Animal Cognition

Spatiotemporal dynamics of behavior is key to understanding the organism-environment relationship. While often implicitly addressed, its relation to Pavlovian contingencies remains understudied in domestic dogs. This study quantitatively examined the spatial dimension of behavior of three freely moving dogs under pairing and extinction of a tone-food Pavlovian contingency. In pairing, a tone (CS) was paired with food delivery (US) on a fixed-time 60-second (FT 60 s) schedule; in extinction, only the tone was presented. Locomotion was recorded using two-dimensional tracking based on center of mass. During pairing, dogs moved closer to the dispenser, covered greater distances, displayed extended trajectories, and showed a conditional approach pattern to the dispenser during CS presentation. In extinction, they stayed closer to the owner or room periphery, traveled shorter distances, and exhibited more restricted trajectories. These findings show that spatial segments (dispenser area) are integrated into the CS-US relationship, demonstrate the usefulness of continuous recording of spatial behavior in the analysis of Pavlovian contingencies, and suggest potential application in contexts relevant to animal welfare. HighlightsO_LISpatial behavior remains largely unexplored in studies of Pavlovian contingencies C_LIO_LIDog locomotion was continuously tracked during pairing and extinction phases C_LIO_LIDistinct spatial patterns emerged across pairing and extinction phases C_LIO_LIIncorporating space into CS-US relations clarifies behavioral organization C_LIO_LITracking tools expand the experimental and applied scope of Pavlovian research C_LI

Traditionally, primates have been considered primarily visual animals. However, studies across a variety of taxa suggest that, in the context of food evaluation, the reliance on this sense might be more nuanced that previously thought, with dietary specialization and food item properties leading to differences in sensory prioritization. We performed a field-based study assessing the use of sensory cues during food evaluation as well as food-related behaviours such as muzzle contact in two mixed-sex groups of wild vervet monkeys including three age classes over a period of five months (nmonkeys = 44). Using a total of 18868 food evaluation observations collected over 44 hours of focal follows, we found that vervets mainly relied on their sense of vision when evaluating food (96.8% of all instances). Sensory usage varied according to food category and sex differences were only observed in the use of smell for a subset of these. Juveniles initiated muzzle contact and used tactile inspection more often than adults whereas females received muzzle contact more often than males. In addition, the low rejection rates suggest that most food items were familiar to the vervets regardless of age and sex. These findings are in line with optimal foraging theory according to which the food evaluation process should be adapted to the familiarity of food items and allows individuals to maximize their intake of energy and critical nutrients, while minimizing the time and effort in food evaluation.

Reports of honeybees demonstrating abstract concepts like sameness and difference marked a pivotal development in comparative psychology. Subsequent studies expanded the scope of concept learning in honeybee cognition, yet most evidence relies on a single method: the delayed-matching-to-sample task using a Y-maze. Whether this setup is uniquely effective or if alternative approaches could yield similar results remains unresolved. Additionally, the failure of bumblebees (Bombus spp.) to complete this task, despite honeybees demonstrating success, remains unexplained. This study compared the performance of honeybees (Apis mellifera) and bumblebees (Bombus terrestris) across matching-to-sample tasks with varying degrees of physical continuity between sample and target stimuli. The objectives were twofold: to evaluate an alternative method for assessing concept learning in both species and to investigate potential species differences in such tasks. Contrary to prior findings, neither species succeeded at the reported proficiency levels in simultaneous matching-to-sample tasks. Moreover, bumblebees outperformed honeybees in one task. These results are consistent with an explanation based on species-specific differences in visual attention mechanisms, and underscore the need for further research on concept learning in social bees.

