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Inferential Learning of Serial Order of Perceptual Categories by Rhesus Monkeys (Macaca mulatta)

Category learning in animals is typically trained explicitly, in most instances by varying the exemplars of a single category in a matching-to-sample task. Here, we show that rhesus macaques can learn categories by a transitive inference paradigm in which novel exemplars of five categories were presented throughout each training session. Instead of requiring decisions about a constant set of repetitively presented stimuli, we studied the macaques ability to determine the relative order of multiple exemplars of particular stimuli that were rarely repeated. Ordinal decisions generalized both to novel stimuli and, as a consequence, to novel pairings. Thus, we showed that rhesus monkeys could learn to categorize on the basis of implied ordinal position, and that they could then make inferences about category order. Our results challenge the plausibility of association models of category learning and broaden the scope of the transitive inference paradigm.\n\nSignificance StatementThe cognitive abilities of non-human animals are of enduring interest to scientists and the general public because they blur the dividing line between human and non-human intelligence. Categorization and sequence learning are highly abstract cognitive abilities each in their own right. This study is the first to provide evidence that visual categories can be ordered serially by macaque monkeys using a behavioral paradigm that provides no explicit feedback about category or serial order. These results strongly challenge accounts of learning based on stimulus-outcome associations.

animal behavior and cognition↗

Sex differences in vocalizations to familiar or unfamiliar females in mice

Mice, both wild and laboratory strains, emit ultrasound to communicate. The sex differences between male to female (male-female) and female to female (female- female) ultrasonic vocalizations (USVs) have been discussed for decades. In the present study, we compared the number of USVs emitted to familiar and unfamiliar females by both males (male-female USVs) and females (female-female USVs). We found that females vocalized more to unfamiliar than to familiar females. In contrast, males exhibited more USVs to familiar partners. This sexually dimorphic behavior suggests that mice change their vocal behavior in response to the social context, and their perception of the context is based on social cognition and memory. In addition, because males vocalized more to familiar females, USVs appear to be not just a response to novel things or individuals, but also to be a social response.

animal behavior and cognition↗

Sex- and estrous-specific effects of paradoxical (REM) sleep deprivation: Neurobehavioral changes and hippocampal neuroinflammation

With millions suffering from sleep disorders in todays modern society, a better understanding of sleep disruption related cognitive outcomes is urgently needed. To that end, a preclinical investigation into the effects of sleep deprivation (SD) on neurobehavioral outcomes and associated hippocampal neuroinflammation was conducted in male and female rats. Due to epidemiological identification of sex differences in many aspects of sleep disorders, sex and estrous cycle stage factors were investigated. Sprague-Dawley rats underwent 120 h of paradoxical SD using a modified multiple platform flower pot method. At 96 h SD, animals were trained on neurobehavioral Novel Object Recognition (NOR) and Passive-Avoidance Task (PAT) paradigms. Prior to NOR/PAT testing at 120 h SD, Elevated Zero Maze (EZM) assessed anxiolytic-like behavior. SD impaired PAT performance among males and females. Anxiolytic-like and locomotor behavior was increased and NOR performance impaired in males after SD. Estrous cycle stage determined by cytological analysis of daily wet smears found females to exhibit estrous-specific differences across all neurobehavioral paradigms, with increased anxiolytic-like behavior and impaired PAT performance only among SD females in estrus. Immunohistochemical analysis of the hippocampus after 120 h SD found microgliosis, but not astrogliosis, in the CA1/2 of males and females in estrus. This study contributes to a better understanding of sex- and estrous-specific difference in sleep disruption induced neurobehavioral outcomes and associated hippocampal inflammation. Further research is needed to investigate the molecular mechanisms underlying the interaction between estrous cycle, hippocampal microgliosis, and sleep disruption cognitive outcomes.

animal behavior and cognition↗

At the roots of interference control: Conflict task in rats reveals the commonalities of onto- and phylo-genetic development

Responding to internal goals and filtering irrelevant environmental cues is essential for adapted behavior. To better understand their phylogenetic evolution, we explored the underlying mechanisms in rats, thanks to the adaptation of a well-established conflict task in Humans. Besides mean performance, state of the art data analysis based on distribution analysis and formal modeling (diffusion model adapted to conflict tasks that proved very powerful in Humans) revealed mechanisms similar to the ones observed in human adults, grounding any theoretical explanation into an evolutionary perspective. Besides, the dynamics of the underlying processes in this simple conflict task resembles the ones observed in human children performing a more challenging conflict task, including the tendency to loose goals. The present study bridges the gap between onto- and phylo-genetic development of cognitive control, opening new perspectives to understand both their functional and neural implementation.

