*

153. Peroneal tubercle: distinct and separated from cuboid facet (0); part of a flange that grades to cuboid facet (1).

154. Cuboid facet inclination: perpendicular to calcaneal neck (0); plantar inclination (1).

155. Calcaneal neck trend: dorsal and plantar margins parallel (0); tapers proximally to calcaneal tuberosity (1).

156. Astragalar facet angle: oblique orientation to calcaneal neck (0); parallels calcaneal neck (1). (modified Polly 1996: character 58).
### Matthodon menui

Formation: Sables à Unios et Térédines sensu stricto;

Locality: Chavot, Monthelon, Cuis, Mancy and Grauves;

Geological Age: MP10;

Absolute Age: 51-48.6 Ma;

Country: France;

Citation: Solé *et al.* 2014a;

Specimens observed: MNHN.F.1939-586 (Holotype), left mandible with M₃ and roots of M₂ and distal root of M₁; MNHN.F.Cui14838, LP₄; MNHN.FCHO14799, right mandible with canine, P₂ to M₂, and alveolus of P₁; T.S. 914, right maxillary with P³ and P⁴; T.S. 374, right mandible with complete P₂-P₄, and M₃, fragmentary M₁ and M₂, and alveolus of P₁; MNHN.F.Ma14832, RP₄; MNHN.F.L-68-GR, left mandible with M₁ and alveoli of P₂ to P₄ and M₂; MNHN.F.L-58-Ma, RM².

### Paracynohyaenodon magnus

Formation: Quercy;

Locality: Robiac and Le Bretou;

Geological Age: MP16;

Absolute Age: 51-48.6 Ma;

Country: France;

Specimens observed: BRT 939 (Holotype), right maxillary bearing P⁴ to M²; RBN 5434, maxillaries bearing right P¹ to P⁴, and alveoli of the canines and left P¹ to P³; Rbn-NM 1018, right P⁴; right mandible bearing M₁ to M₃; LAV 1310, left maxillary bearing P³ to M²; LAV 1312, left maxillary bearing M² and M³; BRT 936, right mandible bearing P₂ to M₂; RBN 5434, right mandible bearing the canine, and P₁ to P₄ and left mandible bearing M₂ and M₃; LAV 1314, right mandible bearing P₄ to M₂;

Citation: Lange-Badré 1979; Crochet 1991.

### Paracynohyaenodon schlosseri

Formation: Quercy;

Locality: Robiac, Le Bretou, Lavergne, La Bouffie, Les Pradigues, Les Clapiès;

Geological Age: MP16-MP17a;

Absolute Age: 40-33.9 Ma;

Country: France;

Specimens observed: Qu B 393 (Holotype), right mandible bearing M₁ to M₃; LAV 1310, left maxillary bearing P³ to M²; LAV 1312, left maxillary bearing M² and M³; Ma Pq 60, right mandible bearing P₁ to M₃; MNHN.FQU8539, right mandible bearing P₄ and M₁; PLV 117, left mandible bearing P₃ to M₂;

Citation: Lange-Badré 1979; Crochet 1991.

### Matthodon peignei n. sp.

Formation: Lacustrine limestone of Montpellier, proposed as Lutetian in age in previous study but here questioned as late Ypresian/early Lutetian in age;

Locality: Aumelas, Hérault, France;

Geological Age: MP10-MP11?

Absolute Age: 51-40.4 Ma;

Country: France;

Specimens observed: UM-AUM694, left mandible bearing the p2-m3 and the alveoli of the canine and p1; UM-AUM268, left mandible bearing the m2 and m3.

### Oxyaenoides aumelasiensis n. sp.

Formation: Lacustrine limestone of Montpellier, proposed as Lutetian in age in previous study but here questioned as late Ypresian/early Lutetian in age;

Locality: Aumelas, Hérault, France;

Geological Age: MP10-MP11?

Absolute Age: 51-40.4 Ma;

Country: France;

Specimens observed: UM-AUM693, left mandible bearing m3, the anterior root and talonid of the m1, and the posterior alveolus of the p4, and the alveoli of m2; UM-AUM686, right M2; UM-AUM692, combination of a left maxillary fragment bearing the P2-M1, a right maxillary fragment bearing the P3-P4, the protocone of the M1 and the alveoli of the P2; UM-AUM695, labial part of a M1.

