Gavin R. Hunt

Diversification and cumulative evolution in New Caledonian crow tool manufacture.

Many animals use tools but only humans are generally considered to have the cognitive sophistication required for cumulative technological evolution. Three important characteristics of cumulative technological evolution are: (i) the diversification of tool design; (ii) cumulative change; and (iii) high–fidelity social transmission. We present evidence that crows have diversified and cumulatively changed the design of their pandanus tools. In 2000 we carried out an intensive survey in New Caledonia to establish the geographical variation in the manufacture of these tools. We documented the shapes of 5550 tools from 21 sites throughout the range of pandanus tool manufacture. We found three distinct pandanus tool designs: wide tools, narrow tools and stepped tools. The lack of ecological correlates of the three tool designs and their different, continuous and overlapping geographical distributions make it unlikely that they evolved independently. The similarities in the manufacture method of each design further suggest that pandanus tools have gone through a process of cumulative change from a common historical origin. We propose a plausible scenario for this rudimentary cumulative evolution.

Spontaneous metatool use by New Caledonian Crows.

A crucial stage in hominin evolution was the development of metatool use -- the ability to use one tool on another [1, 2]. Although the great apes can solve metatool tasks [3, 4], monkeys have been less successful [5-7]. Here we provide experimental evidence that New Caledonian crows can spontaneously solve a demanding metatool task in which a short tool is used to extract a longer tool that can then be used to obtain meat. Six out of the seven crows initially attempted to extract the long tool with the short tool. Four successfully obtained meat on the first trial. The experiments revealed that the crows did not solve the metatool task by trial-and-error learning during the task or through a previously learned rule. The sophisticated physical cognition shown appears to have been based on analogical reasoning. The ability to reason analogically may explain the exceptional tool-manufacturing skills of New Caledonian crows.

Do new caledonian crows solve physical problems through causal reasoning

The extent to which animals other than humans can reason about physical problems is contentious. The benchmark test for this ability has been the trap-tube task. We presented New Caledonian crows with a series of two-trap versions of this problem. Three out of six crows solved the initial trap-tube. These crows continued to avoid the trap when the arbitrary features that had previously been associated with successful performances were removed. However, they did not avoid the trap when a hole and a functional trap were in the tube. In contrast to a recent primate study, the three crows then solved a causally equivalent but visually distinct problem—the trap-table task. The performance of the three crows across the four transfers made explanations based on chance, associative learning, visual and tactile generalization, and previous dispositions unlikely. Our findings suggest that New Caledonian crows can solve complex physical problems by reasoning both causally and analogically about causal relations. Causal and analogical reasoning may form the basis of the New Caledonian crows exceptional tool skills.

Complex cognition and behavioural innovation in New Caledonian crows

Apes, corvids and parrots all show high rates of behavioural innovation in the wild. However, it is unclear whether this innovative behaviour is underpinned by cognition more complex than simple learning mechanisms. To investigate this question we presented New Caledonian crows with a novel three-stage metatool problem. The task involved three distinct stages: (i) obtaining a short stick by pulling up a string, (ii) using the short stick as a metatool to extract a long stick from a toolbox, and finally (iii) using the long stick to extract food from a hole. Crows with previous experience of the behaviours in stages 1–3 linked them into a novel sequence to solve the problem on the first trial. Crows with experience of only using string and tools to access food also successfully solved the problem. This innovative use of established behaviours in novel contexts was not based on resurgence, chaining and conditional reinforcement. Instead, the performance was consistent with the transfer of an abstract, causal rule: ‘out-of-reach objects can be accessed using a tool’. This suggests that high innovation rates in the wild may reflect complex cognitive abilities that supplement basic learning mechanisms.

Direct observations of pandanus-tool manufacture and use by a New Caledonian crow (Corvus moneduloides)

New Caledonian crows are reported to have impressive pandanus-tool manufacture abilities. These claims are based on an extensive artefact record. However, inferring behavioural and cognitive abilities without direct observation of tool manufacture is problematic. Here we report (and document on video) direct observations of a crow making and using stepped pandanus tools at Pic Ningua. We observed (1) a bias for making tools on left edges consistent with that previously found at the site, (2) faithful manufacture of a stepped design with high overall congruence in the shapes of tools, (3) the use of convergent rips to first form the tapered end working away from the trunk then the wide end working towards the trunk, (4) appropriate functional use of stepped tools by use of the leaf-edge barbs to hook food from holes, and (5) consistent holding of tools on the left side of its head when using them. Our observations verify most of the claims based on the artefact record, but the crow’s exact manufacture technique was slightly different to that inferred previously.

