Creodonta

Creodonta is an extinct, possibly polyphyletic order of carnivorous mammals that lived from the Paleocene to the Miocene epochs. Because they both possess carnassial teeth, creodonts and carnivorans were once thought to have shared a common ancestor, but given that different teeth are involved in making up the carnassials (both between creodonts and carnivorans and between the main groups of creodonts), this appears to be a case of evolutionary convergence. Carnassials are also known in other mammal clades, such as in the extinct bat Necromantis. Two distinct families were historically thought to compose the order: Oxyaenidae and Hyaenodontidae. They may both represent separate orders of fossil mammals related to carnivoramorphs or are descended from more basal taxa. Creodonts had an extensive range, both geographically and temporally. They are known from the late Paleocene through the late Oligocene in North America, the early Eocene through late Oligocene in Europe, from the early Eocene through late Miocene in Asia, and from the Paleocene to the late Miocene in Africa. Creodonts were the first large, obviously carnivorous mammals with the radiation of the oxyaenids in the late Paleocene. During the Paleogene, they were the most abundant form of terrestrial carnivore in the Old World. In Oligocene Africa, they were the dominant predatory group. They competed with the Mesonychids and the Entelodonts and ultimately outlasted them by the start of the Oligocene and by the middle of the Miocene respectively, but lost ground to the carnivorans. The last genus became extinct 11.1 million years ago, and carnivorans now occupy their ecological niches. 'Creodonta' was coined by Edward Drinker Cope in 1875. Cope included the oxyaenids and the viverravid Didymictis but omitted the hyaenodontids. In 1880. he expanded the term to include Miacidae, Arctocyonidae, Leptictidae (now Pseudorhyncocyonidae), Oxyaenidae, Ambloctonidae and Mesonychidae. Cope originally placed creodonts within the Insectivora. In 1884, however, he regarded them as a basal group from which both carnivorans and insectivorans arose. Hyaenodontidae was not included among the creodonts until 1909. Over time, various groups were removed, and by 1969 it contained, as it does today, only the oxyaenids and the hyaenodontids. One view of the position of the group is that Creodonta and Carnivora are sister taxa (within a superorder Ferae). Others have argued that insectivorans are more closely related to carnivorans, and creodonts, therefore, are basal eutherians. Others have suggested that Creodonta might not be monophyletic. Polly has argued that the only available synapomorphy between oxyaenids and hyaenodontids is a large metastylar blade on the first molar (M1), but he believes that that feature is common for all basal eutheria. Separating Oxyaenidae from Hyaenodontidae would also comport with biogeographic evidence, since the first oxyaenid is known from the North American early Paleocene and the first hyaenodontids are from very late Paleocene of North Africa. Complicating this arrangement is the tentative endorsement by Gunnell of the erection of a third family, Limnocyonidae. The group includes taxa that were once considered oxyaenids, such as Limnocyon, Thinocyon and Prolimnocyon Wortman had even erected a subfamily of Limnocyoninae within the oxyaenids. Van Valen nests the same subfamily (including Oxyaenodon) within Hyaenodontidae. Gunnell is agnostic whether Limnocyonidae is a group within Hyaenodontidae (although a sister group to the rest of hyaenodontids) or entirely separate. According to Gunnell, the defining features of the oxyaenids include: A small braincase low in the skull. The occiput wide at base and narrowing dorsally (to give it a triangular shape). The lacrimal bone makes a semicircular expansion on the face. The mandibles have heavy symphysis. M1 and m2 form the carnassials, while M3/m3 are absent. The manus and pes are plantigrade or subplantigrade. The fibula articulates with the calcaneum, and the astragalus articulates with the cuboid bone. The phalanges are compressed and fissured at the tip.