### Leonhardtina meridianum n. sp.

Formation: Lacustrine limestone of Montpellier, proposed as Lutetian in age in previous study but here questioned as late Ypresian/early Lutetian in age;

Locality: Aumelas, Hérault, France; Rouzilhac, Aude, France;

Geological Age: MP10-MP11?

Absolute Age: 51-40.4 Ma;

Country: France;

Specimens observed: UM-AUM54, right mandible bearing p3 to m3; UM-AUM530, right maxillary fragment bearing M2 and M3; MNHN.F.RZ142, right mandible bearing p2-p4 and m3.

### Prodissopsalis eocaenicus

Formation: Karst fillings in an aberrant siderolitic facies

Locality: Geiseltal-Untere-Mittelkohle, Geiseltal-Obere Mittelkohle, Egerkingen γ;

Geological Age: MP12-MP13? (possibly MP14);

Absolute Age: 48.6-40.4 Ma;

Countries: Germany, Switzerland;

Specimens observed: Leo III-394; skull bearing the right P1-P4 and M1-M3, left P3-P4 and M1-M3, and the alveoli of the I2, I3, and canines; XXXV-327, left mandible bearing canine, p3-p4 and m1-m3; XXXVI-255, right mandible bearing p3-p4 and m1-m3.

Citation: Van Valen 1965; Lange-Badré & Haubold 1990.
<figure><img src="image_13.png" /></figure>

APPENDIX 4. — Stratigraphic range (MP levels) of the European hyaenodonts and numerical ages for the MP levels.

RANGE (MP LEVEL) OF THE EUROPEAN
HYAENODONTS

<table><tr><td>Allopterodon torvidus</td><td>MP13-MP14</td></tr><tr><td>Apterodon gaudryi</td><td>Rupelian</td></tr><tr><td>Arfia gingerichi</td><td>MP7</td></tr><tr><td>Boritia duffaudi</td><td>MP8+9</td></tr><tr><td>Cynohyaenodon cayluxi</td><td>MP16-MP17a</td></tr><tr><td>Cynohyaenodon trux</td><td>MP12-MP14</td></tr><tr><td>Eoproviverra eisenmanni</td><td>MP7</td></tr><tr><td>Eurotherium matthesi</td><td>MP11</td></tr><tr><td>Eurotherium theriodis</td><td>MP13-MP14</td></tr><tr><td>Galecyon morloi</td><td>MP7</td></tr><tr><td>Hyaenodon exiguus</td><td>MP22-MP30</td></tr><tr><td>Hyaenodon minor</td><td>MP17a-MP18</td></tr><tr><td>Kerberos langebadreae</td><td>MP16</td></tr><tr><td>Leonhardtina gracilis</td><td>MP12-MP13</td></tr><tr><td>Leonhardtina meridianum n. sp.</td><td>MP10-MP11</td></tr><tr><td>Lesmesodon spp.</td><td>MP11</td></tr><tr><td>Matthodon tritens</td><td>MP11</td></tr><tr><td>Matthodon menui</td><td>MP10</td></tr><tr><td>Matthodon peignei n. sp.</td><td>MP10-MP11</td></tr><tr><td>Molodon vellerei</td><td>MP8+9-MP10</td></tr><tr><td>Oxyaenoides aumelasiensis n. sp.</td><td>MP10-MP11</td></tr><tr><td>Oxyaenoides bicuspidens</td><td>MP11</td></tr><tr><td>Oxyaenoides lindgreni</td><td>MP10</td></tr><tr><td>Paracynohyaenodon spp.</td><td>MP16-MP17a</td></tr><tr><td>Paroxyaena spp.</td><td>MP16-MP17b</td></tr><tr><td>Parvagula palulae</td><td>MP7</td></tr><tr><td>Preregidens langebadrae</td><td>MP8+9</td></tr><tr><td>Prodissopsis eocaenicus</td><td>MP12-MP14</td></tr><tr><td>Prototomus minimus</td><td>MP7-MP8+9</td></tr><tr><td>Proviverra typica</td><td>MP12-MP14</td></tr><tr><td>Pterodon dasyuroides</td><td>MP18-MP20</td></tr><tr><td>Quercytherium simplicidens</td><td>MP17a-MP18</td></tr><tr><td>Quercytherium tenebrosum</td><td>MP16-MP17a</td></tr><tr><td>Thereutherium thylacodes</td><td>MP23-MP26</td></tr></table>

NUMERICAL AGES FOR EACH MP LEVEL

Numerical ages were extracted from Escarguel *et al.* (1997) and Vandenberghe *et al.* (2012).

