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Social learning describes the acquisition of knowledge through observation of other individuals, and it is fundamental for the development of culture and traditions within human groups. Although previous studies suggest that Kea (Nestor notabilis) benefit from social learning, experimental evidence has been inconclusive, as in a recent two-action task, all perceptual factors were ignored. The present study attempts to address this by investigating social learning in Kea with a focus on social enhancement processes. In an experiment with a captive group of Kea, we investigated whether individuals that had the opportunity to observe a conspecific performing a simple task subsequently show better performance in that task than a control group without prior demonstration. This study provides a strong tendency of greater success in skill acquisition in Kea as a result of social learning. Kea that observed a conspecific solving a task showed clear evidence of perceptual factors drawing attention to the relevant parts of the experimental apparatus and manipulated these significantly more (100% of trials) than control birds (77.8% of trials). Combined with a strong trend (p = 0.056) of the test subjects solving the task more than the control subjects, this shows conclusively that Kea, at least when required to solve a task, do attend to perceptual factors of a demonstrated action.

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Social play has been described in many animals. However, much of this social behaviour among birds, particularly in adults, is still relatively unexplored in terms of the environmental, psychological, and social dynamics of play. This paper provides an overview of what we know about adult social play in birds and addresses areas in which subtleties and distinctions, such as in play initiation and social organisation and its relationship to expressions of play, are considered in detail. The paper considers emotional, social, innovative, and cognitive aspects of play, then the environmental conditions and affiliative bonds, suggesting a surprisingly complex framework of criteria awaiting further research. Adult social play has so far been studied in only a small number of avian species, exclusively in those with a particularly large brain relative to body size without necessarily addressing brain functions and lateralization. When lateralization of brain function is considered, it can further illuminate a possibly significant relevance of play behaviour to the evolution of cognition, to management of emotions, and the development of sociality.

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Category learning is often tested with similar images that have no significance outside of the experiment for the subjects. By contrast, in nature animals often need to generalize a behavioral response like "eat" across visually distinct stimuli, such as spiders and seeds. Forming functional categories like "food" and "predator" may require conceptual rather than purely perceptual generalization. We trained free-range chickens to classify images assigned to one of four categories based on putative functional significance: inanimate objects, predators, food, and non-competing vertebrates. Images were visually diverse within each category, discouraging classification by perceptual similarity alone. In Experiment 1, chickens classified 80 images into four categories. Chickens then generalized to 80 new exemplars in each of three successive generalization tests. In Experiment 2, chickens saw new types of images to test whether their generalization was perceptual or functional. For example, chickens saw images of skunks for the predator category after training with images of hawks and snakes. Chickens used the "predator" response with these new images for both predators and non-threatening vertebrates, but not for objects or food, and did not successfully generalize any category other than predator. In Experiment 3, chickens categorized fractals as "food," and three of four chickens categorized a range of vertebrates they had not previously encountered as "predators," suggesting that chickens did not see the images as representing real world objects and animals. These results highlight constraints on the use of computer-generated images to assess categorization of natural stimuli in chickens.

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Object permanence, the ability to mentally represent objects even when they are not directly accessible to the senses, is of vital importance for the survival of both human and non-human animals. The Oriental pied hornbill (Anthracoceros albirostris) is an Asian species of hornbill displaying remarkable adaptability in various environments, yet little is known about their cognitive abilities. Their breeding behaviour is unique, as the female hornbill seals herself inside a cavity before laying eggs and the male feeds her and their offspring without visual contact, strongly suggesting the presence of object permanence to some degree. In this study, six Oriental pied hornbills underwent testing for object permanence, including a series of seven standard Piagetian tasks involving visible and invisible displacements. The subjects consistently demonstrated spontaneous object permanence in all stages leading up to the invisible displacement stage. Half of the subjects achieved full stage 6 double invisible displacement Piagetian object permanence, while the other half reached stage 5 double visible displacement. Breeding behaviour and the duration of developmental stages are proposed as potential factors influencing object permanence ability in this species of hornbill.

