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Recent molecular studies have recognized the skunks (Mephitinae) to be the sister taxon - of a clade comprising the Procyonidae and Mustelidae. These findings are inconsistent with the traditional placement of the skunks among mustelids, which is based on morphological evidence from extant taxa. This paper reports on a well-preserved cranium of a young individual, recovered from the middle Miocene deposits (MN 7+8, about 11-12 Ma) of Steinheim am Albuch, Germany. The fossil is the holotype of Palaeomephitis steinheimensis Jäger, 1839, which is here recognized as a senior subjective synonym of Trochotherium cyamoides Fraas, 1870 (consequently, Palaeomephitis Jäger, 1839 is a senior subjective synonym of Trochotherium Fraas, 1870). The specimen is identified as the oldest and most primitive mephitine cranium known to date, approaching the primitive morphology for the Mephitinae. It exhibits a combination of mephitine (accessory middle-ear chamber, lateral swelling of the squamosal) and mustelid (mustelid suprameatal fossa) synapomorphies, corroborating the view that skunks are derived from a mustelid ancestor. Its auditory bulla shows a slightly inflated and relatively large caudal entotympanic, which indicates that the uninflated and relatively small caudal entotympanics of adult mephitines, as well as their hypertrophied ectotympanics, are not primitive (as hitherto assumed) but derived, providing a synapomorphy that supports a sister-group relationship between the Mephitinae and Lutrinae.
The size and morphological characteristics of a skull of an arctomorph carnivoran mammal from Mouillac (old collection of the Phosphorites du Quercy, of unknown age) in France closely match those of the holotype of the earliest known procyonid Pseudobassaris riggsi and another skull referred to this species, both from old collections of the Phosphorites du Quercy (Caylus and Mouillac), probably earliest Late Oligocene in age. The skull is more primitive in morphology than those of Pseudobassaris riggsi and every other known procyonid, approaching a hypothetical primitive procyonid morphotype. The only, but methodologically fundamental, departure from this morphotype is the lack of the procyonid suprameatal fossa, which is the crucial synapomorphy of the family Procyonidae. To explain the phylogenetic and taxonomic status of the arctomorph represented by the skull, three competing hypotheses are put forward. Hypothesis A, which considers the arctomorph as an individual of Pseudobassaris riggsi, assumes that the procyonid suprameatal fossa first appeared in a common ancestor of Pseudobassaris and other procyonids but was still of variable occurence within Pseudobassaris riggsi. Hypothesis B, which proposes the arctomorph as a member of a new Pseudobassaris species ancestral to Pseudobassaris riggsi, concludes that the procyonid suprameatal fossa arose in Pseudobassaris riggsi and in the Procyonidae independently, excluding Pseudobassaris from the procyonids. Hypothesis C, which recognizes the arctomorph as a representative of a new species of a new genus of the paraphyletic procyonid stem group, presumes that the procyonid suprameatal fossa originated in a common ancestor of Pseudobassaris and other procyonids after the new genus had become detached from the ancestral stock of the Procyonidae.
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