animal behavior and cognition↗

The resource-rational dynamics of evidence accumulation

Evidence accumulation is a fundamental aspect of human decision-making. However, how the precise temporal structure of evidence shapes the accumulation process has not been systematically studied. As a result, current understanding of evidence accumulation remains largely limited to its time-averaged behavior. We tested human subjects in a visual estimation task in which they inferred the angular position of an unknown source from a noisy stimulus sequence. Introducing systematic temporal perturbations, i.e., breaks of different durations and at different positions in the otherwise regular evidence sequence, revealed that subjects actively compensated for the memory loss endured during the break by dynamically enhancing evidence integration and memory maintenance immediately after the break. We derived a new time-continuous Bayesian updating model that is dynamically constrained by optimal performance-effort trade-offs. With two free parameters determining the overall resource-efficiencies of encoding and memory maintenance, the model accurately predicts the rich dependencies of subjects accumulation behavior on the evidence schedule, including subjects individual tendencies to emphasize either early (primacy) or late (recency) samples in the evidence sequence. Our results suggest that evidence accumulation is a non-stationary, dynamically controlled process that optimally balances the information gained from incoming evidence against the cognitive effort required to acquire and maintain it. The proposed model is general and should apply broadly across many task domains.

animal behavior and cognition↗

CapuchinAI: Leveraging AI to study wild primate cognition

Advancing the study of primate cognition requires methods that preserve ecological validity while enabling the experimental control typical of laboratory research. We introduce CapuchinAI, a field-deployable touchscreen system that integrates real-time facial recognition with automated cognitive testing, providing a novel methodological framework for studying cognition in wild primates. Our approach combines a high-performing (>97% accuracy) YOLOv7-based facial recognition model (Multiple Capuchins v1.0) with a portable Raspberry Pi-driven touchscreen-reward apparatus designed for automated operation in natural habitats. The system detects approaching capuchins, initiates video recording, presents touchscreen stimuli, and dispenses food rewards contingent on task performance. During a two-week presentation to two habituated groups of wild white-faced capuchins (Cebus imitator) at the Taboga Forest Reserve, 16 individuals voluntarily interacted with the apparatus, 10 learned to trigger rewards, and 8 formed and retained robust screen-reward associations. The rapid habituation and learning rates demonstrate the feasibility of deploying AI-mediated cognitive experiments in the wild. CapuchinAI addresses several long-standing challenges in field cognition research by enabling: (1) autonomous, individualized task administration without researcher intervention; (2) standardized, repeatable trials across individuals and sessions; (3) scalable deployment across groups and sites; and (4) parallel data collection on behavior, identity, and performance. This methodology provides a blueprint for integrating machine learning, touchscreen testing, and automated reward delivery to study within- and between-individual cognitive variation under natural conditions. CapuchinAI represents a significant step toward long-term comparative research on primate cognition by making laboratory experimental paradigms accessible in the wild, bridging the gap between lab and field. Research HighlightsO_LIWe present CapuchinAI, a field-ready touchscreen testing station that uses real-time facial recognition to study cognition in wild capuchin monkeys C_LIO_LIWe developed a YOLOv7-based facial recognition model (Multiple Capuchins v1.0) that identifies individual capuchins with >97% precision and recall from static images, video, and live footage, enabling fully automated, individualized testing in the wild. C_LIO_LIWe integrated a version of this model into a closed-loop touchscreen-reward pipeline that detects an approaching monkey, presents a basic learning task, and automatically delivers food rewards based on the monkeys responses. C_LIO_LIWild capuchins rapidly habituated and learned touchscreen-reward associations, showing that AI-enabled touchscreens provide a scalable field method for deploying lab-style cognitive tests and mapping individual differences across tasks, species, and sites. C_LI

animal behavior and cognition↗

Distinct networks of expressed genes are associated with neophobia in the hippocampus of male and female Eurasian tree sparrows (Passer montanus)

