Object play and problem solving in infancy: Insights into tool use.

Tool use is primarily, but not exclusively, present in species with otherwise advanced cognitive traits. However, the interaction between such traits and conspecific inter-individual variation in the presence, complexity, or intensity of tool use is far from being established. We addressed this matter among human infants, seeking factors that relate to differences in tool use. We examined, both correlationally and experimentally, whether the propensity to engage in object combinations predicts performance in means-end problem-solving tasks involving or not involving the use of a tool. We tested 71 infants aged 15, 18, 21, and 24 months, dividing them into two subgroups: one exposed to an adult demonstrating object-object combinations (i.e., "prompting" infants to combine objects together) and another with comparable social exposure but where the adult demonstrated single-object manipulations. We found a correlation between the combined level of spontaneous and prompted object combinations and problem-solving performance regardless of the involvement of tools in the problem. However, we did not find differences in tool-use performance between the two demonstration subgroups. The correlational analysis suggests that complexity of play, as measured by the frequency of combining objects, is linked to infants' problem-solving skills rather than being specifically associated with tool use, as previously suggested in the literature.

Memory for own actions in parrots.

The ability to recall one's past actions is a crucial prerequisite for mental self-representation and episodic memory. We studied whether blue-throated macaws, a social macaw species, can remember their previous actions. The parrots were trained to repeat four previously learned actions upon command. Test sessions included repeat trials, double repeat trials and trials without repeat intermixed to test if the parrots repeated correctly, only when requested and not relying on a representation of the last behavioral command. Following their success, the parrots also received sessions with increasing time delays preceding the repeat command and successfully mastered 12-15 s delays. The parrots successfully transferred the repeat command spontaneously at first trial to three newly trained behaviors they had never repeated before, and also succeeded in a second trial intermixed with already trained actions (untrained repeat tests). This corroborates that successful repeating is not just an artifact of intense training but that blue-throated macaws can transfer the abstract "repeat rule" to untrained action. It also implies that an important aspect of self-representation has evolved in this avian group and might be adaptive, which is consistent with the complex socio-ecological environment of parrots and previous demonstrations of their complex cognition.

Intra- and interspecific variation in self-control capacities of parrots in a delay of gratification task.

Forgoing immediate satisfaction for higher pay-offs in the future (delayed gratification) could be adaptive in situations that wild animals may encounter. To explain species-differences in self-control, hypotheses based on social complexity, feeding ecology, brain size and metabolic rate have been proposed. To explore these hypotheses in a comparative setting, we tested three macaw species (neotropical parrots)-great green macaws (N = 8), blue-throated macaws (N = 6), blue-headed macaws (N = 6)-and the distantly related African grey parrots (afrotropical parrots; N = 8) in a modified rotating tray task, in which subjects are required to inhibit consuming a constantly available low-quality reward in favour of a high-quality reward that becomes available only after an increasing delay (min. 5 s, max. 60 s). All four species successfully waited for a minimum of 8.3 s ± 11.7 s (group level mean ± SD) with African greys reaching a delay of 29.4 ± 15.2 s, and great green macaws-as best performing macaw species-tolerating delays of 20 s ± 8 s. The best performing African grey individual reached a maximum delay of 50 s, whereas, a great green and a blue-throated macaw tolerated a delay of 30 s max. Females tolerated higher maximum delays than males. Engaging in distraction behaviours enhanced waiting performance across species and all birds were able to anticipate the waiting duration. Our results suggest that both feeding and socio-ecological complexity may be a factor in self-control, but further systematically collected comparative data on self-control of different (parrot) species are required to test the evolutionary hypotheses rigorously.

Complex nests but no use of tools: An investigation of problem solving in weaverbirds (Ploceidae).

Few avian species use tools in the wild. Yet, several birds build nests of great complexity and many aspects of tool use may also apply to nest building. It has long been hypothesised that tool use may select for specialised cognitive adaptations or even general cognitive skills. This might similarly pertain to species that build complex nests. In this study, we investigated the problem-solving capacity of a complex nest builder, a weaverbird species, in a foraging context that either required or did not require the use of tools. First, we tested the capacity of yellow-crowned bishops (Euplectes afer ssp. afer) to use a tool for retrieving an out-of-reach reward during three problem-solving tasks offering different nest/non-nest materials (Experiment 1). Next, subjects were confronted with two problem-solving tasks that required no tools (Experiment 2). No subject was able to use a tool in Experiment 1. However, 11 out of 12 subjects succeeded in using their beak in the first problem-solving task, and 9 in the second problem-solving task of Experiment 2. These results suggest that weaverbirds showed flexible problem-solving if the use of tools was not required.