There is evidence that appetites for specific nutrients can guide foraging behaviour and aid in dietary regulation through associative learning processes that link stimuli to nutrient-specific outcomes. However, most, if not all, examples of such behaviour can be interpreted as being stimulus-bound habits, i.e., reflexive responses induced by environmental stimuli. The control of identified goal-directed actions by nutrient-specific appetites has not been directly assessed. To address this question, we trained rats to press a lever for a high protein reward (whey protein shake) and another lever for a high carbohydrate reward (polycose solution). They were then tested under extinction conditions in which both levers were available following the extended exposure to meals that were high in protein or carbohydrate. When otherwise food-deprived rats had been selectively satiated on protein immediately prior to test, they pressed more on the lever they learned had produced polycose, whereas they pressed the lever they learned had produced whey protein more if they had instead been satiated on carbohydrate. Crucially, the same pattern emerged whether the satiety manipulation was achieved using the same nutrient sources that rats had earned during training (i.e., whey or polycose) or with foods high in the relevant nutrients, indicating that these behaviours were under goal-directed control and sensitive to nutritional state. These results show that actions can be motivated by the nutritional relevance of the instrumental outcome to specific appetites, a relationship that may guide natural foraging decisions.

Urbanisation is one of the most important forms of human-driven landscape change, altering wildlife populations in unprecedented ways. In terms of behaviour, for example, urbanisation is hypothesised to increase the likelihood of observing urban populations touching, exploring, and solving novel food-related tasks compared to rural areas. However, little is known about the impact of spatiotemporal patterns of urbanisation, particularly historical patterns of change, on these behaviours. We tested this in the worlds most urbanised carnivore, the red fox (Vulpes vulpes), by introducing novel food-related tasks (puzzle feeders) to 284 sites throughout Great Britain. We compared tactile and problem-solving behaviours in rural populations, recently colonised urban populations, and long-established urban populations (>40 years). Foxes from 27.4% of locations touched the tasks, foxes from 12.4% of locations solved them. Urban foxes were more likely to touch tasks compared to rural populations. Exploration time, exploratory diversity, and latency to touch tasks did not significantly differ across urban and rural locations. Urbanisation rate from 1994 to 2020 (26 years) did not significantly predict the likelihood of foxes touching or solving tasks across locations. Older urban populations - particularly from London - spent more time exploring tasks and displayed greater exploratory diversity and higher problem-solving success, despite more recent urban populations being equally likely to touch them. Collectively, our findings suggest that certain population characteristics, such as the likelihood of touching/engaging with novelty, potentially emerge early in urbanisation while other characteristics, such as greater exploratory and innovative behaviours, may emerge after long-term urban exposure across many decades. HighlightsO_LIHistorical impacts of urbanisation on wild animal behaviour are unclear. C_LIO_LIWe tested this with wild red foxes responses to novel food objects. C_LIO_LIUrban foxes were more likely to touch and exploit objects, especially from London. C_LIO_LIOlder urban foxes displayed more exploratory and innovative behaviours. C_LIO_LILength of urban exposure may help predict behavioural responses to novelty. C_LI

Group living often improves decision accuracy in animals, yet whether increasing group size buffers against context-dependent biases remains unexplored. One such bias is the decoy effect, where the presence of a third option shifts preferences between two alternatives. Here, we tested whether introducing a decoy shoal influences collective preference for the larger female shoal in adult male zebrafish (Danio rerio), and whether the strength of this effect depends on group size. Groups of two, three, or four males were presented with female shoals under two choice contexts: a dichotomous contrast (four vs. two fish) and a trichotomous contrast including an additional alternative of one, three, or five fish. The order of presentation (dichotomous-first or trichotomous-first) was counterbalanced, and multi-animal tracking was used to quantify group-level shoal-size preference (time spent near shoals), inter-individual distance (IID), polarization, and swimming speed. In the dichotomous-first order, across all group sizes, subject shoals consistently showed a baseline preference for the larger shoal. Adding a third decoy option altered this preference only when the decoy was extreme (one or five fish relative to the 4 vs. 2 alternatives), reducing relative preference toward indifference. Group IIDs were associated with context-dependent shifts in relative preference in the dichotomous-first order, whereas polarization and swimming speed were not. In the trichotomous-first order, groups showed no preference for the larger shoal, with or without a decoy shoal. Our results demonstrate that context-dependent biases, shape collective shoal choice, with effects driven by choice structure and order of presentation of options than by group size.

