RESUMEN
The development of pierce-feeding and loss of oral processing represented major adaptations for underwater feeding in marine mammals. We examined the evolution of pierce-feeding and its association with changes in tooth spacing and tooth size to determine whether pierce-feeding was practiced by the earliest known pinnipeds. Data on crown size and spacing in postcanine dentition were collected and 1) analysed by principal components analysis (PCA) to determine the tooth morphospace of arctoid carnivores, 2) analysed by least squares (LS) regression and phylogenetic independent contrasts (PIC) to determine what morphological variables were associated with increases in tooth spacing, and 3) used to reconstruct the evolution of feeding related traits within a phylogenetic context. The PCA analysis revealed that within arctoid carnivores, the greatest differences in morphospace were associated with pierce-feeding, and the early-diverging seal Enaliarctos was placed within the pinniped morphospace. Increased tooth spacing within Pinnipedia is a result of decreased postcanine crown size. When the evolution of dental characters is reconstructed, 'enaliarctines' were found to represent an intermediate stage in evolution between 'fissiped' and pinniped carnivores. They retained the limited tooth spacing of terrestrial carnivores, possessed postcanine crown lengths intermediate in size between pinnipeds and fissipeds, and possessed reduced heterodonty characteristic of crown pinnipeds. Our study indicated that pierce-feeding evolved early within pinnipeds. This suggested either that pierce-feeding evolved prior to the loss of mastication, or that pierce-feeding evolved at the same time as loss of mastication, and well before simplification of the dentition was completed.