New Fossil Giant Panda Relatives (Ailuropodinae, Ursidae): A Basal Lineage of Gigantic Mio-Pliocene Cursorial Carnivores

ABSTRACT Among the fossil members of the giant panda subfamily of ursid carnivorans, Ailuropodinae, one group of species is of giant size, those of Indarctos. Indarctos species have some dental resemblances to and may be closely related to Agriotherium, although there are other clear differences between these taxa, and no known species has definitive shared derived traits that could link these two genera. Here we describe a rich suite of fossil material from both North America and eastern Asia, all belonging to a new genus, Huracan, which possesses characters shared with both Agriotherium and Indarctos but also has diagnostic autapomorphies. The new taxon was distributed widely in the Holarctic during the latest Miocene, including at least four species: the type species Huracan schneideri (previously Agriotherium schneideri) from the latest Hemphillian (Hh4) and possibly early Blancan North American Land Mammal “Ages” (NALMAs), North America; H. coffeyi from the early Late Hemphillian (Hh3) NALMA, North America; H. qiui, sp. nov., from the Baodean Asian Land Mammal “Age” (ALMA), northern China; and H. roblesi from the MN13 zone (latest Miocene–earliest Pliocene) of Spain. Huracan is the nearest sister taxon to Agriotherium, the latter herein considered to be an ailuropodine (in the tribe Agriotheriini) rather than a hemicyonid, and the common ancestor of both genera evolved from Indarctos (with resultant paraphyly of that taxon) or another Indarctos-like ailuropodine bear, likely in eastern Asia. The dentitions of Huracan and Agriotherium both are more specialised for carnivory than most Indarctos species, indicating a radiation of diverse ecological carnivores earlier in the history of the later-diverging, highly specialized herbivores in the giant panda lineage. Their postcranial morphology suggests that species in both genera (Huracan and Agriotherium) were more cursorial than species assigned to Indarctos, and thus well adapted to more open habitats. These derived traits may explain the worldwide replacement of Indarctos species by Huracan and Agriotherium species during the latest Miocene, in response to significant global cooling and expansion of C4 grasslands that occurred at that time.