Characteristics of urban environments and novel problem-solving performance in Eurasian red squirrels.

Urban environments can be deemed 'harsh' for some wildlife species, but individuals frequently show behavioural flexibility to cope with challenges and demands posed by life in the city. For example, urban animals often show better performance in solving novel problems than rural conspecifics, which helps when using novel resources under human-modified environments. However, which characteristics of urban environments fine-tune novel problem-solving performance, and their relative importance, remain unclear. Here, we examined how four urban environmental characteristics (direct human disturbance, indirect human disturbance, size of green coverage and squirrel population size) may potentially influence novel problem-solving performance of a successful 'urban dweller', the Eurasian red squirrel, by presenting them with a novel food-extraction problem. We found that increased direct human disturbance, indirect human disturbance and a higher squirrel population size decreased the proportion of solving success at the population level. At the individual level, an increase in squirrel population size decreased the latency to successfully solve the novel problem the first time. More importantly, increased direct human disturbance, squirrel population size and experience with the novel problem decreased problem-solving time over time. These findings highlight that some urban environmental characteristics shape two phenotypic extremes in the behaviour-flexibility spectrum: individuals either demonstrated enhanced learning or they failed to solve the novel problem.

Innovative problem solving in macaws.

Behavioural innovations with tool-like objects in non-habitually tool-using species are thought to require complex physical understanding, but the underlying cognitive processes remain poorly understood. A few parrot species are capable of innovating tool-use and borderline tool-use behaviours. We tested this capacity in two species of macaw (Ara ambiguus, n = 9; Ara glaucogularis, n = 8) to investigate if they could solve a problem-solving task through manufacture of a multi-stone construction. Specifically, after having functional experience with a pre-inserted stick tool to push a reward out of a horizontal tube, the subjects were required to insert five stones consecutively from one side to perform the same function as the stick tool with the resulting multi-component construction. One Ara glaucogularis solved the task and innovated the stone construction after the experience with the stick tool. Two more subjects (one of each species) did so after having further functional experience of a single stone pushing a reward out of a shortened tube. These subjects were able to consistently solve the task, but often made errors, for example counter-productive stone insertions from the opposing end, even in some of the successful trials. Conversely, multiple trials without errors also suggested a strong goal direction. Their performance in the follow-up tasks was inconclusive since they sometimes inserted stones into un-baited or blocked 'dummy tubes', but this could have been an attention-deficit behaviour as subjects had not encountered these 'dummy tubes' before. Overall, the successful subjects' performance was so erratic that it proved difficult to conclude whether they had functional understanding of their multi-stone constructions.

Comparative cognition: Practical shortcomings and some potential ways forward.

The objectives in the field of comparative cognition are clear; efforts are devoted to revealing the selection pressures that shape the brains and cognitive abilities of different species and understanding cognitive processes in differently structured brains. However, our progress on reaching these objectives is slow, mostly because of several major practical challenges. In this review, we discuss 2 major shortcomings: (a) the poor systematics and low magnitude of the phylogenetic comparisons made, and (b) the weak comparability of the results caused by interfering species-specific confounding factors (perceptual, motivational, and morphological) alongside an insufficient level of standardisation of the methodologies. We propose a multiple-level comparative approach that emphasises the importance of achieving more direct comparisons within taxonomic groups at genus or family level as the first step before comparing between distantly related groups. We also encourage increasing interdisciplinary efforts to execute "team-science" approach in building a systematic and direct large-scale phylogenetic comparisons of bigger cognitive test batteries that produce reliable species-representative data. We finally revisit some existing suggestions that allow us to maximise standardisation while minimising species-specific confounding factors. (PsycInfo Database Record (c) 2020 APA, all rights reserved).

Genetic monogamy despite frequent extrapair copulations in "strictly monogamous" wild jackdaws.

