The axial skeleton of Tiktaalik roseae.

The axial columns of the earliest limbed vertebrates show distinct patterns of regionalization as compared to early tetrapodomorphs. Included among their novel features are sacral ribs, which provide linkage between the vertebral column and pelvis, contributing to body support and propulsion by the hindlimb. Data on the axial skeletons of the closest relatives of limbed vertebrates are sparce, with key features of specimens potentially covered by matrix. Therefore, it is unclear in what sequence and under what functional context specializations in the axial skeletons of tetrapods arose. Here, we describe the axial skeleton of the elpistostegalian Tiktaalik roseae and show that transformations to the axial column for head mobility, body support, and pelvic fin buttressing evolved in finned vertebrates prior to the origin of limbs. No atlas-axis complex is observed; however, an independent basioccipital-exoccipital complex suggests increased mobility at the occipital vertebral junction. While the construction of vertebrae in Tiktaalik is similar to early tetrapodomorphs, its ribs possess a specialized sacral domain. Sacral ribs are expanded and ventrally curved, indicating likely attachment to the expanded iliac blade of the pelvis by ligamentous connection. Thus, the origin of novel rib types preceded major alterations to trunk vertebrae, and linkage between pelvic fins and axial column preceded the origin of limbs. These data reveal an unexpected combination of post-cranial skeletal characters, informing hypotheses of body posture and movement in the closest relatives of limbed vertebrates.

An extinct north American porcupine with a South American tail.

New World porcupines (Erethizontinae) originated in South America and dispersed into North America as part of the Great American Biotic Interchange (GABI) 3-4 million years ago.1 Extant prehensile-tailed porcupines (Coendou) today live in tropical forests of Central and South America.2,3 In contrast, North American porcupines (Erethizon dorsatum) are thought to be ecologically adapted to higher-latitude temperate forests, with a larger body, shorter tail, and diet that includes bark.4,5,6,7 Limited fossils8,9,10,11,12,13 have hindered our understanding of the timing of this ecological differentiation relative to intercontinental dispersal during the GABI and expansion into temperate habitats.14,15,16,17,18 Here, we describe functionally important features of the skeleton of the extinct Erethizon poyeri, the oldest nearly complete porcupine skeleton documented from North America, found in the early Pleistocene of Florida. It differs from extant E. dorsatum in having a long, prehensile tail, grasping foot, and lacking dental specializations for bark gnawing, similar to tropical Coendou. Results from phylogenetic analysis suggest that the more arboreal characteristics found in E. poyeri are ancestral for erethizontines. Only after it expanded into temperate, Nearctic habitats did Erethizon acquire the characteristic features that it is known for today. When combined with molecular estimates of divergence times, results suggest that Erethizon was ecologically similar to a larger species of Coendou when it crossed the Isthmus of Panama by the early Pleistocene. It is likely that the range of this more tropically adapted form was limited to a continuous forested biome that extended from South America through the Gulf Coast.

A new tuskless walrus from the Miocene of Orange County, California, with comments on the diversity and taxonomy of odobenids.

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.