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 crow's exceptional tool skills.     

New Caledonian crows learn the functional properties of novel tool types

New Caledonian crows were presented with Bird and Emery's (2009a) Aesop's fable paradigm, which requires stones to be dropped into a water-filled tube to bring floating food within reach. The crows did not spontaneously use stones as tools, but quickly learned to do so, and to choose objects and materials with functional properties. Some crows discarded both inefficient and non-functional objects before observing their effects on the water level. Interestingly, the crows did not learn to discriminate between functional and non-functional objects and materials when there was an arbitrary, rather than causal, link between object and reward. This finding suggests that the crows' performances were not based on associative learning alone. That is, learning was not guided solely by the covariation rate between stimuli and outcomes or the conditioned reinforcement properties acquired by functional objects. Our results, therefore, show that New Caledonian crows can process causal information not only when it is linked to sticks and stick-like tools but also when it concerns the functional properties of novel types of tool.     

Does absolute brain size really predict self-control? Hand-tracking training improves performance on the A-not-B task

Large-scale, comparative cognition studies are set to revolutionize the way we investigate and understand the evolution of intelligence. However, the conclusions reached by such work have a key limitation: the cognitive tests themselves. If factors other than cognition can systematically affect the performance of a subset of animals on these tests, we risk drawing the wrong conclusions about how intelligence evolves. Here, we examined whether this is the case for the A-not-B task, recently used by MacLean and co-workers to study self-control among 36 different species. Non-primates performed poorly on this task; possibly because they have difficulty tracking the movements of a human demonstrator, and not because they lack self-control. To test this, we assessed the performance of New Caledonian crows on the A-not-B task before and after two types of training. New Caledonian crows trained to track rewards moved by a human demonstrator were more likely to pass the A-not-B test than birds trained on an unrelated choice task involving inhibitory control. Our findings demonstrate that overlooked task demands can affect performance on a cognitive task, and so bring into question MacLean''s conclusion that absolute brain size best predicts self-control.     

New Caledonian Crows Rapidly Solve a Collaborative Problem without Cooperative Cognition

There is growing comparative evidence that the cognitive bases of cooperation are not unique to humans. However, the selective pressures that lead to the evolution of these mechanisms remain unclear. Here we show that while tool-making New Caledonian crows can produce collaborative behavior, they do not understand the causality of cooperation nor show sensitivity to inequity. Instead, the collaborative behavior produced appears to have been underpinned by the transfer of prior experience. These results suggest that a number of possible selective pressures, including tool manufacture and mobbing behaviours, have not led to the evolution of cooperative cognition in this species. They show that causal cognition can evolve in a domain specific manner–understanding the properties and flexible uses of physical tools does not necessarily enable animals to grasp that a conspecific can be used as a social tool.     

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.     

New Caledonian crows plan for specific future tool use

The ability to plan for future events is one of the defining features of human intelligence. Whether non-human animals can plan for specific future situations remains contentious: despite a sustained research effort over the last two decades, there is still no consensus on this question. Here, we show that New Caledonian crows can use tools to plan for specific future events. Crows learned a temporal sequence where they were (a) shown a baited apparatus, (b) 5 min later given a choice of five objects and (c) 10 min later given access to the apparatus. At test, these crows were presented with one of two tool–apparatus combinations. For each combination, the crows chose the right tool for the right future task, while ignoring previously useful tools and a low-value food item. This study establishes that planning for specific future tool use can evolve via convergent evolution, given that corvids and humans shared a common ancestor over 300 million years ago, and offers a route to mapping the planning capacities of animals.     

