Systematic and evolutionary relationships of the hipparionine horses from Maragheh, Iran (Late Miocene, Turolian age)

Raymond L. Bernor

A systematic analysis of an hipparionine horse assemblage from Maragheh, Iran is made. A brief orientation to systematic philosophy and informal superspecific characterizations of some Old World hipparionines is given as a background to this work. A character state analysis of skulls is made, and has revealed five distinct species. A character state and stratigraphic trend analysis of isolated check tooth and postcranial remains, with known provenance, is also made. These two combined analyses reveal that the most resolute discrimination of hipparionine species and their evolutionary relationships occurs when multiple character complexes of associated skulls, maxillary and mandibular dentitions are made. When this is not possible, skulls have provided the best basis for discriminating species and their evolutionary relationships. Traditional characters of isolated cheek teeth and postcranial remains are shown here to offer limited information content for hipparionine phylogenetic systematics. The systematic portion of this study includes a comprehensive description of cranial and postcranial remains, and has further corroborated the distinction of five species which belong to at least three superspecific groups including: «Hipparion» geltyi sp. nov., Group 1; Hipparion prostylum (s. l.), and Hipparion campbelli sp. nov., Group 3; «Hipparíon» aff. moldavicum and «Hipparion» ?matthewi, Group 2. These species stratigraphic ranges and evolutionary relationships are also given here and argued to be important for establishing future hipparionine geochronologic correlations between a number of Eurasian late Miocene provinces.