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The capacity to create new behaviors is influenced by environmental factors such as foraging ecology, which can lead to phylogenetic variation in innovativeness. Alternatively, these differences may arise due to the selection of the underlying mechanisms, collaterally affecting innovativeness. To understand the evolutionary pathways that might enhance innovativeness, we examined the role of diet breadth and degree of extractive foraging, as well as a range of intervening cognitive and behavioral mechanisms (neophilia, neophobia, flexibility, motivation, and persistence). Darwin's finches are very suitable to this purpose: the clade is composed of closely related species that vary in their feeding habits and capacity to develop food innovations. Using a multi-access box, we conducted an interspecies comparison on innovative problem-solving between two diet specialists, extractive foragers (woodpecker and cactus finch), and two diet generalists, non-extractive foragers (small and medium ground finch). We predicted that if extractive foraging was associated with high innovativeness, variation would be best explained by species differences in persistence and motivation, whereas if diet generalism was the main driver, then variation would be due to differences in flexibility and responses to novelty. We found a faster capacity to innovate and a higher persistence for extractive foragers, suggesting that persistence might be adaptive to extractive foraging and only secondarily to innovation. Our findings also show that diet generalism and some variables linking it to innovation were unrelated to innovativeness and call for the development of joint experimental approaches that capture the diversity of factors giving rise to novel behaviors.

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Categorization represents one cognitive ability fundamental to animal behavior. Grouping of elements based on perceptual or semantic features helps to reduce processing resources and facilitates appropriate behavior. Corvids master complex categorization, yet the detailed categorization learning strategies are less well understood. We trained two jackdaws on a delayed match to category paradigm using a novel, artificial stimulus type, RUBubbles. Both birds learned to differentiate between two session-unique categories following two distinct learning protocols. Categories were either introduced via central category prototypes (low variability approach) or using a subset of diverse category exemplars from which diagnostic features had to be identified (high variability approach). In both versions, the stimulus similarity relative to a central category prototype explained categorization performance best. Jackdaws consistently used a central prototype to judge category membership, regardless of whether this prototype was used to introduce distinct categories or had to be inferred from multiple exemplars. Reliance on a category prototype occurred already after experiencing only a few trials with different category exemplars. High stimulus set variability prolonged initial learning but showed no consistent beneficial effect on later generalization performance. High numbers of stimuli, their perceptual similarity, and coherent category structure resulted in a prototype-based strategy, reflecting the most adaptive, efficient, and parsimonious way to represent RUBubble categories. Thus, our birds represent a valuable comparative animal model that permits further study of category representations throughout learning in different regions of a brain producing highly cognitive behavior.

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A recent study by Johnston, Brecht, and Nieder (2023, Current Biology, 33, 3238-3243) finds that carrion crows associate varying rates of reinforcement with novel arbitrary stimuli and make optimal decisions when they must later choose between stimulus pairs. These results demonstrate that crows are capable of not only storing information about reward probabilities in their memory but also making optimal choices based on this information even a month later.

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Few month old human infants are able to detect the social roles of artificial agents and consistently choose the object behaving as 'approacher' rather than 'repulser'. This preference has been considered evidence of a pre-linguistic and pre-cultural origin of the social mind. Similar preferences have not been described in other species, though comparative data could help clarify the nature of this phenomenon and its evolutionary origin. In this study, we investigated sensitivity to the social role of an artificial agent in domestic chicks. Birds offer an excellent model to study the evolutionary roots of cognitive abilities, since they separated from mammals over 300 Ma. Moreover, the investigation of newly hatched chicks allows control for previous experience. After being exposed to computer-presented animations depicting an interaction among two agents, chicks underwent a free choice test among those same objects. While no initial evidence of a clear preference emerged from the planned analysis, chicks in the experimental condition showed a preference for the 'approacher' when controlling for side bias, mirroring human infants behaviour. This suggests the existence of an early ability to discriminate agents from their interactions, independent from any social experience.