On the evolutionary and ontogenetic origins of tool‐oriented behaviour in New Caledonian crows (Corvus moneduloides)

New Caledonian crows (Corvus moneduloides) are prolific tool users in captivity and in the wild, and have an inherited predisposition to express tool‐oriented behaviours. To further understand the evolution and development of tool use, we compared the development of object manipulation in New Caledonian crows and common ravens (Corvus corax), which do not routinely use tools. We found striking qualitative similarities in the ontogeny of tool‐oriented behaviour in New Caledonian crows and food‐caching behaviour in ravens. Given that the common ancestor of New Caledonian crows and ravens was almost certainly a caching species, we therefore propose that the basic action patterns for tool use in New Caledonian crows may have their evolutionary origins in caching behaviour. Noncombinatorial object manipulations had similar frequencies in the two species. However, frequencies of object combinations that are precursors to functional behaviour increased in New Caledonian crows and decreased in ravens throughout the study period, ending 6 weeks post‐fledging. These quantitative observations are consistent with the hypothesis that New Caledonian crows develop tool‐oriented behaviour because of an increased motivation to perform object combinations that facilitate the necessary learning. © 2011 The Linnean Society of London, Biological Journal of the Linnean Society, 2011, 102 , 870–877.     

An end to insight? New Caledonian crows can spontaneously solve problems without planning their actions

Animals rarely solve problems spontaneously. Some bird species, however, can immediately find a solution to the string-pulling problem. They are able to rapidly gain access to food hung on the end of a long string by repeatedly pulling and then stepping on the string. It is currently unclear whether these spontaneous solutions are produced by insight or by a perceptual-motor feedback loop. Here, we presented New Caledonian crows and humans with a novel horizontal string-pulling task. While the humans succeeded, no individual crow showed a significant preference for the connected string, and all but one failed to gain the food even once. These results clearly show that string pulling in New Caledonian crows is generated not by insight, but by perceptual feedback. Animals can spontaneously solve problems without planning their actions.     

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.     

Hook tool manufacture in New Caledonian crows: behavioural variation and the influence of raw materials

Background

New Caledonian crows use a range of foraging tools, and are the only non-human species known to craft hooks. Based on a small number of observations, their manufacture of hooked stick tools has previously been described as a complex, multi-stage process. Tool behaviour is shaped by genetic predispositions, individual and social learning, and/or ecological influences, but disentangling the relative contributions of these factors remains a major research challenge. The properties of raw materials are an obvious, but largely overlooked, source of variation in tool-manufacture behaviour. We conducted experiments with wild-caught New Caledonian crows, to assess variation in their hooked stick tool making, and to investigate how raw-material properties affect the manufacture process.

Results

In Experiment 1, we showed that New Caledonian crows’ manufacture of hooked stick tools can be much more variable than previously thought (85 tools by 18 subjects), and can involve two newly-discovered behaviours: ‘pulling’ for detaching stems and bending of the tool shaft. Crows’ tool manufactures varied significantly: in the number of different action types employed; in the time spent processing the hook and bending the tool shaft; and in the structure of processing sequences. In Experiment 2, we examined the interaction of crows with raw materials of different properties, using a novel paradigm that enabled us to determine subjects’ rank-ordered preferences (42 tools by 7 subjects). Plant properties influenced: the order in which crows selected stems; whether a hooked tool was manufactured; the time required to release a basic tool; and, possibly, the release technique, the number of behavioural actions, and aspects of processing behaviour. Results from Experiment 2 suggested that at least part of the natural behavioural variation observed in Experiment 1 is due to the effect of raw-material properties.

Conclusions

Our discovery of novel manufacture behaviours indicates a plausible scenario for the evolutionary origins, and gradual refinement, of New Caledonian crows’ hooked stick tool making. Furthermore, our experimental demonstration of a link between raw-material properties and aspects of tool manufacture provides an alternative hypothesis for explaining regional differences in tool behaviours observed in New Caledonian crows, and some primate species.

     

Cognitive Processes Associated with Sequential Tool Use in New Caledonian Crows

Background

Using tools to act on non-food objects—for example, to make other tools—is considered to be a hallmark of human intelligence, and may have been a crucial step in our evolution. One form of this behaviour, ‘sequential tool use’, has been observed in a number of non-human primates and even in one bird, the New Caledonian crow (Corvus moneduloides). While sequential tool use has often been interpreted as evidence for advanced cognitive abilities, such as planning and analogical reasoning, the behaviour itself can be underpinned by a range of different cognitive mechanisms, which have never been explicitly examined. Here, we present experiments that not only demonstrate new tool-using capabilities in New Caledonian crows, but allow examination of the extent to which crows understand the physical interactions involved.

