Macroscelideans (elephant shrews or sengis) are small-bodied (25–540 g), cursorial (running) and saltatorial (jumping), insectivorous and omnivorous1 placental mammals represented by at least 15 extant African species classified in four genera2. Macroscelidea is one of several morphologically diverse but predominantly African placental orders classified in the superorder Afrotheria by molecular phylogeneticists3,4. The distribution of modern afrotheres, in combination with a basal position for Afrotheria within Placentalia and molecular divergence-time estimates, has been used to link placental diversification with the mid-Cretaceous separation of South America and Africa4. Morphological phylogenetic analyses do not support Afrotheria5,6,7 and the fossil record favours a northern origin of Placentalia8. Here we describe fossil postcrania that provide evidence for a close relationship between North American Palaeocene–Eocene apheliscine ‘hyopsodontid’ ‘condylarths’ (early ungulates or hoofed mammals) and extant Macroscelidea. Apheliscine postcranial morphology is consistent with a relationship to other ungulate-like afrotheres (Hyracoidea, Proboscidea) but does not provide support for a monophyletic Afrotheria. As the oldest record of an afrothere clade, identification of macroscelidean relatives in the North American Palaeocene argues against an African origin for Afrotheria, weakening support for linking placental diversification to the break-up of Gondwana.
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We would like to thank: T. M. Bown, for collection of the original Apheliscus skeletal association; D. Diveley, J. Meng (American Museum of Natural History), R. Emry, L. Gordon, H. Kafka, J. Mead, R. Purdy (USNM) and P. Houde (New Mexico State University) for access to specimens; D. M. Boyer for helpful discussions; J. C. Mussell for advice and comments on the manuscript; and D. B. Weishampel for access to equipment and facilities. We would also like to thank G. B. Rathbun for access to high quality images of sengis. Support of fieldwork leading to these discoveries has been provided by the US National Science Foundation and the Bureau of Land Management.
The authors declare that they have no competing financial interests.
This file contains: principal specimens examined; extant comparative material examined; modifications to existing character-taxon matrices; new character-taxon matrix; analyses performed and results (including Supplementary Figs 1-3); and Supplementary bibliography. (DOC 1736 kb)
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Zack, S., Penkrot, T., Bloch, J. et al. Affinities of ‘hyopsodontids’ to elephant shrews and a Holarctic origin of Afrotheria. Nature 434, 497–501 (2005). https://doi.org/10.1038/nature03351
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