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The use of abstract rules in behavioral decisions is considered evidence of executive functions associated with higher-level cognition. Laboratory studies across taxa have shown that animals may be capable of learning abstract concepts, such as the relationships between items, but often use simpler cognitive abilities to solve tasks. Little is known about whether or how animals learn and use abstract rules in natural environments. Here, we tested whether wild, food-caching mountain chickadees (Poecile gambeli) could learn an abstract rule in a spatial-temporal task in which the location of a food reward rotated daily around an 8-feeder square spatial array for up to 34 days. Chickadees initially searched for the daily food reward by visiting the most recently rewarding locations and then moving backward to visit previously rewarding feeders, using memory of previous locations. But by the end of the task, chickadees were more likely to search forward in the correct direction of rotation, moving away from the previously rewarding feeders. These results suggest that chickadees learned the direction rule for daily feeder rotation and used this to guide their decisions while searching for a food reward. Thus, chickadees appear to use an executive function to make decisions on a foraging-based task in the wild. VIDEO ABSTRACT.

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Categorizing individuals on the basis of familiarity is an adaptive way of dealing with the complexity of the social environment. It requires the use of conceptual familiarity and is considered higher order learning. Although, it is common among many species, ecological need might require and facilitate individual differentiation among heterospecifics. This may be true for laboratory populations just as much as for domesticated species and those that live in urban contexts. However, with the exception of a few studies, populations of laboratory animals have generally been given less attention. The study at hand, therefore, addressed the question whether a laboratory population of kea parrots (Nestor notabilis) were able to apply the concept of familiarity to differentiate between human faces in a two-choice discrimination task on the touchscreen. The results illustrated that the laboratory population of kea were indeed able to differentiate between familiar and unfamiliar human faces in a two-choice discrimination task. The results provide novel empirical evidence on abstract categorization capacities in parrots while at the same time providing further evidence of representational insight in kea.

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Mirror exposure elicits a wide range of behavioral responses, some of which have been considered as part of possible evidence of mirror self-recognition (MSR). These responses can range from social behaviors, indicating that an animal considers its own reflection as a conspecific, to mirror-guided and self-directed actions. Evidence of MSR has been found categorically in only a few species, such as in magpies, chimpanzees, horses, and elephants. Evidence in corvids is currently debated due to inconsistent findings. In this study, we investigated the reaction of Eurasian jays when presenting them with three mirror-stimulation tasks. Based on the overall behavioral patterns across these three tasks, conclusions about birds' understanding of a reflective surface, and their perception of the reflection as either themselves or as a conspecific, appear premature. We highlight how the high neophobia of corvids and other methodological constraints might have hindered the likelihood to approach and explore a mirror, preventing the emergence of behaviors typically associated with MSR. Furthermore, we discuss how motivational factors, methodological constraints and species differences should be considered when interpreting behavioral responses to mirrors.

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Logical inference, once strictly associated with spoken language, is now reported in some non-human animals. One aspect of logical inference, reasoning by exclusion, has been traditionally explored through the use of the cups task (cup A and cup B, if not cup A, then exclude cup A and select cup B). However, to fully understand the factors that drove the evolution of logical processes in animals, this latter paradigm needs to cover a taxonomically broader spectrum of species. In this study, we aimed to test the capacity of Southern ground-hornbills (Bucorvus leadbeateri) to show exclusion performance in a two-way object-choice task. First, we determined whether subjects could perceive and choose correctly between two containers (one rewarded, one unrewarded) using visual or acoustic cues (sensory phase). If successful, individuals were then presented with three experimental conditions (test phase): Full information (content of both cups revealed), Exclusion (content of the empty cup revealed), and Control (no content revealed). During the sensory phase, ground-hornbills succeeded in choosing the rewarded container only in the visual modality. Birds were able to select the rewarded container more than would be expected by chance in the Full information condition, but their performances were equal to chance in the Control condition. The without-learning performance of two individuals within the Exclusion condition indicates that this task is not trivial, which invites further investigation on this species' capacity to represent the dependent relationship between the cups (true logical inference).