<table><thead><tr><th>MP</th><th>Start</th><th>End</th><th>ICS stages</th></tr></thead><tbody><tr><td>30</td><td>23.2</td><td>23.03</td><td>Chattian</td></tr><tr><td>29</td><td>23.9</td><td>23.2</td><td>Chattian</td></tr><tr><td>28</td><td>24.6</td><td>23.9</td><td>Chattian</td></tr><tr><td>27</td><td>25.4</td><td>24.6</td><td>Chattian</td></tr><tr><td>26</td><td>27.3</td><td>25.4</td><td>Chattian</td></tr><tr><td>25</td><td>28.8</td><td>27.3</td><td>Chattian-Rupelian</td></tr><tr><td>24</td><td>30</td><td>28.8</td><td>Lutetian</td></tr><tr><td>23</td><td>30.9</td><td>30</td><td>Rupelian</td></tr><tr><td>22</td><td>32.5</td><td>30.9</td><td>Rupelian</td></tr><tr><td>21</td><td>33.9</td><td>32.5</td><td>Priabonian</td></tr><tr><td>20</td><td>34</td><td>33.9</td><td>Priabonian</td></tr><tr><td>19</td><td>35</td><td>34</td><td>Priabonian</td></tr><tr><td>18</td><td>37</td><td>35</td><td>Priabonian</td></tr><tr><td>17b</td><td>37.5</td><td>37</td><td>Priabonian</td></tr><tr><td>17a</td><td>37.8</td><td>37.5</td><td>Priabonian</td></tr><tr><td>16</td><td>40</td><td>37.8</td><td>Bartonian</td></tr><tr><td>15</td><td>41.2</td><td>40</td><td>Lutetian</td></tr><tr><td>14</td><td>43.5</td><td>41.2</td><td>Lutetian</td></tr><tr><td>13</td><td>44.9</td><td>43.5</td><td>Lutetian</td></tr><tr><td>12</td><td>46.3</td><td>44.9</td><td>Lutetian</td></tr><tr><td>11</td><td>47.8</td><td>46.3</td><td>Lutetian</td></tr><tr><td>10</td><td>50.8</td><td>47.8</td><td>Ypresian</td></tr><tr><td>8+9</td><td>55.2</td><td>50.8</td><td>Ypresian</td></tr><tr><td>7</td><td>56</td><td>55.2</td><td>Ypresian</td></tr><tr><td>6</td><td>57.2</td><td>56</td><td>Thanetian</td></tr></tbody></table>
http://sciencepress.mnhn.fr/sites/default/files/documents/en/g2020v42a13_appendix_5.txt

This file is the input code used to perform the analysis that includes 19 ordered characters (listed after "ctype ordered" near the end of the file). It also lists all parameters input into MrBayes, including clock rate, burn-in period, and number of repli-cations. The file can be used directly to run the analysis in MrBayes. With the MrBayes window open, type in the command "execute", type a quotation ("), then drag the tip-dating analysis file, close with a second quotation mark (""); you should see "MrBayes > execute" location_on_ the_computer_with_file_name" on the MrBayes window. Then push the "enter" button to perform the analysis.

http://sciencepress.mnhn.fr/sites/default/files/documents/en/g2020v42a13_appendix_6.txt

The following method follows the one proposed by Borths & Stevens (2017c: supplementary file 3).

This file contains the tree and all summary tree statistics output as part of the summary tree file inferred from the analysis
performed in MrBayes. You can open the file directly in the tree-viewing software FigTree to open the tree.

To adjust the age estimates to an absolute timescale rather than an internal hyaenodont timescale, subtract the age (mean or median) listed for the fixed taxon (*Pakakali*) from 25 (the fixed age used for *Pakakali*) and add the result to each age. For example, to calculate the median date of the divergence of *Prionogale* from *Namasector*: calculate the difference between the fixed age of *Pakakali* (25 Ma) from the age of *Pakakali* on the tree (13.61 Ma) which is 11.39 Ma. Add 11.39 Ma to the age of the node that unites Prionogalidae (12.82 + 11.39) to get the estimated, absolute divergence date between *Prionogale* and *Namasector* (24.21 Ma).

