Mental template matching is a potential cultural transmission mechanism for New Caledonian crow tool manufacturing traditions

Cumulative cultural evolution occurs when social traditions accumulate improvements over time. In humans cumulative cultural evolution is thought to depend on a unique suite of cognitive abilities, including teaching, language and imitation. Tool-making New Caledonian crows show some hallmarks of cumulative culture; but this claim is contentious, in part because these birds do not appear to imitate. One alternative hypothesis is that crows' tool designs could be culturally transmitted through a process of mental template matching. That is, individuals could use or observe conspecifics' tools, form a mental template of a particular tool design, and then reproduce this in their own manufacture - a process analogous to birdsong learning. Here, we provide the first evidence supporting this hypothesis, by demonstrating that New Caledonian crows have the cognitive capacity for mental template matching. Using a novel manufacture paradigm, crows were first trained to drop paper into a vending machine to retrieve rewards. They later learnt that only items of a particular size (large or small templates) were rewarded. At test, despite being rewarded at random, and with no physical templates present, crows manufactured items that were more similar in size to previously rewarded, than unrewarded, templates. Our results provide the first evidence that this cognitive ability may underpin the transmission of New Caledonian crows' natural tool designs.Tis study was supported by the NZ Marsden fund (RDG), a Rutherford Discovery Fellowship and the 2015 Prime Ministers MacDiarmid Emerging Scientist prize (AHT). SAJ thanks the ERC (Grant Agreement No. 3399933) for funding

Modifications to the Aesop's Fable paradigm change New Caledonian crow performances

While humans are able to understand much about causality, it is unclear to what extent non-human animals can do the same. The Aesop's Fable paradigm requires an animal to drop stones into a water-filled tube to bring a floating food reward within reach. Rook, Eurasian jay, and New Caledonian crow performances are similar to those of children under seven years of age when solving this task. However, we know very little about the cognition underpinning these birds' performances. Here, we address several limitations of previous Aesop's Fable studies to gain insight into the causal cognition of New Caledonian crows. Our results provide the first evidence that any non-human animal can solve the U-tube task and can discriminate between water-filled tubes of different volumes. However, our results do not provide support for the hypothesis that these crows can infer the presence of a hidden causal mechanism. They also call into question previous object-discrimination performances. The methodologies outlined here should allow for more powerful comparisons between humans and other animal species and thus help us to determine which aspects of causal cognition are distinct to humans.Publisher PDFPeer reviewe

Using the Aesop's fable paradigm to investigate causal understanding of water displacement by New Caledonian crows.

Understanding causal regularities in the world is a key feature of human cognition. However, the extent to which non-human animals are capable of causal understanding is not well understood. Here, we used the Aesop's fable paradigm--in which subjects drop stones into water to raise the water level and obtain an out of reach reward--to assess New Caledonian crows' causal understanding of water displacement. We found that crows preferentially dropped stones into a water-filled tube instead of a sand-filled tube; they dropped sinking objects rather than floating objects; solid objects rather than hollow objects, and they dropped objects into a tube with a high water level rather than a low one. However, they failed two more challenging tasks which required them to attend to the width of the tube, and to counter-intuitive causal cues in a U-shaped apparatus. Our results indicate that New Caledonian crows possess a sophisticated, but incomplete, understanding of the causal properties of displacement, rivalling that of 5-7 year old children

Decision-making flexibility in New Caledonian crows, young children and adult humans in a multi-dimensional tool-use task.

The ability to make profitable decisions in natural foraging contexts may be influenced by an additional requirement of tool-use, due to increased levels of relational complexity and additional work-effort imposed by tool-use, compared with simply choosing between an immediate and delayed food item. We examined the flexibility for making the most profitable decisions in a multi-dimensional tool-use task, involving different apparatuses, tools and rewards of varying quality, in 3-5-year-old children, adult humans and tool-making New Caledonian crows (Corvus moneduloides). We also compared our results to previous studies on habitually tool-making orangutans (Pongo abelii) and non-tool-making Goffin's cockatoos (Cacatua goffiniana). Adult humans, cockatoos and crows, but not children and orangutans, did not select a tool when it was not necessary, which was the more profitable choice in this situation. Adult humans, orangutans and cockatoos, but not crows and children, were able to refrain from selecting non-functional tools. By contrast, the birds, but not the primates tested, struggled to attend to multiple variables-where two apparatuses, two tools and two reward qualities were presented simultaneously-without extended experience. These findings indicate: (1) in a similar manner to humans and orangutans, New Caledonian crows and Goffin's cockatoos can flexibly make profitable decisions in some decision-making tool-use tasks, though the birds may struggle when tasks become more complex; (2) children and orangutans may have a bias to use tools in situations where adults and other tool-making species do not

Self‐control in crows, parrots and nonhuman primates

Self‐control is critical for both humans and nonhuman animals because it underlies complex cognitive abilities, such as decision‐making and future planning, enabling goal‐directed behavior. For instance, it is positively associated with social competence and life success measures in humans. We present the first review of delay of gratification as a measure of self‐control in nonhuman primates, corvids (crow family) and psittacines (parrot order): disparate groups that show comparable advanced cognitive abilities and similar socio‐ecological factors. We compare delay of gratification performance and identify key issues and outstanding areas for future research, including finding the best measures and drivers of delayed gratification. Our review therefore contributes to our understanding of both delayed gratification as a measure of self‐control and of complex cognition in animals

Performance in Object-Choice Aesop's Fable Tasks Are Influenced by Object Biases in New Caledonian Crows but not in Human Children

The ability to reason about causality underlies key aspects of human cognition, but the extent to which non-humans understand causality is still largely unknown. The Aesop's Fable paradigm, where objects are inserted into water-filled tubes to obtain out-of-reach rewards, has been used to test casual reasoning in birds and children. However, success on these tasks may be influenced by other factors, specifically, object preferences present prior to testing or arising during pre-test stone-dropping training. Here, we assessed this 'object-bias' hypothesis by giving New Caledonian crows and 5-10 year old children two object-choice Aesop's Fable experiments: sinking vs. floating objects, and solid vs. hollow objects. Before each test, we assessed subjects' object preferences and/or trained them to prefer the alternative object. Both crows and children showed pre-test object preferences, suggesting that birds in previous Aesop's Fable studies may also have had initial preferences for objects that proved to be functional on test. After training to prefer the non-functional object, crows, but not children, performed more poorly on these two object-choice Aesop's Fable tasks than subjects in previous studies. Crows dropped the non-functional objects into the tube on their first trials, indicating that, unlike many children, they do not appear to have an a priori understanding of water displacement. Alternatively, issues with inhibition could explain their performance. The crows did, however, learn to solve the tasks over time. We tested crows further to determine whether their eventual success was based on learning about the functional properties of the objects, or associating dropping the functional object with reward. Crows inserted significantly more rewarded, non-functional objects than non-rewarded, functional objects. These findings suggest that the ability of New Caledonian crows to produce performances rivaling those of young children on object-choice Aesop's Fable tasks is partly due to pre-existing object preferences.This research was funded by the European Research Council under the European Union's Seventh Framewor k Programme (FP7/ 2007-2013)/ERC Grant Agreement No. 3399933, awarded to NSC (funding RM, SAJ, EL & NSC). AHT was funded by a Rutherford Discovery Fellowship from the Royal Society of New Zealand

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