The skull of Arsinoitherium (Mammalia, Embrithopoda) and the higher order interrelationships of ungulates

Nicholas Court

 Detailed anatomical description of arsinoithere cranial remains from the Lower Oligocene, Fayum Depression, Egypt, provides the basic data for a systematic investigation. All cranial and some postcranial features are assessed from a phylogenetic standpoint. Several soft tissue characters are then added to a cladistic analysis based on 54 derived ungulate morphological characters. The resulting phylogenetic hypothesis implies that perissodactyls, sirenians, proboscideans and arsinoitheres constitute a monophyletic unit (5 synapomorphies). However, increasing the tree length by 3 steps reveals a closer association between hyraxes and perissodactyls. Nevertheless, 13 synapomorphies link proboscideans, sirenians and  arsinoitheres to the exclusion of all other ungulates. Form of the sphenopalatine and ethmoid foramina, recurved posttympanic process, absence of a fenestra rotundum in the petrosal, vestigial paroccipital process of the exoccipital and the highly unusual absence of a hypoglossal foramen in the skull, imply a robust sister-group relationship between arsinoitheres and proboscideans. In this analysis artiodactyls share only one derived character with all other ungulates studied. Monophyly of Ungulata, including Artiodactyla, is therefore only weakly supported. It is argued that pedal anatomy of hyraxes is non-homologous with that of Tethytheria. Arsinoitherium should now be classified within Tethytheria, sharing a sister-group relationship with Proboscidea. Hyraxes are excluded, thus refuting the concept of Paenungulata. However, monophyly of the wider concept, Pantomesaxonia, containing hyraxes, perissodactyls, sirenians, proboscideans and now, arsinoitheres, is supported by this study.