Methodology/Principal Findings

In two experiments, we tested seven captive New Caledonian crows in six tasks requiring the use of up to three different tools in a sequence to retrieve food. Our study incorporated several novel features: (i) we tested crows on a three-tool problem (subjects were required to use a tool to retrieve a second tool, then use the second tool to retrieve a third one, and finally use the third one to reach for food); (ii) we presented tasks of different complexity in random rather than progressive order; (iii) we included a number of control conditions to test whether tool retrieval was goal-directed; and (iv) we manipulated the subjects'' pre-testing experience. Five subjects successfully used tools in a sequence (four from their first trial), and four subjects repeatedly solved the three-tool condition. Sequential tool use did not require, but was enhanced by, pre-training on each element in the sequence (‘chaining’), an explanation that could not be ruled out in earlier studies. By analyzing tool choice, tool swapping and improvement over time, we show that successful subjects did not use a random probing strategy. However, we find no firm evidence to support previous claims that sequential tool use demonstrates analogical reasoning or human-like planning.

Conclusions/Significance

While the ability of subjects to use three tools in sequence reveals a competence beyond that observed in any other species, our study also emphasises the importance of parsimony in comparative cognitive science: seemingly intelligent behaviour can be achieved without the involvement of high-level mental faculties, and detailed analyses are necessary before accepting claims for complex cognitive abilities.     

Tentative evidence for inequity aversion to unequal work-effort but not to unequal reward distribution in Goffin's cockatoos

One hallmark in the evolution of cooperation is the ability to evaluate one's own payoff for a task against that of another person. To trace its evolutionary history, there has recently been a surge in comparative studies across different species. In non-human animals, evidence of inequity aversion has so far been identified in several primate species, dogs, and rats. Research in birds revealed mixed findings so far: among corvids, crows and ravens did react sensitively to unequal payoffs and work-effort, while New Caledonian crows did not. Among psittacids, kea were studied so far: Yet, despite the fact that they live in large, hierarchically organized social groups that show complex interactions, they did not show a significant reaction to inequitable payoffs. Here we tested for the first time a Cacatua, the Goffin's cockatoo, using a standardized token exchange paradigm in which first the partner and then the subject could exchange a token for a food reward. Our results show that subjects did not react to unequal reward distributions. However, in comparison to the Equity Condition, the likelihood to exchange was lower in the condition in which the partner received the same reward as a gift (without having to work for it) whereas the subject had to perform a task involving substantial work-effort, suggesting that the Goffin's cockatoos do react aversively to work-effort inequity. In a follow-up experiment, subjects never received a reward but observed a conspecific receive a high-quality reward depending on condition. We found again no evidence for an aversion for the unequal reward distribution, but only that, independent of condition, subjects quickly lost their motivation to participate due to not receiving a reward. In summary, Goffins showed some sensitivity to increased unequal work-effort, but did not react to unequal reward distribution.     

Context-dependent tool use in New Caledonian crows

Humans and chimpanzees both exhibit context-dependent tool use. That is, both species choose to use tools when food is within reach, but the context is potentially hazardous. Here, we show that New Caledonian crows used tools more frequently when food was positioned next to a novel model snake than when food was positioned next to a novel teddy bear or a familiar food bowl. However, the crows showed no significant difference in their neophobic reactions towards the teddy bear and the model snake. Therefore, the crows used tools more in response to a risky object resembling a natural predator than to a less-threatening object that provoked a comparable level of neophobia. These results show that New Caledonian crows, like humans and chimpanzees, are capable of context-dependent tool use.     

Rooks perceive support relations similar to six-month-old babies

Some corvids have demonstrated cognitive abilities that rival or exceed those of the great apes; for example, tool use in New Caledonian crows, and social cognition, episodic-like memory and future planning in Western scrub-jays. Rooks appear to be able to solve novel tasks through causal reasoning rather than simple trial-and-error learning. Animals with certain expectations about how objects interact would be able to narrow the field of candidate causes substantially, because some causes are simply ‘impossible’. Here we present evidence that rooks hold such expectations and appear to possess perceptual understanding of support relations similar to that demonstrated by human babies, which is more comprehensive than that of chimpanzees.     