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Mirror tasks can be used to investigate whether animals can instrumentally use a mirror to solve problems and can understand the correspondence between reflections and the real objects they represent. Two bird species, a corvid (New Caledonian crow) and a parrot (African grey parrot), have demonstrated the ability to use mirrors instrumentally in mirror-mediated spatial locating tasks. However, they have not been challenged with a mirror-guided reaching task, which involves a more complex understanding of the mirror's properties. In the present study, a task approximating the mirror-guided reaching task used in primate studies was adapted for, and given to, a corvid species (Eurasian jay) using a horizontal string-pulling paradigm. Four birds learned to pull the correct string to retrieve a food reward when they could see the food directly, whereas none used the reflected information to accomplish the same objective. Based on these results, it cannot be concluded whether these birds understand the correspondence between the location of the reward and its reflected information, or if the relative lack of visual-perceptual motor feedback given by the setup interfered with their performance. This novel task is posited to be conceptually more difficult compared to mirror-mediated spatial locating tasks, and should be used in avian species that have previously been successful at using the mirror instrumentally. This would establish whether these species can still succeed at it, and thus whether the task does indeed pose additional cognitive demands.

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Inferential reasoning by exclusion allows responding adaptively to various environmental stimuli when confronted with inconsistent or partial information. In the experimental context, this mechanism involves selecting correctly between an empty option and a potentially rewarded one. Recently, the increasing reports of this capacity in phylogenetically distant species have led to the assumption that reasoning by exclusion is the result of convergent evolution. Within one largely unstudied avian order, i.e. the Charadriiformes, brown skuas (Catharacta antarctica ssp lonnbergi) are highly flexible and opportunistic predators. Behavioural flexibility, along with specific aspects of skuas' feeding ecology, may act as influencing factors in their ability to show exclusion performance. Our study aims to test whether skuas are able to choose by exclusion in a visual two-way object-choice task. Twenty-six wild birds were presented with two opaque cups, one covering a food reward. Three conditions were used: 'full information' (showing the content of both cups), 'exclusion' (showing the content of the empty cup), and 'control' (not showing any content). Skuas preferentially selected the rewarded cup in the full information and exclusion condition. The use of olfactory cues was excluded by results in the control condition. Our study opens new field investigations for testing further the cognition of this predatory seabird.

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Cognition influences how individuals interact with the environment, affecting the ecology of species. Gaining insight into the proficiency of relevant cognitive abilities provides an indication of the processes necessary for a species' survival and reproduction. Many birds have "slow" life-histories and complex social environments suggestive of high cognitive ability. Little, however, is known about the cognition of most birds with these traits, thus studying cognition in seabirds with these traits provides insight into how slow life-histories and complex social environments relate more generally to predicting cognitive ability. Object permanence is a cognitive ability shared by highly intelligent animals and could be an ecologically relevant ability for many seabirds. I used a simple experimental setup in a semi-controlled environment to test object permanence in captive horned puffins (Fratercula corniculata) by hiding a reward to be retrieved, first partially and then completely. I discovered that the horned puffins performed poorly on the object permanence task when the reward was hidden completely. I discuss briefly how the slow life-histories of many seabirds probably evolved due to the stochastic conditions associated with their marine environment, which in turn may cause an energetic bottleneck that limits the allocation of resources to certain cognitive abilities.

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Macphail famously criticized two foundational assumptions that underlie the evolutionary approach to comparative psychology: that there are differences in intelligence across species, and that intelligent behavior in animals is based on more than associative learning. Here, we provide evidence from recent work in avian cognition that supports both these assumptions: intelligence across species varies, and animals can perform intelligent behaviors that are not guided solely by associative learning mechanisms. Finally, we reflect on the limitations of comparative psychology that led to Macphail's claims and suggest strategies researchers can use to make more advances in the field.

