Paenanthracotherium bergeri, Scherler & Lihoreau & Becker, 2019

PAENANTHRACOTHERIUM BERGERI SP. NOV.

( FIGS 5–7)

urn:lsid:zoobank.org:act:81139BAC-A379-41DC-A9AB-9FD12BB5B20A Anthracotherium magnum Cuvier from Auvergne; De

Blainville, 1848: 126, 129, 131, 133, pl. ‘ A. magnum ,

systême dentaire: d’Auvergne’ [viewed].

Anthracotherium magnum Cuvier from Moyssac; De Blainville, 1848: 127, pl. ‘ A. magnum , systême dentaire: du Tarn et Garonne’.

Anthracotherium magnum Cuvier from Digoin; De Blainville, 1848: 127, 129, 130, pl. ‘ A. magnum , systême dentaire: de Digoin’ [viewed].

Anthracotherium magnum Cuvier ; Leymerie, 1851: 388–392 [viewed].

‘Un Anthracotherium different de Anthracotherium magnum ’; Gervais, 1852: 97 [viewed].

Anthracotherium magnum Cuvier ; Rütimeyer, 1857: 29.

Anthracotherium Cuvier ; Kowalevsky, 1873: pl. 11, figs 37, 52–54, pl. 14, fig. 96.

Anthracotherium bumbachense ; Stehlin, 1910: 168–169.

Anthracotherium cf. bumbachense Stehlin ; Depéret, 1913: 350–351 [viewed].

Anthracotherium bumbachense* Stehlin ; Roman & Boucher, 1936: 10–38, figs 2–5, 9, 11.2, 13, pl. 1–4 [* holotype, viewed].

‘ Anthracotherium jeune de St-Henri’; Roman & Boucher, 1936: fig. 7 [viewed].

Anthracotherium Cuvier ; Lavocat, 1951: 130 [viewed in part].

Anthracotherium cuvieri Gaudry ; Brunet & Vianey-Liaud, 1987: 31 [viewed].

Anthracotherium bumbachense Stehlin ; Brunet & Vianey-Liaud, 1987: 31 [viewed].

Anthracotherium magnum Cuvier ; Sudre, 1995: 214– 216 [viewed].

Anthracotherium bumbachense Stehlin ; Sudre, 1995: 216 [viewed].

Anthracotherium cf. magnum ; BiochroM’97, 1997: 791 [viewed].

Anthracotherium bumbachense Stehlin ; Hugueney, 1997: 420, table 4 [viewed].

Anthracotherium magnum Cuvier ; Hugueney, 1997: 420 [viewed].

Anthracotherium sp. ; Becker et al., 2004: 6–7 [viewed].

Elliotherium murphyi nomen nudum; Scherler, 2011: 85–93, figs 5.2, 5.5, 5.6, 5.10, table 5.1 [viewed].

Anthracotherium bumbachense Stehlin ; Alloing-Séguier et al., 2014: 670–671, 678, table 2 [viewed].

Elliotherium murphyi nomen nudum; Pickford, 2016: 249 [viewed].

Differential diagnosis

Large Paenanthracotherium ( Fig. 4), differing from other species of the genus by the presence of an ectometafossid on the lower molars (200), a shallow and constant cingulid in front of the transverse valley of the lower molars (371), and sometimes the presence of an ectocristylid (381), a less prominent parastyle (681), a very small metastyle on the upper molars (700), and a division into two of the postprotocrista on P4 (1051). Paenanthracotherium bergeri differs from P. strategus in having the posthypocristid on m1 and m2 joining the postentocristid (331), no clear junction between the protocone and the metaconule on M1 and M2 (520), no extension of the postparacristule in the transverse valley (590), the postmetacrista connecting labially to the metastyle (651), and a strong development of the distostyle on P4 (1031). Paenanthracotherium bergeri differs from P. hippoideum in having a larger size ( Fig. 4), a distolingual cingulid on p4 forming a continuous wall until the distostylid (70), a strong premetacristid on the lower molars (150), a lack of connection between the premetacristid and the preprotocristid on the lower molars (161), a slightly inflated prehypocristid (301), an endohypocristid on the lower molars (341), the presence of a unique accessory cristulid mesially directed in front of the hypoconulid (between the pre- and the posthypocristulid) on m3 (460), the premetacristule divided into two mesial arms on the upper molars (531), a fused symphysis in adult specimens (741), the presence of a p2–p3 diastema (870), and a postparafossa on P3 (950).

Holotype

Skull with left I2–P2, M3 and right I1–M2 (FSL-213772) ( Figs 5, 7).