Animal cognition: crows spontaneously solve a metatool task

A recent study found that tool-manufacturing New Caledonian crows spontaneously solved a metatool task in which the birds used a tool to obtain a second, longer tool that could then be used to obtain food that was otherwise out of reach.     

Of babies and birds: complex tool behaviours are not sufficient for the evolution of the ability to create a novel causal intervention

Humans are capable of simply observing a correlation between cause and effect, and then producing a novel behavioural pattern in order to recreate the same outcome. However, it is unclear how the ability to create such causal interventions evolved. Here, we show that while 24-month-old children can produce an effective, novel action after observing a correlation, tool-making New Caledonian crows cannot. These results suggest that complex tool behaviours are not sufficient for the evolution of this ability, and that causal interventions can be cognitively and evolutionarily disassociated from other types of causal understanding.     

Parallel tool industries in New Caledonian crows

Individual specialization in the use of foraging tools occurs in hunter-gatherer societies but is absent in non-human primate tool use. 'Parallel tool industries' in hunter-gatherers are mainly based on strict sexual division of labour that is highly reliant on social conformity. Here, we show that 12 individuals in a population of New Caledonian crows on Maré Island had strong preferences for either stick tools or pandanus tools. Eight of the 12 crows had exclusive preferences. The individual specialization that we found is probably associated with different foraging niches. However, in spite of sexual size dimorphism there was no significant association between the sex of crows and their tool preferences. Our findings demonstrate that highly organized, strict sexual division of labour is not a necessary prerequisite for the evolution of parallel tool industries.     

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.     

Flexibility in problem solving and tool use of kea and New Caledonian crows in a multi access box paradigm

Parrots and corvids show outstanding innovative and flexible behaviour. In particular, kea and New Caledonian crows are often singled out as being exceptionally sophisticated in physical cognition, so that comparing them in this respect is particularly interesting. However, comparing cognitive mechanisms among species requires consideration of non-cognitive behavioural propensities and morphological characteristics evolved from different ancestry and adapted to fit different ecological niches. We used a novel experimental approach based on a Multi-Access-Box (MAB). Food could be extracted by four different techniques, two of them involving tools. Initially all four options were available to the subjects. Once they reached criterion for mastering one option, this task was blocked, until the subjects became proficient in another solution. The exploratory behaviour differed considerably. Only one (of six) kea and one (of five) NCC mastered all four options, including a first report of innovative stick tool use in kea. The crows were more efficient in using the stick tool, the kea the ball tool. The kea were haptically more explorative than the NCC, discovered two or three solutions within the first ten trials (against a mean of 0.75 discoveries by the crows) and switched more quickly to new solutions when the previous one was blocked. Differences in exploration technique, neophobia and object manipulation are likely to explain differential performance across the set of tasks. Our study further underlines the need to use a diversity of tasks when comparing cognitive traits between members of different species. Extension of a similar method to other taxa could help developing a comparative cognition research program.     

Vice-anvil use in nut processing by two Corvus species

Woodpeckers and certain passerine species secure encased food in the environment in various ways to facilitate the extraction of the contents with their bills. They do this by securing the food items in locations such as crevices and holes, newly defined in this paper as ‘vice-anvils’. Here I report that free-living New Caledonian crows (Corvus moneduloides) and rooks (Corvus frugilegus, in New Zealand) also use vice-anvils to process candlenuts and walnuts, respectively. New Caledonian crows placed candlenut sections in vice-anvils to aid kernel extraction, after the candlenuts had been dropped onto an anvil to break them open. In contrast, rooks used vice-anvils to secure walnuts while they broke the shell with their bills. Long-term use by rooks of a vice-anvil in a tree had produced a ‘purpose-made’ nut-cracking site. My findings extend the persistent use of specific vice-anvils to Corvus species and further demonstrate their innovative and flexible foraging behaviour.