Previous hypotheses for the origin and diversification of pinnipeds have followed a narrative approach based mostly on dispersalist (i.e., center of origin) explanations. Using an analytical approach, we present a testable hypothesis to explain the evolutionary biogeography of pinnipedimorphs (fur seals, sea lions, walruses, seals, and their fossil relatives) based on both dispersal and vicariant events in the context of a species-level phylogenetic framework. This integrated hypothesis considers many lines of evidence, including physical and ecologic factors controlling modern pinniped distributions, past geologic events related to opening and closing of seaways, paleoceanographic models, the improving pinniped fossil record, and pinniped phylogenetic analyses based on both morphologic and molecular data sets. Oceanic biogeographic regions and faunal provinces are defined and oceanic circulation patterns discussed with reference to the distribution of extant and fossil species. Paleobiogeographic hypotheses for each of the major pinniped lineages are presented using area cladograms and paleogeographic maps showing oceanographic and tectonic changes during successive intervals of the Cenozoic.
Our biogeographic hypothesis supports an eastern North Pacific origin for pinnipedimorphs during the late Oligocene coincident with initiation of glaciation in Antarctica. During the early Miocene, pinnipedimorphs remained restricted to the eastern North Pacific, where they began to diversify. Otariids (fur seals and sea lions) are first known from the late Miocene in the North Pacific, where they remained restricted until the late Pliocene. A transequatorial dispersal into the western South Pacific at this time preceded the rapid diversification of this group that occurred during the Pleistocene in the Southern Ocean. Odobenids (walruses) evolved in the North Pacific during the late early Miocene and underwent dramatic diversification in the late Miocene with later members of the odobenine lineage dispersing into the North Atlantic, most likely via an Arctic route. Extinct archaic phocoids, the desmatophocids, known only from the early to late Miocene, were confined to the eastern and western North Pacific. Phocids, although postulated here to have a North Pacific origin, are first known as fossils from the middle Miocene in the eastern and western North Atlantic region, as well as the Paratethys. Both monachine and phocine seals are distinct lineages beginning in the middle Miocene in the eastern and western provinces of the North Atlantic. During the late Miocene, phocids underwent a dramatic diversification. The early biogeographic history of phocine seals is centered in the Arctic and North Atlantic. Subsequent dispersal of phocines into the Paratethys and Pacific occurred during the Pleistocene. In contrast, monachine seals have a southern hemisphere center of diversity, especially the lobodontines of the Southern Ocean. Southern dispersal of this clade most likely occurred through the Neogene Central American Seaway prior to its closure in the mid-Pliocene. The pagophilic nature of extant phocine and lobodontine seals is largely a function of Pleistocene glacioeustatic events.