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Individual recognition is the ability to differentiate between conspecifics based on their individual features. It forms the basis of many complex communicative and social behaviours. Here, we review studies investigating individual recognition in the auditory and visual domain in birds. It is well established that auditory signals are used by many birds to discriminate conspecifics. In songbirds, the neuronal structures underpinning auditory recognition are associated with the song system. Individual recognition in the visual domain has mainly been explored in chickens and pigeons, and is less well understood. Currently it is unknown which visual cues birds use to identify conspecifics, and whether they have cortical areas dedicated to processing individual features. Moreover, whether birds can recognise themselves visually, as evidenced by mirror self-recognition, remains controversial. In the auditory domain, the responses of neurons in the song system suggest identification of the bird's own song. The surveyed behavioural and neural findings can provide a framework for more controlled investigations of individual recognition in birds and other species.

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After re-reading Macphail's (1987) essay "The Comparative Psychology of Intelligence" with all the associated commentaries, I was struck by how contemporary many of the arguments and counter-arguments still appear. Of course, we now know much more about the abilities of many more species (including their neurobiology) and fewer researchers currently favor explanations of behavior based solely on associative processes; however, the role of contextual variables in comparative psychology still remains cloudy. I discuss these issues briefly. Given my research interests involving the cognitive and communicative abilities of Grey parrots, the one aspect of the original article upon which I feel I can comment in depth involves Macphail's claims about the importance of language-and specifically syntax-in problem-solving and thus in placing humans above all other creatures. Granted, no other species has (or in my opinion is likely ever to acquire) everything that goes into what is considered "human language." Nevertheless, several other species have acquired symbolic representation, and considerable information now exists upon which to base an argument that such acquisition by itself enables more complex and "human-like" cognitive processes. Such processes may form the basis of the kind of intelligence that is measured-not surprisingly-with human-based tasks, including the use of such representations as a means to directly query non-human subjects in ways not unlike those used with young children.

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Studying animal cognition is meaningful because it helps us understand how animals adapt to the natural environment. Many birds build nests, clean their nests and reject foreign objects from their nests, which provide an optimal opportunity for studying their cognition toward foreign objects in nests. However, hand-made models used in previous studies have many deficiencies that considerably constrain our capacity to understand the evolution of avian cognition of foreign objects because they are unquantifiable and dependent on different features. We established a 3D modelling and printing method to manipulate one geometric dimension of a model while controlling for others, which allowed us to investigate avian cognition for different dimensions independently. Here we introduce this method, conduct an empirical study as an example, and discuss its applications to further studies.

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Impulsivity-the extent to which a reward is devalued by the amount of time until it is realized-can be affected by an individual's current energetic state and long-term developmental history. In European starlings (Sturnus vulgaris), a previous study found that birds that were lighter for their skeletal size, and birds that had undergone greater shortening of erythrocyte telomeres over the course of development, were more impulsive as adults. Here, we studied the impulsivity of a separate cohort of 29 starlings hand-reared under different combinations of food amount and begging effort. The task involved repeated choice between a key yielding one pellet after 3 s and another key yielding two pellets after 8 s. Impulsivity was operationalised as the proportion of choices for the short-delay option. We found striking variation in impulsivity. We did not replicate the results of the previous study concerning developmental telomere attrition, though combining all the evidence to date in a meta-analysis did support that robustness of that association. We also found that early-life conditions and mass for skeletal size interacted in predicting impulsivity. Specifically, birds that had experienced the combination of high begging effort and low food amount were less impulsive than other groups, and the usual negative relationship between impulsivity and body mass was abolished in birds that had experienced high begging effort. We discuss methodological differences between our study and studies that measure impulsivity using an adjusting-delay procedure.

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