To calculate evolutionary rate of change select "ratelgrBrlens_mean" or "ratelgrBrlensmedian." These display the relative
rate of evolutionary change for the tree. To convert the relative evolutionary rate into absolute evolutionary rate (% change/
site/Ma), multiply the "ratelgrBrlens" value by the mean or median clockrate ouput in the .pstat file from MrBayes. In the
case of the ordered analysis, the median clockrate is 0.007547 and the mean clockrate is 0.01022. Multiply the "ratelgrBrlens"
by the clockrate value then multiply by 100 to express absolute % change/site/Ma. For example, to calculate the median %
change/site/Ma for the branch supporting Prionogalidae: 1.14 × 0.007547 × 100 = 0.86 % change/site/Ma.
<figure><img src="image_14.png" /><figcaption>APPENDIX 7. — Estimations of the body mass of selected hyaenodonts, oxyaenids, and mesonychids.</figcaption></figure>

<table><thead><tr><th>Taxa 1</th><th>Taxa 3</th><th>Species</th><th>Bodymass (kg)</th><th>Bodymass (g)</th><th>Ln bodymass</th><th>FAD mp</th><th>LAD mp</th></tr></thead><tbody><tr><td>Hyaenodonta</td><td>Proviverrinae</td><td>Alienetherium buxwilleri</td><td>5.5</td><td>5500</td><td>8.612503371</td><td>13</td><td>13</td></tr><tr><td>Hyaenodonta</td><td>Proviverrinae</td><td>Allopterodon bulbosus</td><td>0.73</td><td>730</td><td>6.593044534</td><td>17b</td><td>19</td></tr><tr><td>Hyaenodonta</td><td>Proviverrinae</td><td>Allopterodon minor</td><td>2.1</td><td>2100</td><td>7.649692624</td><td>16</td><td>19</td></tr><tr><td>Hyaenodonta</td><td>Proviverrinae</td><td>Allopterodon torvidus</td><td>0.58</td><td>580</td><td>6.363028104</td><td>13</td><td>14</td></tr><tr><td>Hyaenodonta</td><td>Arfiinae</td><td>Arfia gingerichi</td><td>1.63</td><td>1630</td><td>7.396335294</td><td>7</td><td>7</td></tr><tr><td>Hyaenodonta</td><td>Proviverrinae</td><td>Boritia duffaudi</td><td>2.58</td><td>2580</td><td>7.855544678</td><td>8</td><td>9</td></tr><tr><td>Hyaenodonta</td><td>Proviverrinae</td><td>Cartierodon egerkingensis</td><td>28.65</td><td>28650</td><td>10.26290872</td><td>13</td><td>13</td></tr><tr><td>Hyaenodonta</td><td>Proviverrinae</td><td>Cynohyaenodon blainvillei</td><td>0.44</td><td>440</td><td>6.086774727</td><td>19</td><td>19</td></tr><tr><td>Hyaenodonta</td><td>Proviverrinae</td><td>Cynohyaenodon cayluxi</td><td>1.73</td><td>1730</td><td>7.455876687</td><td>16</td><td>17a</td></tr><tr><td>Hyaenodonta</td><td>Proviverrinae</td><td>Cynohyaenodon lautricensis</td><td>3</td><td>3000</td><td>8.006367568</td><td>16</td><td>16</td></tr><tr><td>Hyaenodonta</td><td>Proviverrinae</td><td>Cynohyaenodon rutimeyeri</td><td>1.89</td><td>1890</td><td>7.544332108</td><td>14</td><td>14</td></tr><tr><td>Hyaenodonta</td><td>Proviverrinae</td><td>Cynohyaenodon trux</td><td>1.17</td><td>1170</td><td>7.064759028</td><td>12</td><td>14</td></tr><tr><td>Mesonychia</td><td>Mesonychidae</td><td>Dissacus blayaci</td><td>39.08</td><td>39080</td><td>10.57336611</td><td>10</td><td>10</td></tr><tr><td>Mesonychia</td><td>Mesonychidae</td><td>Dissacus europaeus</td><td>9.25</td><td>9250</td><td>9.132378831</td><td>6</td><td>6</td></tr><tr><td>Mesonychia</td><td>Mesonychidae</td><td>Dissacus filholi</td><td>16.89</td><td>16890</td><td>9.73447701</td><td>10</td><td>10</td></tr><tr><td>Mesonychia</td><td>Mesonychidae</td><td>Dissacus progressus</td><td>56.59</td><td>56590</td><td>10.94358757</td><td>10</td><td>10</td></tr><tr><td>Mesonychia</td><td>Mesonychidae</td><td>Dissacus raslanloubatieri</td><td>13.14</td><td>13140</td><td>9.483416292</td><td>8</td><td>9</td></tr><tr><td>Mesonychia</td><td>Mesonychidae</td><td>Dissacus