We describe Titanotaria orangensis (gen. et. sp. nov.), a new species of walrus (odobenid) from the upper Miocene Oso Member of the Capistrano Formation of Orange County, California. This species is important because: (1) It is one of the best-known and latest-surviving tuskless walruses; (2) It raises the number of reported odobenid taxa from the Oso Member to four species making it one of the richest walrus assemblages known (along with the basal Purisima of Northern California); (3) It is just the second record of a tuskless walrus from the same unit as a tusked taxon. Our phylogenetic analysis places T. orangensis as sister to a clade that includes Imagotaria downsi, Pontolis magnus, Dusignathus spp., Gomphotaria pugnax, and Odobeninae. We propose new branch-based phylogenetic definitions for Odobenidae, Odobeninae, and a new node-based name (Neodobenia) for the clade that includes Dusignathus spp., G. pugnax, and Odobeninae. A richness analysis at the 0.1 Ma level that incorporates stratigraphic uncertainty and ghost lineages demonstrates maximum peaks of richness (up to eight or nine coeval lineages) near the base of Odobenidae, Neodobenia, and Odobenini. A more conservative minimum curve demonstrates that standing richness may have been much lower than the maximum lineage richness estimates that are biased by stratigraphic uncertainty. Overall the odobenid fossil record is uneven, with large time slices of the record missing on either side of the Pacific Ocean at some times and biases from the preserved depositional environments at other times. We recognize a provisional timescale for the transition of East Pacific odobenid assemblages that include “basal odobenids” (stem neodobenians) from the Empire and older formations (>7 Ma), to a mixture of basal odobenids and neodobenians from the Capistrano and basal Purisima (7–5 Ma), and then just neodobenians from all younger units (<5 Ma). The large amount of undescribed material will add new taxa and range extensions for existing taxa, which will likely change some of the patterns we describe.

The origin of quadrupedal locomotion in tetrapods entailed the evolution of a regionalized axial skeleton with sacral ribs. These ribs provide linkage between the pelvis and vertebral column and contribute to body support and propulsion by the hind limb. The closest relatives of limbed vertebrates are not known to possess such a connection and, therefore, have typically been described as primarily supporting their bodies against the substrate with pectoral fins. However, data on the axial skeletons of stem tetrapods are sparce, with key features of specimens potentially covered by matrix. Here we provide micro-computed tomography data of the axial skeleton of Tiktaalik roseae and show that its vertebrae and ribs are regionalized along the craniocaudal axis, including expanded and ventrally curved ribs in the sacral region. The sacral ribs would have contacted the expanded iliac blade of the pelvis in a soft tissue connection. No atlas-axis complex is observed, however the basioccipital-exoccipital complex is deconsolidated from the rest of the neurocranium, suggesting increased mobility at occipital-vertebral junction. Thus, axial regionalization that allowed for innovations in head mobility, body support and buttressing the pelvic fin evolved prior to the origin of limbs.

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