The evolution of cost ef®cient swimming in marine mammals: limits to energetic optimization

Mammals re-entered the oceans less than 60 million years ago. The transition from a terrestrial to an aquatic lifestyle required extreme morphological and behavioural modi¢cations concomitant with fundamentally di¡erent locomotor mechanisms for moving on land and through water. Energetic transport costs typically re£ect such di¡erent locomotor modes, but can not be discerned from the fossil record. In this study the energetic challenges associated with changing from terrestrial to aquatic locomotion in primitive marine mammals are examined by comparing the transport, maintenance and locomotor costs of extant mammals varying in degree of aquatic specialization. The results indicate that running and swimming specialists have converged on an energetic optimum for locomotion. An allometric expression, COTTOTˆ7.79 massÿ0:29 (r 2ˆ0.83, nˆ6 species), describes the total cost of transport in J kgÿ1 mÿ1 for swimming marine mammals ranging in size from 21kg to 15 000 kg. This relation is indistinguishable from that describing total transport costs in running mammals. In contrast, the transitional lifestyle of semi-aquatic mammals, similar to that of ancestral marine mammals, incurs costs that are 2.4^5.1 times higher than locomotor specialists. These patterns suggest that primitive marine mammals confronted an energetic hurdle before returning to costs reminiscent of their terrestrial ancestry, and may have reached an evolutionary limit for energetic optimization during swimming.