Neophobia, avoidance of novel stimuli, is an ecologically and evolutionarily relevant behavioral trait that varies among individuals and across species. Especially among wild animals, the neuromolecular mechanisms underlying individual variation in neophobia have not been well characterized. We examined three neophobic behaviors in captive female and male Eurasian tree sparrows (Passer montanus) from a wild population introduced to the USA in 1870: responses towards novel objects, novel foods, and repeated presentations of the same initially novel object. We compared transcriptomic patterns associated with neophobia in three brain regions, the striatum, dorsal hippocampus, and rostral hippocampus, using differential expression and co-expression network analyses. We found that the striatum and hippocampus had distinct transcriptomic profiles, as did the rostral and caudal subregions of the hippocampus, supporting recent hypotheses that these subregions are functionally specialized. Despite the absence of sex differences in neophobic behaviors, neophobia-associated gene modules revealed sex-specific patterns within brain regions. For females, neophobic behaviors more strongly correlated with gene modules in the caudal hippocampus, a region involved in stress and anxiety, whereas for males, neophobic behaviors correlated with gene modules in the rostral hippocampus, a region that may play a larger role in spatial cognition. These modules exhibited significant overlap, suggesting that neophobic behaviors in both females and males are driven by shared neurobiological mechanisms, though they exhibit sex-specific patterns of brain region localization. Further, this work highlights the importance of examining both male and female animals in neurobiological research.

animal behavior and cognition↗

A farewell to EQ: A new brain size measure for comparative primate cognition

Both absolute and relative brain size vary greatly among and within the major vertebrate lineages. Scientists have long debated how larger brains in primates and hominins translate into greater cognitive performance, and in particular how to control for the relationship between the non-cognitive functions of the brain and body size. One solution to this problem is to establish the slope of cognitive equivalence, that is the line connecting organisms with an identical bauplan but different body sizes. Here, we suggest that intraspecific slopes provide the best available estimate of this measure. This approach was abandoned because slopes were too low by an unknown margin due to estimation error. We control for the error problem by focusing on highly dimorphic primate species with large sample sizes and fitting a line through the mean values for adult females and males. We obtain the best estimate for the slope of ca 0.27, a value much lower than those constructed using all mammal species, and close to the value expected based on the genetic correlation between brain size and body size. We also find that the estimate of cognitive brain size based on cognitive equivalence fits empirical cognitive studies better than the encephalization quotient (EQ), which should therefore be avoided in future studies on primates, and presumably mammals and birds in general. The use of residuals from the line of cognitive equivalence may change conclusions concerning the cognitive abilities of extant and extinct primate species, including hominins.

animal behavior and cognition↗

Image classification and cognition using contour curvature statistics

Although the higher order mechanisms behind object representation and classification in the visual system are still not well understood, there are hints that simple shape primitives such as "curviness" might activate neural activation and guide this process. Drawing on elementary invariance principles, we propose that a statistical geometric object, the probability distribution of the normalized contour curvatures (NCC) in the intensity field of a planar image, has the potential to represent and classify categories of objects. We show that NCC is sufficient for discriminating between cognitive categories such as animacy, size and type, and demonstrate the robustness of this metric to variation in illumination and viewpoint, consistent with neurobiological constraints and psychological experiments. A generative model for producing artificial images with the observed NCC distributions highlights the key features that our metric captures and just as importantly, those that it does not. More broadly, our study points to the need for statistical geometric approaches to cognition that build in both the statistics and the natural invariances of the sensory world.

animal behavior and cognition↗

Social complexity affects cognitive abilities but not brain structure in a Poecilid fish

Complex cognitive performance is suggested to be the out-turn of complex social life, allowing individuals to achieve higher fitness through sophisticated "Machiavellian" strategies. Although there is ample support for this concept, especially when comparing species, most of the evidence is correlative. Here we provide an experimental investigation of how group size and composition may affect brain and cognitive development in the guppy (Poecilia reticulata). For six months, we reared sexually mature female guppies in one of three different social treatments: (i) three female guppies; (ii) three female guppies mixed with three female splash tetras (Copella arnoldi), a species that co-occurs with the guppy in the wild; and (iii) six female guppies. We then tested the guppies performance in inhibitory control, associative learning and reversal learning tasks to evaluate their self-control, operant conditioning and cognitive flexibility capabilities. Afterwards, we estimated their brain size and the size of major brain regions using X-ray imaging technology. We found that individuals in larger groups of six individuals, in both same and mixed species treatments, outperformed individuals from the smaller groups of three guppies in reversal learning, with no apparent differences in the inhibitory control and associative learning tasks. This is rare evidence of how living in larger social groups improves cognitive flexibility, supporting that social pressures play an important role in shaping individual cognitive development. Interestingly, social manipulation had no apparent effect on brain morphology, but relatively larger telencephalons were associated with better individual performance in reversal learning. This suggests alternative mechanisms beyond brain region size enabled greater cognitive flexibility in individuals from larger groups.