rougierae</td><td>9.01</td><td>9014</td><td>9.106573419</td><td>7</td><td>7</td></tr><tr><td>Hyaenodonta</td><td>Proviverrinae</td><td>Eoproviverra eisenmanni</td><td>0.05</td><td>50</td><td>3.912023005</td><td>7</td><td>7</td></tr><tr><td>Hyaenodonta</td><td>Proviverrinae</td><td>Eurotherium matthesi</td><td>6.47</td><td>6470</td><td>8.774931387</td><td>11</td><td>11</td></tr><tr><td>Hyaenodonta</td><td>Proviverrinae</td><td>Eurotherium theriodis</td><td>6.88</td><td>6880</td><td>8.836373931</td><td>13</td><td>14</td></tr><tr><td>Hyaenodonta</td><td>Sinopinae</td><td>Galecyon gallus</td><td>0.77</td><td>770</td><td>6.646390515</td><td>7</td><td>7</td></tr><tr><td>Hyaenodonta</td><td>Sinopinae</td><td>Galecyon morloi</td><td>0.6</td><td>600</td><td>6.396929655</td><td>7</td><td>7</td></tr><tr><td>Hyaenodonta</td><td>Hyaenodontinae</td><td>Hyaenodon brachyrhynchus</td><td>17.06</td><td>17060</td><td>9.744491821</td><td>17</td><td>23</td></tr><tr><td>Hyaenodonta</td><td>Hyaenodontinae</td><td>Hyaenodon gervaisi</td><td>50.36</td><td>50360</td><td>10.82695249</td><td>17b</td><td>29</td></tr><tr><td>Hyaenodonta</td><td>Hyaenodontinae</td><td>Hyaenodon heberti</td><td>30.53</td><td>30530</td><td>10.32646509</td><td>17b</td><td>19</td></tr><tr><td>Hyaenodonta</td><td>Hyaenodontinae</td><td>Hyaenodon minor</td><td>7.65</td><td>7650</td><td>8.942460927</td><td>17a</td><td>18</td></tr><tr><td>Hyaenodonta</td><td>Hyaenodontinae</td><td>Hyaenodon requieni</td><td>38.3</td><td>38300</td><td>10.55320518</td><td>17a</td><td>20</td></tr><tr><td>Hyaenodonta</td><td>Hyaenodontinae</td><td>Hyaenodon rossignoli</td><td>4.48</td><td>4480</td><td>8.407378325</td><td>18</td><td>19</td></tr><tr><td>Hyaenodonta</td><td>Hyainailourinae</td><td>Kerberos langebadreae</td><td>87.69</td><td>87690</td><td>11.38156315</td><td>16</td><td>16</td></tr><tr><td>Hyaenodonta</td><td>Proviverrinae</td><td>Leonhardtina godinoti</td><td>0.99</td><td>990</td><td>6.897704943</td><td>10</td><td>10</td></tr><tr><td>Hyaenodonta</td><td>Proviverrinae</td><td>Leonhardtina meridianum n. sp.</td><td>0.84</td><td>840</td><td>6.733401892</td><td>10</td><td>10</td></tr><tr><td>Hyaenodonta</td><td>Proviverrinae</td><td>Leonhardtina gracilis</td><td>1.73</td><td>1730</td><td>7.455876687</td><td>12</td><td>13</td></tr><tr><td>Hyaenodonta</td><td>Proviverrinae</td><td>Lesmesodon behnkeae</td><td>8.31</td><td>8310</td><td>9.025214888</td><td>11</td><td>11</td></tr><tr><td>Hyaenodonta</td><td>Proviverrinae</td><td>Lesmesodon edingeri</td><td>2.21</td><td>2210</td><td>7.700747795</td><td>11</td><td>11</td></tr><tr><td>Hyaenodonta</td><td>Proviverrinae</td><td>Matthodon menui</td><td>12.31</td><td>12310</td><td>9.418167219</td><td>10</td><td>10</td></tr><tr><td>Hyaenodonta</td><td>Proviverrinae</td><td>Matthodon peignei n. sp.</td><td>15.75</td><td>15750</td><td>9.664595644</td><td>10</td><td>10</td></tr><tr><td>Hyaenodonta</td><td>Proviverrinae</td><td>Matthodon tritens</td><td>19.15</td><td>19150</td><td>9.860057995</td><td>11</td><td>11</td></tr><tr><td>Hyaenodonta</td><td>Proviverrinae</td><td>Minimovellentodon russelli</td><td>1.18</td><td>1180</td><td>7.073269717</td><td>8</td><td>9</td></tr><tr><td>Hyaenodonta</td><td>Proviverrinae</td><td>Morlodon vellerei</td><td>2.25</td><td>2250</td><td>7.718685495</td><td>8</td><td>10</td></tr><tr><td>Oxyaenidae</td><td>Oxyaeninae</td><td>Oxyaena gulo</td><td>12.3</td><td>12300</td><td>9.417354541</td><td>8</td><td>9</td></tr><tr><td>Oxyaenidae</td><td>Oxyaeninae</td><td>Oxyaena woutersi</td><td>6.48</td><td>6480</td><td>8.776475789</td><td>7</td><td>7</td></tr><tr><td>Hyaenodonta</td><td>Proviverrinae</td><td>Oxyaenoides aumelasiensis n. sp.