Abstract: Summary Are complex, species-specific behaviors in animals reinforced by material reward alone or do they also induce positive emotions? Many adaptive human behaviors are intrinsically motivated: they not only improve our material outcomes, but improve our affect as well [1, 2, 3, 4, 5, 6, 7, 8]. Work to date on animal optimism, as an indicator of positive affect, has generally focused on how animals react to change in their circumstances, such as when their environment is enriched [9, 10, 11, 12, 13, 14] or they are manipulated by humans [15, 16, 17, 18, 19, 20, 21, 22, 23], rather than whether complex actions improve emotional state. Here, we show that wild New Caledonian crows are optimistic after tool use, a complex, species-specific behavior. We further demonstrate that this finding cannot be explained by the crows needing to put more effort into gaining food. Our findings therefore raise the possibility that intrinsic motivation (enjoyment) may be a fundamental proximate cause in the evolution of tool use and other complex behaviors. Video Abstract

Abstract: Despite numerous attempts to define animal tool use over the past four decades, the definition remains elusive and the behaviour classification somewhat subjective. Here, we provide a brief review of the definitions of animal tool use and show how those definitions have been modified over time. While some aspects have remained constant (i.e., the distinction between 'true' and 'borderline' tool use), others have been added (i.e., the distinction between 'dynamic' and 'static' behaviours). We present an updated, comprehensive catalog of documented animal tool use that indicates whether the behaviours observed included any 'true' tool use, whether the observations were limited to captive animals, whether tool manufacture has been observed, and whether the observed tool use was limited to only one individual and, thus, 'anecdotal' (i.e., N = 1). Such a catalog has not been attempted since Beck (1980). In addition to being a useful reference for behaviourists, this catalog demonstrates broad tool use and manufacture trends that may be of interest to phylogenists, evolutionary ecologists, and cognitive evolutionists. Tool use and tool manufacture are shown to be widespread across three phyla and seven classes of the animal kingdom. Moreover, there is complete overlap between the Aves and Mammalia orders in terms of the tool use categories (e.g., food extraction, food capture, agonism) arguing against any special abilities of mammals. The majority of tool users, almost 85% of the entries, use tools in only one of the tool use categories. Only members of the Passeriformes and Primates orders have been observed to use tools in four or more of the ten categories. Thus, observed tool use by some members of these two orders (e.g., Corvus, Papio) is qualitatively different from that of all other animal taxa. Finally, although there are similarities between Aves and Mammalia, and Primates and Passeriformes, primate tool use is qualitatively different. Approximately 35% of the entries for this order demonstrate a breadth of tool use (i.e., three or more categories by any one species) compared to other mammals (0%), Aves (2.4%), and the Passeriformes (3.1%). This greater breadth in tool use by some organisms may involve phylogenetic or cognitive differences � or may simply reflect differences in length and intensity of observations. The impact that tool usage may have had on groups' respective ecological niches and, through niche-construction, on their respective evolutionary trajectories remains a subject for future study.

Abstract: Chimpanzees, Pan troglodytes, are the most sophisticated tool-users among all nonhuman primates. From an evolutionary perspective, it is therefore puzzling that the tool use behaviour of their closest living primate relative, the bonobo, Pan paniscus, has been described as particularly poor. However, only a small number of bonobo groups have been studied in the wild and only over comparably short periods. Here, we show that captive bonobos and chimpanzees are equally diverse tool-users in most contexts. Our observations illustrate that tool use in bonobos can be highly complex and no different from what has been described for chimpanzees. The only major difference in the chimpanzee and bonobo data was that bonobos of all age–sex classes used tools in a play context, a possible manifestation of their neotenous nature. We also found that female bonobos displayed a larger range of tool use behaviours than males, a pattern previously described for chimpanzees but not for other great apes. Our results are consistent with the hypothesis that the female-biased tool use evolved prior to the split between bonobos and chimpanzees.

Abstract: Abstract
Several comparative research programs have focusedon the cognitive, life history and ecological traits thataccount for variation in brain size. We review one ofthese programs, a program that uses the reported frequencyof behavioral innovation as an operational measureof cognition. In both birds and primates, innovationrate is positively correlated with the relative size of associationareas in the brain, the hyperstriatum ventrale andneostriatum in birds and the isocortex and striatum inprimates. Innovation rate is also positively correlatedwith the taxonomic distribution of tool use, as well asinterspecific differences in learning. Some features ofcognition have thus evolved in a remarkably similar wayin primates and at least six phyletically-independent avianlineages. In birds, innovation rate is associated withthe ability of species to deal with seasonal changes in theenvironment and to establish themselves in new regions,and it also appears to be related to the rate atwhich lineages diversify. Innovation rate provides a usefultool to quantify inter-taxon differences in cognitionand to test classic hypotheses regarding the evolution ofthe brain.