Foraging bee decision-making research has focused on choice determinants, and the variability and underlying causes of pre-choice latency remain understudied. Here, individual bumblebees (Bombus impatiens) were trained to associate one colored stimulus with a medium-value reward and another with a novel, higher-value reward. The experimental design consists of seven blocks, each containing four consecutive single-stimulus presentations followed by a forced binary choice. The latency to choose a stimulus and the type of choice during dual-choice trials were analysed. In dual-choice trials, bees in the yellow-high reward group showed a slower increase in high-reward selection than those in the blue-high group, suggesting persistent innate color bias. Response latencies for the low-reward stimulus systematically increased across blocks, indicating progressive devaluation. Early learning phases showed a temporary increase in response latency, extending previous findings on experience-dependent adjustments in acceptance thresholds. Latency in single-stimulus trials correlated with binary choice results, though choice proved a stronger indicator of preference than latency. Certain options elicited faster responses when presented with an alternative than when presented alone. Together, these findings support a deliberative model of bumblebee decision-making, in which pre-choice latency is modulated by innate preferences, associative learning, and immediate context.

The midsession reversal (MSR) task is frequently used to study behavioral flexibility and decision strategies in animals. In a typical version of the task, subjects complete 80 trials in which they choose between two simultaneously presented stimuli, S1 and S2. During the first 40 trials, responses to S1 are reinforced, whereas responses to S2 are not. The contingencies then reverse without warning: From trial 41 to 80, only responses to S2 are reinforced. In birds, performance in this task is often characterized by anticipatory and perseverative errors around the reversal point, suggesting a reliance on elapsed time since the session began. In contrast, rats tested in operant conditioning chambers typically show near-optimal performance with few errors, a pattern often interpreted as evidence that rats rely primarily on local reinforcement cues rather than temporal information. The present study investigated whether rats exclusively rely on local cues in the MSR task or whether temporal information also contributes to the decision process. Two groups of rats were trained with different intertrial intervals (ITIs; 5 s or 10 s) while the reversal point remained fixed at Trial 41. During acquisition, both groups diplayed similar learning rates and near-optimal steady-state performance with minimal anticipatory or perseverative errors. However, when the ITI was manipulated in probe sessions, systematic shifts in switching behavior emerged. Rats adjusted their choices according to the temporal midpoint experienced during training rather than the nominal trial number of the reversal. These results suggest that rats rely on a mixed strategy that integrates local reinforcement cues with global timing information. Temporal control may therefore be present even when it is not expressed during standard training conditions.

Optimal foraging behavior is a key component of successful adaptations to natural environments. Understanding how animals decide to stay near food or to leave it for another food patch gives us insights into the underlying cognitive mechanisms that govern adaptive behaviors. 3D pose tracking was used to determine how pigeons exploit a 4 square meter arena with two separate platforms (i.e. food patches) whose absolute and relative elevations were manipulated. Detailed kinematic features of foraging and traveling behaviors were quantified using automated video tracking, without a need for manual coding. Our computational approach captured continuous, high-dimensional movement patterns and enabled precise quantification of travel costs between patches. Combined with mixed-effects survival analysis, our fine-grained behavioral tracking provided detailed insight into the moment-by-moment dynamics of patch-leaving decisions of pigeons. As expected from behavior optimization models, our results showed a preference to visit a ground food platform first, and longer latencies to leave an elevated platform. Foraging activity significantly decreased throughout the session, with shorter visits, less pecks per visit, and a decrease in inter-peck variability. However, a mixed-effects Cox regression modeled pigeons patch-leaving probability, demonstrating that current and cumulative foraging parameters between patches significantly enhanced the models predictive power beyond patch accessibility (i.e., beyond travel costs). This suggests that pigeons integrate both current environmental cues and their individual foraging history when making patch-leaving decisions. Our findings are discussed in relation to the marginal value theorem and optimal foraging theory.

Communication allows organisms to quickly convey information vital for survival or fitness. Chemical communication and speed-accuracy trade-offs are ubiquitous in animal decision making. Most studies have used species which forage mainly above-ground species, tested in an epigean setting, but it remains unclear whether below-ground species behave similarly. Here, we use the below-ground ant Tetramorium alpestre to assess the efficacy of above- vs. below-ground mazes, the accuracy of decisions when using natural vs. artificial pheromones, the presence of a speed-accuracy trade-off, and the pheromones effect on aggression. Ants decided more quickly under below-ground than above-ground conditions, indicating they may be distracted by above-ground stimuli. Ants followed both natural and artificial trails but in direct competition preferred artificial trails, likely due to a higher pheromone concentration. Surprisingly, no speed-accuracy trade off was observed during path decision-making. Lastly, population origin but not pheromones affected if and how aggression occurred in presence of trail and home-range marking pheromones. We argue that the design of behavioural tests should match the lifestyle of the focal organism. We further speculate that speed-accuracy trade-offs likely are highly species- and context-specific and other factors besides chemicals seem important to trigger aggression, at least in this species.