</td><td>4.24</td><td>4240</td><td>8.352318548</td><td>10</td><td>10</td></tr><tr><td>Hyaenodonta</td><td>Proviverrinae</td><td>Oxyaenoides bicuspidens</td><td>14.19</td><td>14190</td><td>9.56029277</td><td>11</td><td>11</td></tr><tr><td>Hyaenodonta</td><td>Proviverrinae</td><td>Oxyaenoides lindgreni</td><td>9.71</td><td>9710</td><td>9.180911561</td><td>10</td><td>10</td></tr><tr><td>Hyaenodonta</td><td>Proviverrinae</td><td>Oxyaenoides schlosseri</td><td>16.93</td><td>16930</td><td>9.736842475</td><td>13</td><td>14</td></tr><tr><td>Mesonychia</td><td>Mesonychidae</td><td>Pachyaena cf. gracilis</td><td>38.98</td><td>38980</td><td>10.57080397</td><td>8</td><td>9</td></tr><tr><td>Mesonychia</td><td>Mesonychidae</td><td>Pachyaena cf. ossifraga</td><td>74.89</td><td>74890</td><td>11.22377565</td><td>8</td><td>9</td></tr><tr><td>Mesonychia</td><td>Mesonychidae</td><td>Pachyaena gigantea</td><td>162.44</td><td>162440</td><td>11.99806398</td><td>8</td><td>9</td></tr><tr><td>Hyaenodonta</td><td>Proviverrinae</td><td>Paenoxyaenoides liguritor</td><td>18.02</td><td>18020</td><td>9.799237531</td><td>18</td><td>18</td></tr><tr><td>Oxyaenidae</td><td>Ambloctinae</td><td>Palaeonictis gigantea</td><td>12.04</td><td>12040</td><td>9.395989719</td><td>7</td><td>9</td></tr><tr><td>Hyaenodonta</td><td>Proviverrinae</td><td>Paracynohyaenodon magnus</td><td>3.85</td><td>3850</td><td>8.255828427</td><td>16</td><td>16</td></tr><tr><td>Hyaenodonta</td><td>Proviverrinae</td><td>Paracynohyaenodon schlosseri</td><td>2.97</td><td>2970</td><td>7.996317232</td><td>16</td><td>17a</td></tr><tr><td>Hyaenodonta</td><td>Hyainailourinae</td><td>Parapterodon lostangensis</td><td>103.62</td><td>103620</td><td>11.54848564</td><td>19</td><td>19</td></tr><tr><td>Hyaenodonta</td><td>Hyainailourinae</td><td>Paroxyaena galliae</td><td>52.78</td><td>52780</td><td>10.87388761</td><td>16</td><td>17b</td></tr><tr><td>Hyaenodonta</td><td>Proviverrinae</td><td>Parvagula palulae</td><td>0.06</td><td>60</td><td>4.094344562</td><td>7</td><td>7</td></tr><tr><td>Hyaenodonta</td><td>Proviverrinae</td><td>Praecodens acutus</td><td>1.09</td><td>1090</td><td>6.993932975</td><td>13</td><td>13</td></tr><tr><td>Hyaenodonta</td><td>Proviverrinae</td><td>Preregidens langebadrae</td><td>3.53</td><td>3530</td><td>8.16905315</td><td>8</td><td>9</td></tr><tr><td>Hyaenodonta</td><td>Proviverrinae</td><td>Prodissopalis eocaenicus</td><td>18.02</td><td>18020</td><td>9.799237531</td><td>12</td><td>14</td></tr><tr><td>Hyaenodonta</td><td>Proviverrinae</td><td>Protoproviverra palaeonictides</td><td>0.91</td><td>910</td><td>6.8134446</td><td>10</td><td>10</td></tr><tr><td>Hyaenodonta</td><td>Sinopinae</td><td>Prototomus girardoti</td><td>0.45</td><td>450</td><td>6.109247583</td><td>7</td><td>9</td></tr><tr><td>Hyaenodonta</td><td>Sinopinae</td><td>Prototomus minimus</td><td>0.24</td><td>240</td><td>5.480638923</td><td>7</td><td>9</td></tr><tr><td>Hyaenodonta</td><td>Proviverrinae</td><td>Proviverra typica</td><td>0.42</td><td>420</td><td>6.040254711</td><td>12</td><td>13</td></tr><tr><td>Hyaenodonta</td><td>Hyainailourinae</td><td>Pterodon dasyuroides</td><td>51.56</td><td>51560</td><td>10.85050146</td><td>18</td><td>20</td></tr><tr><td>Hyaenodonta</td><td>Proviverrinae</td><td>Quercytherium simplicidens</td><td>3.12</td><td>3120</td><td>8.045588281</td><td>17a</td><td>18</td></tr><tr><td>Hyaenodonta</td><td>Proviverrinae</td><td>Quercytherium tenebrosum</td><td>2.89</td><td>2890</td><td>7.969011781</td><td>16</td><td>17a</td></tr></tbody></table>
*Alienetherium buxwilleri* Lange-Badré, 1981

