Stable isotope evidence for resource partitioning in extinct marine carnivores

Abstract

Living pinnipeds play essential roles in marine ecosystems and display divergent lineage-specific foraging and habitat preferences. The origin of these ecological strategies remains unclear. We analyzed original (n = 22) and published (n = 93) stable carbon (δ13C) and oxygen (δ18O) isotope compositions of tooth enamel from fossil pinnipeds and coeval marine and terrestrial mammals from the late Early Miocene to early Middle Miocene Temblor Formation, the early Middle Miocene Round Mountain Silt Formation at Sharktooth Hill Bonebed, the Middle Miocene of lower levels of the Monterey Formation, and the late Middle Miocene Santa Margarita Formation from the eastern North Pacific Ocean; and original (n = 43) δ13C and δ18O values from the Early Pliocene Yorktown Formation in the western North Atlantic. As expected for aquatic mammals, pinnipeds and control marine mammals had low δ18O variability relative to coeval terrestrial mammals, indicating minimal diagenetic alteration. Among late Early Miocene and Middle Miocene pinnipeds, Allodesmus had lower δ13C values than coeval taxa indicating offshore foraging, in agreement with predictions based on skeletal morphology. However, Allodesmus from the late Middle Miocene Santa Margarita Formation had overlapping δ13C values with co-occurring cetaceans and pinnipeds suggesting preferential nearshore habitat. The stem otariid Pithanotaria had higher enamel δ13C values than co-occurring pinnipeds, indicating nearshore foraging. The stem odobenid Neotherium had intermediate δ13C values supporting foraging between nearshore and offshore predators. Conversely, the stem odobenid cf. Imagotaria had lower enamel δ18O but comparable δ13C values to coexisting pinnipeds, suggesting the exploitation of estuarine resources. At least two ecologically distinct pinniped groups also occurred in the Yorktown Formation. In this unit, the odobenid Ontocetus and some monachinae phocids were predominantly nearshore foragers, with higher enamel δ13C and δ18O values than simultaneously occurring marine mammals. The second group was exclusively composed of phocids, which had comparatively low δ13C and δ18O values, compatible with northward foraging movements along the western North Atlantic coast. The paleoecological profiles of fossils bear strong similarities to the life modes of extant pinniped communities, implying that extant foraging modes appeared early in pinniped evolution and that resource partitioning contributed to their community structure over time.