Dunking rusk: innovative food soaking behaviour in Goffin's cockatoos (Cacatua goffiniana).

Foraging innovations in animals involving the processing of resources that are already edible in an unprocessed state, yet of improved quality in a processed state, are rare but important to study the evolution of food preparation. Here, we present the first scientific report of food dunking behaviours in parrots by Goffin's cockatoos, a model species for innovative problem solving. Observations during lunch showed seven out of 18 cockatoos placing their food into water and soaking it prior to consumption. This was largely done with dry rusk which was eaten almost exclusively when dunked. Furthermore, their transport effort and waiting times before retrieving food from the water indicate their willingness to invest considerable time to prepare a soaked rusk piece of a higher texture quality. Our present results suggest that the function of this behaviour is to soak the food. Because only some individuals dunked food and dunking has not been observed in the wild, we believe this to be a spontaneous foraging innovation either by one or multiple individuals.

Individual Goffin´s cockatoos (Cacatua goffiniana) show flexible targeted helping in a tool transfer task.

Flexible targeted helping is considered an advanced form of prosocial behavior in hominoids, as it requires the actor to assess different situations that a conspecific may be in, and to subsequently flexibly satisfy different needs of that partner depending on the nature of those situations. So far, apart from humans such behaviour has only been experimentally shown in chimpanzees and in Eurasian jays. Recent studies highlight the prosocial tendencies of several bird species, yet flexible targeted helping remained untested, largely due to methodological issues as such tasks are generally designed around tool-use, and very few bird species are capable of tool-use. Here, we tested Goffin's cockatoos, which proved to be skilled tool innovators in captivity, in a tool transfer task in which an actor had access to four different objects/tools and a partner to one of two different apparatuses that each required one of these tools to retrieve a reward. As expected from this species, we recorded playful object transfers across all conditions. Yet, importantly and similar to apes, three out of eight birds transferred the correct tool more often in the test condition than in a condition that also featured an apparatus but no partner. Furthermore, one of these birds transferred that correct tool first more often before transferring any other object in the test condition than in the no-partner condition, while the other two cockatoos were marginally non-significantly more likely to do so. Additionally, there was no difference in the likelihood of the correct tool being transferred first for either of the two apparatuses, suggesting that these birds flexibly adjusted what to transfer based on their partner´s need. Future studies should focus on explanations for the intra-specific variation of this behaviour, and should test other parrots and other large-brained birds to see how this can be generalized across the class and to investigate the evolutionary history of this trait.

Object manufacture based on a memorized template: Goffin's cockatoos attend to different model features.

Although several nonhuman animals have the ability to recognize and match templates in computerized tasks, we know little about their ability to recall and then physically manufacture specific features of mental templates. Across three experiments, Goffin cockatoos (Cacatua goffiniana), a species that can use tools in captivity, were exposed to two pre-made template objects, varying in either colour, size (long or short) or shape (I or L-shaped), where only one template was rewarded. Birds were then given the opportunity to manufacture versions of these objects themselves. We found that all birds carved paper strips from the same colour material as the rewarded template, and half were also able to match the size of a template (long and short). This occurred despite the template being absent at test and birds being rewarded at random. However, we found no evidence that cockatoos could carve L-shaped pieces after learning that L-shaped templates were rewarded, though their manufactured strips were wider than in previous tests. Overall, our results show that Goffin cockatoos possess the ability to physically adjust at least the size dimension of manufactured objects relative to a mental template. This ability has previously only been shown in New Caledonian crows, where template matching was suggested as a potential mechanism allowing for the cumulative cultural transmission of tool designs. Our results show that within avian tool users, the ability to recreate a physical template from memory does not seem to be restricted to species that have cumulative tool cultures.

Tool making cockatoos adjust the lengths but not the widths of their tools to function.

The ability to innovatively use or even manufacture different tools depending on a current situation can be silhouetted against examples of stereotyped, inborn tool use/manufacture and is thus often associated to advanced cognitive processing. In this study we confronted non-specialized, yet innovative tool making birds, Goffin's cockatoos (Cacatua goffiniana), with an apparatus featuring an out-of-reach food reward that could be placed at different distances from a tool opening. Alternatively, the food stayed at a constant distance but the tool opening in the front of the apparatus had different diameters. We used a novel material for tool manufacture (cardboard) that demanded an incrementally increased manufacturing effort from the actor, depending on the length of the tool required. We found that our subjects used two strategies to succeed in this tasks: either by making carboard-stripe tools using the full length of the material sheets originally offered or by adjusting the lengths of their tools to different goal distances. Subjects also discarded cardboard stripes that were too short to reach the goal prior to use and discarded longer pieces when the goal was further away than when it was close. Nevertheless, likely due to morphological constraints, the birds failed to adjust the widths of their tools depending on the diameter of the tool opening.

Spontaneous innovation of hook-bending and unbending in orangutans (Pongo abelii).

Betty the crow astonished the scientific world as she spontaneously crafted hook-tools from straight wire in order to lift a basket out of vertical tubes. Recently it was suggested that this species' solution was strongly influenced by predispositions from behavioural routines from habitual hook-tool manufacture. Nevertheless, the task became a paradigm to investigate tool innovation. Considering that young humans had surprising difficulties with the task, it was yet unclear whether the innovation of a hooked tool would be feasible to primates that lacked habitual hook making. We thus tested five captive orangutans in a hook bending and unbending task. Orangutans are habitually tool-using primates that have been reported to use but not craft hooked tools for locomotion in the wild. Two orangutans spontaneously innovated hook tools and four unbent the wire from their first trial on. Pre-experience with ready-made hooks had some effect but did not lead to continuous success. Further subjects improved the hook-design feature when the task required the subjects to bent the hook at a steeper angle. Our results indicate that the ability to represent and manufacture tools according to a current need does not require stereotyped behavioural routines, but can indeed arise innovatively. Furthermore, the present study shows that the capacity for hook tool innovation is not limited to large brained birds within non-human animals.