"Monogamy" refers to different components of pair exclusiveness: the social pair, sexual partners, and the genetic outcome of sexual encounters. Avian monogamy is usually defined socially or genetically, whereas quantifications of sexual behavior remain scarce. Jackdaws (Corvus monedula) are considered a rare example of strict monogamy in songbirds, with lifelong pair bonds and little genetic evidence for extrapair (EP) offspring. Yet jackdaw copulations, although accompanied by loud copulation calls, are rarely observed because they occur visually concealed inside nest cavities. Using full-day nest-box video surveillance and on-bird acoustic bio-logging, we directly observed jackdaw sexual behavior and compared it to the corresponding genetic outcome obtained via molecular parentage analysis. In the video-observed nests, we found genetic monogamy but frequently detected forced EP sexual behavior, accompanied by characteristic male copulation calls. We, thus, challenge the long-held notion of strict jackdaw monogamy at the sexual level. Our data suggest that male mate guarding and frequent intrapair copulations during the female fertile phase, as well as the forced nature of the copulations, could explain the absence of EP offspring. Because EP copulation behavior appeared to be costly for both sexes, we suggest that immediate fitness benefits are an unlikely explanation for its prevalence. Instead, sexual conflict and dominance effects could interact to shape the spatiotemporal pattern of EP sexual behavior in this species. Our results call for larger-scale investigations of jackdaw sexual behavior and parentage and highlight the importance of combining social, sexual, and genetic data sets for a more complete understanding of mating systems.

Social learning in great white pelicans (Pelecanus onocrotalus): A preliminary study.

Great white pelicans (Pelecanus onocrotalus) exhibit life-history parameters and ecological traits thought to be associated with social learning, and advanced cognitive processing more generally. In this study we investigated whether this species can acquire novel behavior socially in a foraging context. Birds from the test group watched a trained conspecific opening an opaque box containing a food reward by using its beak, whereas the control group had no demonstrator but saw the box for an equivalent time span. Individuals from both groups were subsequently allowed access to the box. Subjects of the test group performed significantly better than the control group. This is the first experimental evidence of social learning in a cooperatively hunting bird. Further studies are needed in order to shed light on the factors favoring the evolution of this capacity, by testing different pelican species that vary in their ecology.

Parrots Voluntarily Help Each Other to Obtain Food Rewards.

Helping others to obtain benefits, even at a cost to oneself, poses an evolutionary puzzle [1]. While kin selection explains such "selfless" acts among relatives, only reciprocity (paying back received favors) entails fitness benefits for unrelated individuals [2]. So far, experimental evidence for both prosocial helping (providing voluntary assistance for achieving an action-based goal) and reciprocity has been reported in a few mammals but no avian species [3]. In order to gain insights into the evolutionary origins of these behaviors, the capacity of non-mammalian species for prosociality and for reciprocity needs to be investigated. We tested two parrot species in an instrumental-helping paradigm involving "token transfer." Here, actors could provide tokens to their neighbor, who could exchange them with an experimenter for food. To verify whether the parrots understood the task's contingencies, we systematically varied the presence of a partner and the possibility for exchange. We found that African grey parrots voluntarily and spontaneously transferred tokens to conspecific partners, whereas significantly fewer transfers occurred in the control conditions. Transfers were affected by the strength of the dyads' affiliation and partially by the receivers' attention-getting behaviors. Furthermore, the birds reciprocated the help once the roles were reversed. Blue-headed macaws, in contrast, transferred hardly any tokens. Species differences in social tolerance might explain this discrepancy. These findings show that instrumental helping based on a prosocial attitude, accompanied but potentially not sustained by reciprocity, is present in parrots, suggesting that this capacity evolved convergently in this avian group and mammals.

Assessing African grey parrots' prosocial tendencies in a token choice paradigm.

Prosociality is defined as a voluntary, typically low-cost behaviour that benefits another individual. Social tolerance has been proposed as a potential driver for its evolution, both on the proximate and on the ultimate level. Parrots are an interesting species to study such other-regarding behaviours, given that they are highly social and stand out in terms of relative brain size and cognitive capacity. We tested eight African grey parrots in a dyadic prosocial choice test. They faced a choice between two different tokens, a prosocial (actor and partner rewarded) and a selfish (only actor rewarded) one. We found that the birds did not behave prosocially when one subject remained in the actor role; however, when roles were alternated, the birds' prosocial choices increased. The birds also seemed to reciprocate their partner's choices, given that a contingency between choices was observed. If the food provisioned to the partner was of higher quality than that the actor obtained, actors increased their willingness to provide food to their partner. Nonetheless, the control conditions suggest that the parrots did not fully understand the task's contingencies. In sum, African grey parrots show the potential for prosociality and reciprocity; however, considering their lack of understanding of the contingencies of the particular tasks used in this study, the underlying motivation for the observed behaviour remains to be addressed by future studies, in order to elucidate the phylogenetic distribution of prosociality further.

Emotional responses to conspecific distress calls are modulated by affiliation in cockatiels (Nymphicus hollandicus).