animal behavior and cognition↗

Stable long-term individual variation in chimpanzee technological efficiency

Using tools to access hard-to-reach and high-quality resources, such as termites, honey, and nuts, initiated a fundamental adaptive shift in human and nonhuman primate cognitive and behavioural evolution. Variation in the efficiency of extracting calorie-rich and nutrient-dense resources directly impacts energy expenditure, and potentially has significant repercussions for cultural transmission where model selection biases are employed during skill acquisition. Assessing variation in efficiency is key to understanding the evolution of complex behavioural traits in primates. Yet, individual-level differences beyond age- and sex-class in primate extractive foraging efficiency have never been investigated. Here, we used 25 years (1992- 2017) of video data of the Bossou chimpanzee community (Guinea), to investigate whether individual differences in nut-cracking efficiency exist across the life span of chimpanzees aged [≥] 6 years. Data from 3,882 oil-palm nut-cracking bouts from over 800 hours of observation were collected. We found long-term stable and reliable individual differences in four (out of five) measures of efficiency. We found no sex effect, challenging previous research on a female bias in chimpanzee tool use. These life-long differences in extractive foraging impacts daily energy budgets, which potentially have significant individual fitness and life history consequences. Additionally, the establishment of long-term individual variation in chimpanzee stone tool use has implications for interpreting archaeological records of hominins. Our findings highlight the importance of longitudinal data from long-term field sites when investigating underlying cognitive and behavioural diversity across individual lifespans and between populations.

animal behavior and cognition↗

Smells allow for tapping into episodic-like memory of minipigs

Episodic-like memory is considered as one of the most advanced cognitive capacities and could therefore have major implications on how animals with this capacity are treated in respect to their welfare. Yet, episodic-like memory has only been substantiated in a few vertebrate species. These species are quite diverse, originate from several taxa, and the question is raised whether episodic-like memory is indeed a rather specific capacity or whether this capacity turns out to be more common when further species are investigated. In general, intensively foraging animals that rely on some replenishing resources are promising species to investigate episodic-like memory. Here, we conducted a pilot study on minipigs (Sus scrofa f. domestica), replicating an approach used in dogs. Both, dogs and pigs are macrosmate species, making the comparison of their memory of olfactory stimuli promising. The minipigs learnt to differentiate the sequence of encountered location-smell combinations as easily as dogs. This makes minipigs a promising species and the use of smells a promising approach to investigate episodic-like memory in more detail. This study also hints towards a cognitive capacity of pigs, which raises further concerns regarding intensive and barren housing conditions.

animal behavior and cognition↗

Innate building blocks underly socially learned call sequences

Recent evidence of extensive call sequence use in non-human primates has led to the theory that syntax evolved to mitigate the constraints of their genetically fixed repertoires, before vocal production learning later emerged in humans. However, evidence of similarly extensive sequence repertoires in an open-ended vocal production learner--the Western Australian magpie (Gymnorhinatibicendorsalis)--offers a unique opportunity to explore potential alternative pathways to syntactic communication. Our previous work revealed fledgling magpies learn group-specific repertoires of structured call sequences from their social contacts, with more sociable individuals acquiring larger repertoires earlier in development. Notably however, the individual vocal segments that combine to form their calls and call sequences were shared across groups and emerged as early as the first week post-fledging--suggesting the underlying vocal elements may not be learned. Here we utilised acoustic neighbourhood-based dimensionality reduction to compare clustering patterns of vocal segments across magpie fledgling developmental stages, and between fledglings and adults. We found no evidence of acoustic development over time, and no significant distinction between fledgling and adult productions of the same vocal segments. The same coarticulatory effects--where a vocal element is produced differently when combined with another--and geographic variation established previously in adults were supported in fledglings too. These findings support that the vocal building blocks underpinning magpie call sequences are innate, suggesting usage learning better explains how fledglings learn to combine calls. In a species capable of open-ended production learning, this suggests learning to combine existing signals may be more adaptive than productively learning new ones. Rather than evolving solely to compensate for genetically fixed repertoires, syntax may have evolved as a flexible, convergent solution to the various challenges of expanding communicative capacity--whether due to genetic constraints, cognitive limitations or the cost of establishing new meaning in novel signals.

