Petrosal and bony labyrinth morphology of the stem paenungulate mammal (Paenungulatomorpha) Ocepeia daouiensis from the Paleocene of Morocco.

Based on high-resolution computed tomography, we describe in detail the petrosal and inner ear anatomy of one of the few known African stem paenungulates (Paenungulatomorpha), Ocepeia daouiensis from the Selandian of the Ouled Abdoun phosphate basin (Morocco). The petrosal of Ocepeia displays some remarkable, probably derived features (among eutherians) such as relatively small pars cochlearis, pars canalicularis labyrinth (including small semicircular canals), a large wing-like pars mastoidea, a large and inflated tegmen tympani, and the dorsoventral orientation of the large canal for the ramus superior. The presence of small semicircular canals in Ocepeia is an interesting shared trait with tenrecoidean afrotherians. Otherwise, and consistent with a general primitive skull morphology, the middle ear and labyrinth of Ocepeia daouiensis is characterised by many plesiomorphic traits close to the eutherian generalised plan. This adds to the rather generalised morphology of the earliest crown paenungulates such as Eritherium, Phosphatherium and Seggeurius to support an ancestral paenungulatomorph morphotype poorly derived from the eutherian pattern. As a result, Ocepeia provides key morphological and fossil data to test phylogenetic relationships of the Afrotheria (including Paenungulatomorpha) at the placental root mostly inferred from molecular studies.

Early African Fossils Elucidate the Origin of Embrithopod Mammals.

Modern mammals rapidly evolved in the early Cenozoic in all continental provinces, including in Africa, with one of the first placental branches, the Afrotheria [1, 2]. Afrotherian evolution is at the origin of the major radiation of African ungulate-like mammals, including extant hyrax, elephant, and sea cow orders, which all belong to the Paenungulata. The paenungulate radiation also includes the extinct order Embrithopoda of uncertain interordinal relationships, which is best known for the giant and strangely specialized Oligocene genus Arsinoitherium. The Ouled Abdoun basin, Morocco, yielded exceptional Paleocene-Eocene fossils documenting the early paenungulate evolution [3-8]. Here we report two new small Ypresian species, Stylolophus minor n.g., n.sp. and cf. Stylolophus sp., which are the earliest and most primitive embrithopods. The cladistic analysis relates the Embrithopoda to crown paenungulates as the stem-group of the Tethytheria, which makes crown tethytherians restricted to extant elephant and sea cow orders. The Embrithopoda is therefore an early tethytherian offshoot predating the elephant and sea cow divergence. The resulting phylogeny supports a strictly African early radiation of the paenungulates excluding the Phenacolophidae and Anthracobunia. It sustains an at least early Paleocene African origin of the Embrithopoda. The unique tooth pattern of the embrithopods (hyperdilambdodont and pseudolophodont molars) is resolved as evolving early and directly from the dilambdodont (W-shaped labial molar crests) ancestral paenungulate morphotype. The specialized upper molar morphology with two transverse crests is convergent and non-homologous in embrithopods and crown Tethytheria. These convergences for specialized folivorous diet were driven by free herbivorous African niches in the early Paleogene.

Convergence of Afrotherian and Laurasiatherian Ungulate-Like Mammals: First Morphological Evidence from the Paleocene of Morocco.

Molecular-based analyses showed that extant "ungulate" mammals are polyphyletic and belong to the two main clades Afrotheria (Paenungulata) and Laurasiatheria (Euungulata: Cetartiodactyla-Perissodactyla). However, paleontological and neontological studies hitherto failed to demonstrate the morphological convergence of African and Laurasian "ungulate" orders. They support an "Altungulata" group including the Laurasian order Perissodactyla and the African superorder Paenungulata and characterized especially by quadritubercular and bilophodont molars adapted for a folivorous diet. We report new critical fossils of one of the few known African condylarth-like mammal, the enigmatic Abdounodus from the middle Paleocene of Morocco. They show that Abdounodus and Ocepeia display key intermediate morphologies refuting the homology of the fourth main cusp of upper molars in Paenungulata and Perissodactyla: Paenungulates unexpectedly have a metaconule-derived pseudohypocone, instead of a cingular hypocone. Comparative and functional dental anatomy of Abdounodus demonstrates indeed the convergence of the quadritubercular and bilophodont pattern in "ungulates". Consistently with our reconstruction of the structural evolution of paenungulate bilophodonty, the phylogenetic analysis relates Abdounodus and Ocepeia to Paenungulata as stem taxa of the more inclusive new clade Paenungulatomorpha which is distinct from the Perissodactyla and Anthracobunidae. Abdounodus and Ocepeia help to identify the first convincing synapomorphy within the Afrotheria-i.e., the pseudohypocone-that demonstrates the morphological convergence of African and Laurasian ungulate-like placentals, in agreement with molecular phylogeny. Abdounodus and Ocepeia are the only known representatives of the early African ungulate radiation predating the divergence of extant paenungulate orders. Paenungulatomorpha evolved in Africa since the early Tertiary independently from laurasiatherian euungulates and "condylarths" such as apheliscids. The rapid early Tertiary radiation of the Afrotheria and Paenungulatomorpha, as illustrated by the Paleocene Moroccan mammals, is concurrent with that of the Laurasiatheria in a general, explosive mammal evolution in both the South and North Tethyan continents following the K/Pg event.

The ear region of earliest known elephant relatives: new light on the ancestral morphotype of proboscideans and afrotherians.

One of the last major clades of placental mammals recognized was the Afrotheria, which comprises all main endemic African mammals. This group includes the ungulate-like paenungulates, and among them the elephant order Proboscidea. Among afrotherians, the petrosal anatomy remains especially poorly known in Proboscidea. We provide here the first comparative CT scan study of the ear region of the two earliest known proboscideans (and paenungulates), Eritherium and Phosphatherium, from the mid Palaeocene and early Eocene of Morocco. It is helpful to characterize the ancestral morphotype of Proboscidea to understand petrosal evolution within proboscideans and afrotherians. The petrosal structure of these two taxa shows several differences. Eritherium is more primitive than Phosphatherium and closer to the basal paenungulate Ocepeia in several traits (inflated tegmen tympani, very deep fossa subarcuata and ossified canal for ramus superior of stapedial artery). Phosphatherium, however, retains plesiomorphies such as a true crus commune secundaria. A cladistic analysis of petrosal traits of Eritherium and Phosphatherium among Proboscidea results in a single tree with a low level of homoplasy in which Eritherium, Phosphatherium and Numidotherium are basal. This contrasts with previous phylogenetic studies showing homoplasy in petrosal evolution among Tethytheria. It suggests that evolutionary modalities of petrosal characters differ with the taxonomic level among Afrotheria: noticeable convergences occurred among the paenungulate orders, whereas little homoplasy seems to have occurred at intra-ordinal level in orders such as Proboscidea. Most petrosal features of both Eritherium and Phosphatherium are primitive. The ancestral petrosal morphotype of Proboscidea was not specialized but was close to the generalized condition of paenungulates, afrotherians, and even eutherians. This is consistent with cranial and dental characters of Eritherium, suggesting that the ancestral morphotypes of the different paenungulate orders were close to each other. Specializations occurred rapidly after the ordinal radiation of Paenungulata.