*Allopterodon* Ginsburg, 1978

*Allopterodon bulbosus* (Lange-Badré, 1979)

*Allopterodon minor* (Filhol, 1877)

*Allopterodon torvidus* (Van Valen, 1965)

*Anchomomys* Stehlin, 1916

*Apterodon gaudryi* Fischer, 1880

*Apterodon intermedius* Lange-Badré & Böhme, 2005

*Arfia gingerichi* Smith & Smith, 2001

*Aumelasia gabineaudi* Sudre, 1980

*Boritia duffaudi* Solé, Falconnet & Laurent, 2014

*Cartierodon egerkingensis* Solé & Mennecart, 2019

*Cynohyaenodon* Filhol, 1873

*Cynohyaenodon blainvillei* Brunet, 1977

*Cynohyaenodon cayluxi* Filhol, 1873

*Cynohyaenodon lautricensis* Lange-Badré, 1978

*Cynohyaenodon rutimeyeri* Van Valen, 1965

*Cynohyaenodon trux* Van Valen, 1965

*Dichobune robertiana* Gervais, 1848

*Dissacus blayaci* Stehlin, 1926

*Dissacus europaeus* Lemoine, 1891

*Dissacus filholi* (Lemoine, 1891)

*Dissacus progressus* Crusafont Pairó & Golpe-Posse, 1968

*Dissacus raslanloubatieri* Solé, Godinot, Laurent, Galoyer & Smith, 2019

*Dissacus rougierae* Solé, Godinot, Laurent, Galoyer & Smith, 2019

*Eoproviverra eisenmanni* (Godinot, 1981)