Vocal communication is used across the animal kingdom to transfer information from emitters to receivers, such as size, sex, age, dominance status or even emotional states. The transmission of an emotional state from one individual to another is called "emotional contagion" and is classified as the first level of empathy. Emotional contagion is thought to be stronger between familiar individuals. While affiliation represents a stronger relation between individuals than mere familiarity, it remains understudied whether affiliation modulates emotional reactions as well. Using cockatiels (Nymphicus hollandicus), we played back three types of audio stimuli to individual birds: a partner's distress call (emitted when birds are caught or forcibly restrained), a non-partner's distress call, and a control sound (white noise). The calls were recorded from familiar birds with either low (non-partners) or high levels of affiliation (partners). The subjects' response was scored using four behavioural parameters: the time spent near the loudspeaker, the amount of movements, the number of calls emitted, and the position of the crest. Across all variables, birds were more attentive and active when confronted to distress calls compared to control sounds, particularly when the distress call was emitted from a partner rather than a non-partner. These results raise the possibility that distress calls do not only function as a stimulus-triggering automatic reaction in cockatiels but also transmit emotions. Moreover, affiliation enhanced emotional reactions to conspecific distress calls. Our data provides first insights into the mechanisms of emotional contagion in parrots.

Economic Decision-Making in Parrots.

Economic decision-making involves weighing up differently beneficial alternatives to maximise payoff. This sometimes requires the ability to forego one's desire for immediate satisfaction. This ability is considered cognitively challenging because it not only requires inhibiting impulses, but also evaluating expected outcomes in order to decide whether waiting is worthwhile. We tested four parrot species in a token exchange task. The subjects were first trained to exchange three types of tokens for a food item of low, medium, and high value and successfully learned to exchange these in an order according to their value. Subsequently, they were confronted with a choice between a food item and a token that could be exchanged for higher-quality food. In additional control conditions however, choosing a token led to an equal or lower payoff. Individuals of all species were capable of deciding economically, yet only large macaws outperformed the other species in one of the crucial controls. For some individuals, particularly African grey parrots, the token apparently had an intrinsic value, which prevented them from choosing economically in some control conditions and which should be considered as potentially confounding by future token exchange studies.

Are parrots poor at motor self-regulation or is the cylinder task poor at measuring it?

The ability to inhibit unproductive motor responses triggered by salient stimuli is a fundamental inhibitory skill. Such motor self-regulation is thought to underlie more complex cognitive mechanisms, like self-control. Recently, a large-scale study, comparing 36 species, found that absolute brain size best predicted competence in motor inhibition, with great apes as the best performers. This was challenged when three Corvus species (corvids) were found to parallel great apes despite having much smaller absolute brain sizes. However, new analyses suggest that it is the number of pallial neurons, and not absolute brain size per se, that correlates with levels of motor inhibition. Both studies used the cylinder task, a detour-reaching test where food is presented behind a transparent barrier. We tested four species from the order Psittaciformes (parrots) on this task. Like corvids, many parrots have relatively large brains, high numbers of pallial neurons, and solve challenging cognitive tasks. Nonetheless, parrots performed markedly worse than the Corvus species in the cylinder task and exhibited strong learning effects in performance and response times. Our results suggest either that parrots are poor at controlling their motor impulses, and hence that pallial neuronal numbers do not always correlate with such skills, or that the widely used cylinder task may not be a good measure of motor inhibition.

Ravens, New Caledonian crows and jackdaws parallel great apes in motor self-regulation despite smaller brains.

Overriding motor impulses instigated by salient perceptual stimuli represent a fundamental inhibitory skill. Such motor self-regulation facilitates more rational behaviour, as it brings economy into the bodily interaction with the physical and social world. It also underlies certain complex cognitive processes including decision making. Recently, MacLean et al. (MacLean et al. 2014 Proc. Natl Acad. Sci. USA 111, 2140-2148. (doi:10.1073/pnas.1323533111)) conducted a large-scale study involving 36 species, comparing motor self-regulation across taxa. They concluded that absolute brain size predicts level of performance. The great apes were most successful. Only a few of the species tested were birds. Given birds' small brain size-in absolute terms-yet flexible behaviour, their motor self-regulation calls for closer study. Corvids exhibit some of the largest relative avian brain sizes-although small in absolute measure-as well as the most flexible cognition in the animal kingdom. We therefore tested ravens, New Caledonian crows and jackdaws in the so-called cylinder task. We found performance indistinguishable from that of great apes despite the much smaller brains. We found both absolute and relative brain volume to be a reliable predictor of performance within Aves. The complex cognition of corvids is often likened to that of great apes; our results show further that they share similar fundamental cognitive mechanisms.