animal behavior and cognition↗

DigiFish: A flexible open-source tool for animating realistic virtual fish and reconstructing visual fields from tracking data

1. For many animals, vision represents the primary sensory modality governing a vast diversity of behaviours. Creating 'virtual twins', or visually realistic digital reconstructions of real individuals interacting within their environments, offers a promising approach for understanding how visual information guides an animal's decisions, behaviours, and interactions. Such systems enable the creation of video scenes from any view, including the viewpoint of the animals themselves. This allows receiver information to then be analysed from the perspective of species-specific visual systems, or for the accurate reconstruction of a real animal's movement to provide social stimuli for behavioural experiments. 2. Despite advances in 3D modelling, photorealistic scene rendering, and animal tracking, efficiently creating realistic digital reconstructions of movement remains challenging for many species. In fishes, a continuously-deforming body due to the absence of discrete joints make it particularly difficult to map tracked body keypoints onto articulated digital models, often resulting in unnatural or erratic animations. 3. Here, we introduce DigiFish, an add-on for the open-source animating software Blender, that generates realistic fish animations from tracking data. By integrating anatomical keypoint tracking with spline-based body curvatures, DigiFish reconstructs biologically-realistic swimming postures and movements of fishes within customisable virtual aquatic environments. The system also supports virtual 'point-of-view' cameras that can be placed arbitrarily within a scene, or dynamically driven by eye-tracking data, enabling unlimited perspectives on how individuals, conspecifics, or landmarks appear against complex backgrounds and how these views change with realistic swimming and eye movements. 4. We provide the Blender add-on and user guide and use DigiFish to reconstruct two different contexts: staged predator-prey interactions in the laboratory and natural schooling of wild fish in the field. Using a further reconstruction of fish swimming in an arena, we also use an Elementary Motion Detector model to show that our virtual reconstructions successfully capture perceptually-realistic movements and postures of individual fish within a digitally reconstructed scene. 5. By integrating photo-realistic scenes with biologically-realistic, simulated motion, DigiFish offers a flexible and accessible platform for investigating visually guided behaviour, perception, and cognition in complex and dynamic environments.

animal behavior and cognition↗

Flexible visual learning and memory in nectar foraging hornets

Pollinators, such as bees, develop flexible memories of colours, patterns and shapes, for efficient flower recognition. Here we tested whether other insects facing the same foraging problem have evolved similar cognitive abilities for flexible learning. We trained wild hornets from two species commonly found in Europe (Vespa velutina nigrithorax and Vespa crabro) to associate sucrose solution rewards to colour stimuli in a Y maze. Hornets of both species succeeded in differential and reversal learning, and developed short and long-term memories of the learnt associations. Thus, just like bees, hornet foragers can learn various visual cue-reward associations and remember them over several hours or days for selecting flowers. Our study in non-model species illustrates how standard conditioning approaches can be used to explore and compare the cognitive abilities of animals with similar foraging ecology.

animal behavior and cognition↗

Early- but not late-adolescent Western diet consumption programs for long-lasting memory impairments in male but not female rats

Early life Western diet (WD) consumption leads to impaired memory function, particularly for processes mediated by the hippocampus. However, the precise critical developmental window(s) during which WD exposure negatively impacts hippocampal function are unknown. Here, we exposed male and female rats to a WD model involving free access to a variety of high-fat and/or high-sugar food and drink items during either the early-adolescent period (postnatal days [PN] 26-41; WD-EA) or late-adolescent period (PN 41-56; WD-LA). Control (CTL) rats were given healthy standard chow throughout both periods. To evaluate long-lasting memory capacity well beyond the early life WD exposure periods, we performed behavioral assessments after both a short (4 weeks for WD-EA, 2 weeks for WD-LA) and long (12 weeks for WD-EA, 10 weeks for WD-LA) period of healthy diet intervention. Results revealed no differences in body weight or body composition between diet groups, regardless of sex. Following the shorter period of healthy diet intervention, both male and female WD-EA and WD-LA rats showed deficits in hippocampal-dependent memory compared to CTL rats. Following the longer healthy diet intervention period, memory impairments persisted in male WD-EA but not WD-LA rats. In contrast, in female rats the longer healthy diet intervention reversed the initial memory impairments in both WD-EA and WD-LA rats. Collectively, these findings reveal that early-adolescence is a critical period of long-lasting hippocampal vulnerability to dietary insults in male but not female rats, thus highlighting developmental- and sex-specific effects mediating the relationship between the early life nutritional environment and long-term cognitive health.