Compound tool construction by New Caledonian crows.

The construction of novel compound tools through assemblage of otherwise non-functional elements involves anticipation of the affordances of the tools to be built. Except for few observations in captive great apes, compound tool construction is unknown outside humans, and tool innovation appears late in human ontogeny. We report that habitually tool-using New Caledonian crows (Corvus moneduloides) can combine objects to construct novel compound tools. We presented 8 naïve crows with combinable elements too short to retrieve food targets. Four crows spontaneously combined elements to make functional tools, and did so conditionally on the position of food. One of them made 3- and 4-piece tools when required. In humans, individual innovation in compound tool construction is often claimed to be evolutionarily and mechanistically related to planning, complex task coordination, executive control, and even language. Our results are not accountable by direct reinforcement learning but corroborate that these crows possess highly flexible abilities that allow them to solve novel problems rapidly. The underlying cognitive processes however remain opaque for now. They probably include the species' typical propensity to use tools, their ability to judge affordances that make some objects usable as tools, and an ability to innovate perhaps through virtual, cognitive simulations.

Can hook-bending be let off the hook? Bending/unbending of pliant tools by cockatoos.

The spontaneous crafting of hook-tools from bendable material to lift a basket out of a vertical tube in corvids has widely been used as one of the prime examples of animal tool innovation. However, it was recently suggested that the animals' solution was hardly innovative but strongly influenced by predispositions from habitual tool use and nest building. We tested Goffin's cockatoo, which is neither a specialized tool user nor a nest builder, on a similar task set-up. Three birds individually learned to bend hook tools from straight wire to retrieve food from vertical tubes and four subjects unbent wire to retrieve food from horizontal tubes. Pre-experience with ready-made hooks had some effect but was not necessary for success. Our results indicate that the ability to represent and manufacture tools according to a current need does not require genetically hardwired behavioural routines, but can indeed arise innovatively from domain general cognitive processing.

Within-group relationships and lack of social enhancement during object manipulation in captive Goffin's cockatoos (Cacatua goffiniana).

Different types of social relationships can influence individual learning strategies in structured groups of animals. Studies on a number of avian species have suggested that local and/or stimulus enhancement are important ingredients of the respective species' exploration modes. Our aim was to identify the role of enhancement during object manipulation in different social contexts. We used focal observations to identify a linear dominance hierarchy as well as affiliative relationships between individuals in a group of 14 Goffin's cockatoos (Cacatua goffiniana, formerly goffini). Thereafter, in an unrewarded object choice task, several pairs of subjects were tested for a possible influence of social enhancement (local vs. stimulus) in three conditions: dominance, affiliation, and kinship. Our results suggest strong individual biases. Whereas previous studies on ravens and kea had indicated that enhancement in a non-food-related task was influenced by the social relationship between a demonstrator and an observer (affiliated - nonaffiliated), we found no such effects in our study group. In this context, Goffin's cockatoos' object learning seems to take place more on an individual level, despite their generally high motivation to manipulate nonfood items.

Flexible decision-making relative to reward quality and tool functionality in Goffin cockatoos (Cacatua goffiniana).

Decisions involving the use of tools may require an agent to consider more levels of relational complexity than merely deciding between an immediate and a delayed option. Using a new experimental approach featuring two different types of tools, two apparatuses as well as two different types of reward, we investigated the Goffin cockatoos' ability to make flexible and profitable decisions within five different setups. Paralleling previous results in primates, most birds overcame immediate drives in favor of future gains; some did so even if tool use involved additional work effort. Furthermore, at the group level subjects maximized their profit by simultaneously considering both the quality of an immediate versus a delayed food reward (accessible with a tool) and the functionality of the available tool. As their performance levels remained stable across trials in all testing setups, this was unlikely the result of a learning effect. The Goffin cockatoos' ability to focus on relevant information was constrained when all task components (both food qualities, both apparatuses and both tools) were presented at the same time.

Social transmission of tool use and tool manufacture in Goffin cockatoos (Cacatua goffini).

Tool use can be inherited, or acquired as an individual innovation or by social transmission. Having previously reported individual innovative tool use and manufacture by a Goffin cockatoo, we used the innovator (Figaro, a male) as a demonstrator to investigate social transmission. Twelve Goffins saw either demonstrations by Figaro, or 'ghost' controls where tools and/or food were manipulated using magnets. Subjects observing demonstrations showed greater tool-related performance than ghost controls, with all three males in this group (but not the three females) acquiring tool-using competence. Two of these three males further acquired tool-manufacturing competence. As the actions of successful observers differed from those of the demonstrator, result emulation rather than high-fidelity imitation is the most plausible transmission mechanism.

Goffin cockatoos wait for qualitative and quantitative gains but prefer 'better' to 'more'.

Evidence for flexible impulse control over food consumption is rare in non-human animals. So far, only primates and corvids have been shown to be able to fully inhibit the consumption of a desirable food item in anticipation for a gain in quality or quantity longer than a minute. We tested Goffin cockatoos (Cacatua goffini) in an exchange task. Subjects were able to bridge delays of up to 80 s for a preferred food quality and up to 20 s for a higher quantity, providing the first evidence for temporal discounting in birds that do not cache food.