Paratypes

Maxilla with left I1–I2, C–M3 and right I1–M3 (FSL-213773), mandible with left i1, p4–m3 and right p2– m3 (FSL-213774), and mandible with left i1, i3–m3 and right i1, i3–c, p2–m3 (FSL-213772) ( Figs 6, 7).

Type locality and horizon

La Bénisson-Dieu, France (late Early Oligocene, MP24).

Etymology

In memory of Jean-Pierre Berger, in recognition of his contribution to palaeontology and for having initiated the review of European anthracotheres.

Distribution

Oligocene of Europe: La Bénissons-Dieu (MP24), Digoin (MP24), Le Garouillas (MP25), Les Milles (MP26), Pech Desse (MP28), La Comberatière (MP29), Moissac (Oligocene), Lamontgie (MP23–29?), France and Petrosani (Oligocene), Romania (e.g. De Blainville, 1848; Stehlin, 1910; Roman & Boucher, 1936; Lavocat, 1951; Sudre, 1995; Hugueney, 1997; Scherler, 2011; Scherler et al., 2013; see Supporting Information, Appendix S5 for complete references).

Remarks

The fossils from La Bénissons-Dieu were previously assigned to A. bumbachense by Roman & Boucher (1936), based on the reduced lateral metapods compared with the medial ones. It appeared that the specimens from Bumbach ( Switzerland), assigned by Stehlin (1910) to the new species A. bumbachense , displayed a similar feature. However, when comparing the different metapods from La Bénissons-Dieu and Bumbach, there is a considerable variation in size and slenderness ( Scherler, 2011). Additionally, owing to the scarcity and poor preservation of the material from Bumbach, the attribution of the skull, maxillae and mandibles from La Bénissons-Dieu to the same species is difficult and ambiguous. Moreover, the specimens from Bumbach cannot be assigned clearly to Anthracotherium for the above-mentioned reasons ( Scherler, 2011). However, three incomplete mandibles from the French localities of Le Garouillas, Quercy (UM-GAR2313; Sudre, 1995), Moissac, Tarn et Garonne (MHNT.PAL.MAM.2002.3; Leymerie, 1851) and Lamontgie near Issoire, Puy de Dôme (figured in De Blainville, 1848: specimen Cr4223MNHN coll. Abbé Croizet in Lavocat, 1951; Hugueney, 1997) can be assigned, without doubt, to the same species based on the size, the more selenodont pattern than in A.magnum and the lack of an entoconulid on m3s. The taxon was named in an unpublished thesis ( Scherler, 2011) and was cited later without formal description, diagnosis or figuration ( Pickford, 2016). We thus consider Elliotherium murphyi to be a nomen nudum and propose Paenanthracotherium bergeri gen. nov., sp. nov. as the new name identifying the largest Paenanthracotherium species from Europe.

Figure 4 illustrates the size variations of the different specimens of Paenanthracotherium by plotting the P4/p4, M1/m1, M2/m2 and M3/m3, and Table 1 shows their mean values. The measurements of all studied specimens are given in the Supporting Information (Appendix S6). The two European species of Paenanthracotherium , P. bergeri and P. hippoideum , can be differentiated easily by their size, with the former being much larger than the latter. Moreover, these measurements help to confirm the attribution of the specimen from Pech Desse, France to P. bergeri .

Description

Upper dentition: The upper dentition formula is complete (3I, 1C, 4P, 3M). The upper incisors display a continuous and slightly developed lingual cingulum. Their lateral outline is triangular, with mesial and distal crests reaching the cingula. The incisor row is continuous, but there is a diastema between I3 and C.The upper canine is very large, with a high and sharp crown. Its transverse section is circular to ellipsoidal. P1 has a triangular occlusal outline and lacks any stylar development. The labial cingulum is weak and discontinuous, and the lingual one is slightly developed and continuous. The pre- and postparacristae are aligned and oblique in relationship to tooth row axis. The postparafossa is simple and weakly marked. P2 is similar to P1 but bears a slightly developed metastyle. The cingula are weak and continuous all around the tooth; the distal one is slightly more developed. It lacks an ectoparacrista, and the distolingual basin (endoparafossa) is simple and slightly developed. On P3, the para- and metastyles are developed. The cingula are continuous, but the mesial and labial ones are less developed compared with the distal and lingual ones. Moreover, the distolingual cingulum does not bear any protostyle, which is well developed in the Anthracotherium . The postparafossa is marked. The para- and metastyles on P4 are labially well developed, which differs in the species of Anthracotherium . The cingulum is present all around the tooth,