Although tool use may enhance resource utilization, its fitness benefits are difficult to measure. By examining longitudinal data from 196 radio-tagged southern sea otters (Enhydra lutris nereis), we found that tool-using individuals, particularly females, gained access to larger and/or harder-shelled prey. These mechanical advantages translated to reduced tooth damage during food processing. We also found that tool use diminishes trade-offs between access to different prey, tooth condition, and energy intake, all of which are dependent on the relative prey availability in the environment. Tool use allowed individuals to maintain energetic requirements through the processing of alternative prey that are typically inaccessible with biting alone, suggesting that this behavior is a necessity for the survival of some otters in environments where preferred prey are depleted. Description Editor’s summary It has long been known that sea otters sometimes use tools like shells and rocks to help them process their often thick-shelled mollusk prey. Law et al. show that animals who make regular use of such tools, especially females, are better able to consume a wide variety of prey, have higher energy consumption rates, and show reduced tooth wear (see the Perspective by Klump). The broadening of prey base facilitated by the use of tools was especially important in cases where the more easily processed prey that are preferred, such as abalone, were in short supply. —Sacha Vignieri

The notion that tool-use is unique to humans has long been refuted by the growing number of observations of animals using tools across various contexts. Yet, the mechanisms behind the emergence and sustenance of these tool-use repertoires are still heavily debated. We argue that the current animal behaviour literature is biased towards a social learning approach, in which animal, and in particular primate, tool-use repertoires are thought to require social learning mechanisms (copying variants of social learning are most often invoked). However, concrete evidence for a widespread dependency on social learning is still lacking. On the other hand, a growing body of observational and experimental data demonstrates that various animal species are capable of acquiring the forms of their tool-use behaviours via individual learning, with (non-copying) social learning regulating the frequencies of the behavioural forms within (and, indirectly, between) groups. As a first outline of the extent of the role of individual learning in animal tool-use, a literature review of reports of the spontaneous acquisition of animal tool-use behaviours was carried out across observational and experimental studies. The results of this review suggest that perhaps due to the pervasive focus on social learning in the literature, accounts of the individual learning of tool-use forms by naïve animals may have been largely overlooked, and their importance under-examined.

Sea otters are marine specialists but diet generalists, which feed primarily on benthic mega-invertebrates (i.e., body dimension >1 cm). They locate and capture epibenthic and infaunal prey with their forepaws by relying on vision and tactile sensitivity during short-duration dives (generally <2 min) in shallow waters (routine dives <30 m and maximum dive depth ~100 m) of the littoral zone. Sea otters have an elevated resting metabolic rate and small or no energy reserves in the form of blubber, so they feed every 3–4 h. Foraging dives often occur in bouts (i.e., two or more consecutive dives), which may last several hours with 1–2 min between dives, depending on the type of prey. Sea otters consume small or soft prey entirely or use their teeth or stone tools to access the flesh of mega-invertebrates with a shell, test, or exoskeleton. The daily percentage of time that sea otters devote to foraging depends on age, sex, presence of a pup, time of year, and prey abundance, which varies geographically, seasonally, and episodically. In areas occupied by sea otters for many years, epifaunal prey generally decline first followed by infaunal species, and this may result in greater foraging effort and diet specialization associated with density-dependent competition for food. Although prey availability strongly influences sea otter carrying capacity, both intrinsic and extrinsic factors influence population equilibrium density, resulting in spatiotemporal variations in foraging behavior.

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