Paenanthracotherium, Scherler & Lihoreau & Becker, 2019

GENUS PAENANTHRACOTHERIUM GEN. NOV.

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Diagnosis ( Fig. 3)

Medium to large Anthracotheriinae s.l. ( Fig. 4) with rather selenodont cheek teeth, p3 with entostylid (61), lower molars that lack clear endometacristid (210) and with main arm of the prehypocristid connecting to the postmetafossid (321), P3 without distolingual cingular style (910) and without ectoparacrista (930), upper molars with labial cusps with pinched ribs (481) and without secondary cristules labial to metaconule (560).

Differential diagnosis ( Fig. 3)

Paenanthracotherium differs from all Anthracotheriinae s.l. by the absence of or reduced endometacristid on the lower molars (210) and the pinched mesiodistal development of the ribs of labial cusps on the upper molars (481). Within Anthracotheriinae s.l., Paenanthracotherium differs from Heptacodon and A. birmanicus by a mesiolingual secondary cristid on p4, connecting the preprotocristid with the lingual margin (81), an entostylid on p4 (111), an incomplete but present ectohypocristulid on m3 (391; absent in Heptacodon and complete in A. birmanicus ) and no cristids on the lower canine (800). It differs from Heptacodon and Prominatherium by the lack of postectoprotocrista on P4 (1010). Moreover, it differs from Heptacodon by lower molars with an ectoentocristid (270), no distal cingulid on m3 (450), no secondary ectometafossule lingual to the ectometacristule (580), unfused lingual roots of the upper molars (710), presence of a diastema between p1 and p2 (861), P1 and P2 (891) and P2 and P3 (901), no postprotofossa on P4 (1060), and by the preprotocrista on P4 that joins the base of the paracone then the mesiostyle (1000). It differs from Anthracotherium and A. birmanicus by the distolabial orientation of the postprotocristid on p3 (22), the lack of postectoprotocristid (170), postprotofossid (180) and posthypofossid on lower molars (350), and a unique prehypocristid, not divided into two mesial arms (291). It differs from Anthracotherium by the lack of entoconulid on m3 (420).

Type species

Paenanthracotherium bergeri sp. nov. from La Bénissons-Dieu ( France), late Early Oligocene .

Other included species

Paenanthracotherium hippoideum ( Rütimeyer, 1857) ; Paenanthracotherium strategus ( Forster-Cooper, 1913) .

Etymology

From the Latin adverb ‘paene’, meaning almost, and the genus name Anthracotherium Cuvier, 1822 , meaning charcoal beast in Latin, in reference to the similarities between both genera and to the inclusion of this genus within Anthracotherium for at least 165 years.

Distribution

Late Early to Late Oligocene in France, Switzerland, Germany, Romania and Pakistan (e.g. Roman & Boucher, 1936; Pickford, 1987; Sudre, 1995; Scherler, 2011; Scherler et al., 2013; see Supporting Information, Appendix S5 for a more exhaustive list of localities with references).

Remark on the included species Paenanthracotherium hippoideum and P. strategus were previously assigned to the genera Anthracotherium and Brachyodus , respectively ( Rütimeyer, 1857; Forster-Cooper, 1913). However, their similar morphology and the inferred phylogeny allow us to gather them in a new genus. The specimens from St Menoux and St Henri, France were attributed to A. cuvieri by Gaudry (1873) and A. bumbachense by Roman & Boucher (1936). Nevertheless, the morphology of the m3 lacking the entoconulid, the general shape of the teeth, and the dimensions similar to those of P. hippoideum from Aarwangen, Switzerland ( Fig. 4) allow us to attribute this material to A. bumbachense .