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While key early(iest) fossils were recently discovered for several crown afrotherian mammal orders, basal afrotherians, e.g., early Cenozoic species that comprise sister taxa to Paenungulata, Afroinsectiphilia or Afrotheria, are nearly unknown, especially in Africa. Possible stem condylarth-like relatives of the Paenungulata (hyraxes, sea-cows, elephants) include only  Abdounodus hamdii  and  Ocepeia daouiensis  from the Selandian of Ouled Abdoun Basin, Morocco, both previously only documented by lower teeth. Here, we describe new fossils of  Ocepeia , including  O.grandis  n. sp., and a sub-complete skull of  O. daouiensis , the first known before the Eocene for African placentals.  O.daouiensis  skull displays a remarkable mosaic of autapomophic, ungulate-like and generalized eutherian-like characters. Autapomorphies include striking anthropoid-like characters of the rostrum and dentition. Besides having a basically eutherian-like skull construction,  Ocepeia daouiensis  is characterized by ungulate-like, and especially paenungulate-like characters of skull and dentition (e.g., selenodonty). However, some plesiomorphies such as absence of hypocone exclude  Ocepeia  from crown Paenungulata. Such a combination of plesiomorphic and derived characters best fits with a stem position of  Ocepeia  relative to Paenungulata. In our cladistic analyses  Ocepeia  is included in Afrotheria, but its shared derived characters with paenungulates are not optimized as exclusive synapomorphies. Rather, within Afrotheria  Ocepeia  is reconstructed as more closely related to insectivore-like afroinsectiphilians (i.e., aardvarks, sengis, tenrecs, and golden moles) than to paenungulates. This results from conflict with undetected convergences of Paenungulata and Perissodactyla in our cladistic analysis, such as the shared bilophodonty. The selenodont pattern best supports the stem paenungulate position of  Ocepeia ; that, however, needs further support. The remarkable character mosaic of  Ocepeia  makes it the first known “transitional fossil” between insectivore-like and ungulate-like afrotherians. In addition, the autapomorphic family Ocepeiidae supports the old – earliest Tertiary or Cretaceous – endemic evolution of placentals in Africa, in contrast to hypotheses rooting afrotherians in Paleogene Laurasian “condylarths”.

Ocepeia (Middle Paleocene of Morocco): The Oldest Skull of an Afrotherian Mammal