Tool Use by the New Caledonian Crow Corvus moneduloides to Obtain Cerambycidae from Dead Wood

Summary From a successful effort in late 1997 to film crow tool-use with the BBC Natural History Unit, I present the first detailed description of New Caledonian Crows Corvus moneduloides using tools to extract larvae of an endemic Cerambycidae:Prioninae from dead wood. I observed birds over a six-week period at Sarraméa where fallen Aleurites moluccana logs were ‘salted’ with the larvae. Crows habitually used mostly A. moluccana leaf-stems to extract larvae from holes and manufacture was also common when birds removed leaves from leaf-stems before using tools. The techniques of adult crows varied but were more proficient than those of a juvenile crow that frequented the site. Tool-use to extract Cerambycidae from dead wood was probably an important part of the foraging behaviour of the crows year-round. My observations here and elsewhere show that crows in different localities can have distinctly different traditions in their tool behaviour.

An investigation into the cognition behind spontaneous string pulling in New Caledonian crows

The ability of some bird species to pull up meat hung on a string is a famous example of spontaneous animal problem solving. The “insight” hypothesis claims that this complex behaviour is based on cognitive abilities such as mental scenario building and imagination. An operant conditioning account, in contrast, would claim that this spontaneity is due to each action in string pulling being reinforced by the meat moving closer and remaining closer to the bird on the perch. We presented experienced and naïve New Caledonian crows with a novel, visually restricted string-pulling problem that reduced the quality of visual feedback during string pulling. Experienced crows solved this problem with reduced efficiency and increased errors compared to their performance in standard string pulling. Naïve crows either failed or solved the problem by trial and error learning. However, when visual feedback was available via a mirror mounted next to the apparatus, two naïve crows were able to perform at the same level as the experienced group. Our results raise the possibility that spontaneous string pulling in New Caledonian crows may not be based on insight but on operant conditioning mediated by a perceptual-motor feedback cycle.

Laterality in tool manufacture by crows

New Caledonian crows (Fig. 1) fashion tapered tools from either the left or the right edge of the long narrow leaves of pandanus trees or screw pines, which they use to extract invertebrates in rainforest vegetation. Although right-handedness is thought to be uniquely human, we show here that crows from different localities display a widespread laterality in making their tools, indicating that this behaviour is unlikely to be attributable to local social traditions or ecological factors. To our knowledge, this is the first demonstration of species-level laterality in manipulatory skills outside humans.

Social learning in New Caledonian crows

New Caledonian (NC) crows are the most sophisticated tool manufacturers other than humans. The diversification and geographical distribution of their three Pandanus tool designs that differ in complexity, as well as the lack of ecological correlates, suggest that cumulative technological change has taken place. To investigate the possibility that high-fidelity social transmission mediated this putative ratchet-like process, we studied the ontogeny of Pandanus tool manufacture and social organization in free-living NC crows. We found that juvenile crows took more than 1 year to reach adult proficiency in their Pandanus tool skills. Although trial-and-error learning is clearly important, juveniles have ample opportunity to learn about Pandanus tool manufacture by both observing their parents and interacting with artifactual material. The crows’ social system seems likely to promote the faithful social transmission of local tool designs by both favoring the vertical transmission of tool information and minimizing horizontal transmission. We suggest that NC crows develop their Pandanus tool skills in a highly scaffolded learning environment that facilitates the cumulative technological evolution of tool designs.

Tool-making New Caledonian crows have large associative brain areas.

Animals with a high rate of innovative and associative-based behavior usually have large brains. New Caledonian (NC) crows stand out due to their tool manufacture, their generalized problem-solving abilities and an extremely high degree of encephalization. It is generally assumed that this increased brain size is due to the ability to process, associate and memorize diverse stimuli, thereby enhancing the propensity to invent new and complex behaviors in adaptive ways. However, this premise lacks firm empirical support since encephalization could also result from an increase of only perceptual and/or motor areas. Here, we compared the brain structures of NC crows with those of carrion crows, jays and sparrows. The brains of NC crows were characterized by a relatively large mesopallium, striatopallidal complex, septum and tegmentum. These structures mostly deal with association and motor-learning. This supports the hypothesis that the evolution of innovative or complex behavior requires a brain composition that increases the ability to associate and memorize diverse stimuli in order to execute complex motor output. Since apes show a similar correlation of cerebral growth and cognitive abilities, the evolution of advanced cognitive skills appears to have evolved independently in birds and mammals but with a similar neural orchestration.