*Eurotherium* Polly & Lange-Badré, 1993

*Eurotherium matthesi* (Lange-Badré & Haubold, 1990)

*Eurotherium theriodis* (Van Valen, 1965)

*Galecyon* Gingerich & Deutsch, 1989

*Galecyon gallus* Solé, Gheerbrant & Godinot, 2013

*Galecyon morloi* Smith & Smith, 2001

*Gazinocyon vulpeculus* (Matthew & Granger, 1915)

*Hyaenodon brachyrhynchus* de Blainville, 1839

*Hyaenodon exiguus* (Gervais, 1872)

*Hyaenodon gervaisi* Martin, 1906

*Hyaenodon heberti* Filhol, 1876

*Hyaenodon minor* Gervais, 1852

*Hyaenodon requieni* Gervais, 1848-1852

*Hyaenodon rossignoli* Lange-Badré, 1979

*Kerberos langebadreae* Solé, Amson, Borths, Vidalenc, Morlo & Bastl, 2015

*Leonhardtina* Matthes, 1952

*Leonhardtina godinoti* Solé, Falconnet & Laurent, 2014

*Leonhardtina gracilis* Matthes, 1952

*Leonhardtina meridianum* n. sp.

*Lesmesodon* Morlo & Habersetzer, 1999

*Lesmesodon behnkeae* Morlo & Habersetzer, 1999

*Lesmesodon edingeri* (Springhorn, 1982)

*Lophiodon isselensis* Fischer, 1929

*Lophiodon parisiense*

*Matthodon* Lange-Badré & Haubold, 1990

*Matthodon menui* (Rich, 1971)

*Matthodon peignei* n. sp.

*Matthodon tritens* Lange-Badré & Haubold, 1990

*Minimovellentodon russelli* Solé, Falconnet & Laurent, 2014

*Molrodon* Solé, 2013

*Molrodon vellerei* Solé, 2013

*Namasector* Morales, Pickford & Salesa, 2008

*Namasector soriae* Morales, Pickford & Salesa, 2008

*Oxyaena gulo* Matthew & Granger, 1915

*Oxyaena woutersi* (Lange-Badré & Godinot, 1982)

*Oxyaenoides* Matthes, 1967

*Oxyaenoides aumelasiensis* n. sp.

*Oxyaenoides bicuspidens* Matthes, 1967

*Oxyaenoides lindgreni* (Rich, 1971)

*Oxyaenoides schlosseri* (Rütimeyer, 1891)

*Pachyaena* Cope, 1874

*Pachyaena gigantea* Osborn & Wortman, 1892

*Paenoxyaenoides liguritor* Lange-Badré, 1979

*Pakakali* Borths & Stevens, 2017

*Pakakali rukwaensis* Borths & Stevens, 2017

*Palaeonictis gigantea* de Blainville, 1842

*Paracynohyaenodon magnus* Crochet, 1888

*Paracynohyaenodon schlosseri* Martin, 1906

*Parapterodon lostangensis* Lange-Badré, 1979

*Paroxyaena galliae* Filhol, 1881

*Parvagula* Lange-Badré, 1987

*Parvagula palulae* Lange-Badré, 1987

*Praecodens acutus* Lange-Badré, 1981

*Preregidens* Solé, Falconnet & Vidalenc, 2015

*Preregidens langebadrae* Solé, Falconnet & Vidalenc, 2015

*Prionogale* Schmidt-Kittler & Heizmann, 1991

*Prionogale breviceps* Schmidt-Kittler & Heizmann, 1991

*Prodissopalsis eocaenicus* Matthes, 1952

*Protoproviverra palaeonictides* (Lemoine, 1880)

*Prototomus girardoti* Smith & Smith, 2001

*Prototomus minimus* Smith & Smith, 2001

*Proviverra* Rütimeyer, 1862

*Proviverra typica* Rütimeyer, 1862

*Pterodon dasyuroides* de Blainville, 1839

*Quercytherium simplicidens* Lange-Badré, 1975

*Quercytherium tenebrosum* Filhol, 1880

*Simbakubwa kutokaafrika* Borths & Stevens, 2019

*Thereutherium* Filhol, 1876

*Thereutherium thylacodes* Filhol, 1876