animal behavior and cognition↗

The prostate cancer therapy enzalutamide compared with abirateroneacetate/prednisone impacts motivation for exploration, spatial learning andalters dopaminergic transmission in aged castrated mice

Cognitive side effects after cancer treatment, threatening quality of life (QoL) and adherence to treatments, now constitute a major challenge in oncology. Abiraterone acetate plus prednisone (AAP) and enzalutamide (ENZ) are next generation therapy (NGT) administered with androgen deprivation therapy to metastatic castration-resistant prostate cancer (mCRPC) patients. NGT significantly improved mCRPC overall survival but neurological side effects such as fatigue and cognitive impairment have been recently reported. We developed a behavioral 17 months-aged and castrated mouse model receiving per os AAP or ENZ during 5 days per week for six consecutive weeks. After behavioral tests, brain and plasma were collected for immunohistochemical studies. The objective was to elucidate the impact of NGT on spontaneous activity, cognitive functions and emotional reactivity, as well as neurobiological functions. ENZ exposure reduced spontaneous activity and exploratory behavior associated with a decreased tyrosine hydroxylase (TH)-dopaminergic activity in the substantia nigra pars compacta and the ventral tegmental area. A decrease in TH+-DA afferent fibers and Phospho-DARPP32-related dopaminergic neuronal activities in the striatum and the ventral hippocampus, highlighted ENZ-induced dopaminergic regulation whithin the nigrostriatal and mesolimbocortical pathways. ENZ and AAP treatments did not substantially modify spatial learning and memory or behavioral flexibility performances, but ENZ led to a thygmotaxis behavior impacting the cognitive score, and reduced c-fos-related activity of NeuN+-neurons in the dorsal hippocampus. These results establish the consequences of the mCRPC treatment ENZ in aged castrated mouse motivation to exploration and cognition, of particular importance for future management of patients elderly postrate cancer patients and their QoL.

animal behavior and cognition↗

Does training method matter?: Evidence for the negative impact of aversive-based methods on companion dog welfare

There is a growing number of dogs kept as companion animals, and the methods by which they are trained range broadly from those using mostly positive punishment and negative reinforcement (aversive-based methods) to those using primarily positive reinforcement (reward-based methods). Although the use of aversive-based methods has been strongly criticized for negatively affecting dog welfare, these claims do not find support in solid scientific evidence. Previous research on the subject lacks companion dog-focused research, investigation of the entire range of aversive-based techniques (beyond shock-collars), objective measures of welfare, and long-term welfare studies. The aim of the present study was to perform a comprehensive evaluation of the short- and long-term effects of aversive- and reward-based training methods on companion dog welfare. Ninety-two companion dogs were recruited from three reward-based (Group Reward, n=42) and four aversive-based (Group Aversive, n=50) dog training schools. For the short-term welfare assessment, dogs were video recorded for three training sessions and six saliva samples were collected, three at home (baseline levels) and three after the training sessions (post-training levels). Video recordings were then used to examine the frequency of stress-related behaviors (e.g., lip lick, yawn) and the overall behavioral state of the dog (e.g., tense, relaxed), and saliva samples were analyzed for cortisol concentration. For the long-term welfare assessment, dogs performed a cognitive bias task. Dogs from Group Aversive displayed more stress-related behaviors, spent more time in tense and low behavioral states and more time panting during the training sessions, showed higher elevations in cortisol levels after training and were more pessimistic in the cognitive bias task than dogs from Group Reward. These findings indicate that the use of aversive-based methods compromises the welfare of companion dogs in both the short- and the long-term.

animal behavior and cognition↗