Combinatory actions during object play in psittaciformes (Diopsittaca nobilis, Pionites melanocephala, Cacatua goffini) and corvids (Corvus corax, C. monedula, C. moneduloides).

The playful (i.e., not overtly functional) combination of objects is considered a potential ontogenetic and phylogenetic precursor of technical problem solving abilities, as it may lead to affordance learning and honing of mechanical skills. We compared such activities in 6 avian species: 3 psittaciforms (black-headed caiques, red-shouldered macaws, and Goffin cockatoos) and 3 corvids (New Caledonian crows, ravens, and jackdaws). Differences in the type and frequency of object combinations were consistent with species' ecology. Object caching was found predominately in common ravens, which frequently cache food. The most intrinsically structured object combinations were found in New Caledonian crows and Goffin cockatoos, which both stand out for their problem solving abilities in physical tasks. Object insertions prevailed in New Caledonian crows that naturally extract food using tools. Our results support the idea that playful manipulations of inedible objects are linked to physical cognition and problem-solving abilities.

Monocular tool control, eye dominance, and laterality in New Caledonian crows.

Tool use, though rare, is taxonomically widespread, but morphological adaptations for tool use are virtually unknown. We focus on the New Caledonian crow (NCC, Corvus moneduloides), which displays some of the most innovative tool-related behavior among nonhumans. One of their major food sources is larvae extracted from burrows with sticks held diagonally in the bill, oriented with individual, but not species-wide, laterality. Among possible behavioral and anatomical adaptations for tool use, NCCs possess unusually wide binocular visual fields (up to 60°), suggesting that extreme binocular vision may facilitate tool use. Here, we establish that during natural extractions, tool tips can only be viewed by the contralateral eye. Thus, maintaining binocular view of tool tips is unlikely to have selected for wide binocular fields; the selective factor is more likely to have been to allow each eye to see far enough across the midsagittal line to view the tool's tip monocularly. Consequently, we tested the hypothesis that tool side preference follows eye preference and found that eye dominance does predict tool laterality across individuals. This contrasts with humans' species-wide motor laterality and uncorrelated motor-visual laterality, possibly because bill-held tools are viewed monocularly and move in concert with eyes, whereas hand-held tools are visible to both eyes and allow independent combinations of eye preference and handedness. This difference may affect other models of coordination between vision and mechanical control, not necessarily involving tools.

Object caching in corvids: incidence and significance.

Food caching is a paramount model for studying relations between cognition, brain organisation and ecology in corvids. In contrast, behaviour towards inedible objects is poorly examined and understood. We review the literature on object caching in corvids and other birds, and describe an exploratory study on object caching in ravens, New Caledonian crows and jackdaws. The captive adult birds were presented with an identical set of novel objects adjacent to food. All three species cached objects, which shows the behaviour not to be restricted to juveniles, food cachers, tool-users or individuals deprived of cacheable food. The pattern of object interaction and caching did not mirror the incidence of food caching: the intensely food caching ravens indeed showed highest object caching incidence, but the rarely food caching jackdaws cached objects to similar extent as the moderate food caching New Caledonian crows. Ravens and jackdaws preferred objects with greater sphericity, but New Caledonian crows preferred stick-like objects (similar to tools). We suggest that the observed object caching might have been expressions of exploration or play, and deserves being studied in its own right because of its potential significance for tool-related behaviour and learning, rather than as an over-spill from food-caching research. This article is part of a Special Issue entitled: CO3 2013.

Object permanence in the Goffin cockatoo (Cacatua goffini).

The ability to represent hidden objects plays an important role in the survival of many species. In order to provide an inclusive synopsis of the current benchmark tasks used to test object permanence in animals for a psittacine representative, we tested eight Goffin cockatoos (Cacatua goffini) on Stages 3-6 of Piagetian object permanence as well as derivations of spatial transposition, rotation, and translocation tasks. Subjects instantly solved visible displacement 3b and 4a but showed an extended plateau for solving Stage 5a at a very late age (10 months). Subjects readily solved most invisible displacement tasks including double hidings and four angles (90°, 180°, 270°, and 360°) of rotation and translocations at high performance levels, although Piagetian Stage 6 invisible displacement tasks caused more difficulties for the animals than transposition, rotations, and translocation tasks.