Many animals use the Earths magnetic field as a directional reference, for long-distance, but also for local movements. Among those are amphibians which can be trained along the y-axis (the shore - deep gradient) in their aquatic environment. We used a light gradient (light-dark axis) to train larvae of the European green toad (Bufotes viridis) towards a magnetic direction, testing their magnetoreception ability. After training we tested the individual animals four times, in the four cardinal magnetic field directions (N, E, S, W). Recent developments in circular statistics allowed us to use the larvaes responses in a mixed effects models (the individual as the random factor) and tease the overall and individual responses apart. We used control simulations to test whether the mixed effects models could produce false positive findings, which confirmed the validity of this approach. Our results clearly show a trained magnetic compass response for the green toad, adding another animal to the list of magnetoreceptive animals. Interestingly, when analyzing just the first choice after release, there was only a magnetic effect. However, over the course of the entire 2 minutes trial, animals also showed untrained magnetic as well as non-magnetic responses, highlighting the complexity of small-scale animal orientation, with many interacting cues and motivations. Switching to repeated measures experimental design together with the newly developed circular statistical approaches can therefore be used to better understand the entirety of the animal orientation strategies, going beyond the overall effect. Our approach has the potential to study different aspects of animal orientation in the same experiment (i.e., magnetic alignment and trained magnetic effects) and therefore bridge the gaps between different lines of research.

The ability to prepare for mutually exclusive outcomes is often considered uniquely human. Solving such problems requires anticipating alternative futures before acting. In the classic forked-tube task, the optimal strategy is to block both exits to secure a reward: children under four years and great apes typically fail, whereas older children succeed. Using this paradigm, we tested three Asian small-clawed otters (Aonyx cinereus) and one Eurasian otter (Lutra lutra) and compared their performance with chimpanzee data. Otters covered both exits significantly more often than chimpanzees, with all individuals succeeding within their first trials. Initial inconsistency in maintaining the strategy appeared linked to anatomical constraints that limited reward success. When retested two months later with an apparatus better suited to otter morphology, individuals adopted and maintained dual coverage as success increased, indicating that the behavior tracked the payoff structure of the task rather than reflecting low-level mechanisms such as trial-and-error learning. Together, these findings indicate that blocking both exits is an adaptive response to the tasks causal structure, supporting the ecological intelligence hypothesis: cognition evolves in response to ecological demands, particularly foraging challenges that place recurrent pressures on memory, decision-making, and executive control, rather than being driven solely by social complexity.

Object play - seemingly non-functional interactions with objects - can promote the development of foraging skills, tool use, and behavioral innovation. Among Catarrhine monkeys, it was described in macaques and baboons. Wild geladas, although closely related to baboons, have been described as lacking object play (observed only in captivity) linked to their specialized grazing ecology. Here, we provide the first evidence of both social and solitary object play in a wild gelada population (NOMUs=13) at Debre Libanos (Ethiopia) and compare it with object play in sympatric olive baboons (Nindividuals=42). Notably, immature geladas engaged in object play both socially and solitarily, but the latter case was most frequent also with novel objects introduced by researchers. Solitary object play occurred at levels comparable to those of baboons, challenging previous reports of limited object interest in geladas. This finding aligns with the occurrence of object play in phylogenetically related species and with the retention in wild geladas of arboreal behavior and fruit consumption and hand morphology enhancing fine manipulation. Hence, object play in geladas under certain environmental conditions may reflect a biologically rooted capacity and underscores the importance of ecological variability, as distinct behavioral ecotypes can emerge across different populations of the same species.

Despite growing evidence that many animals can evaluate quantities, the ecological relevance of numerical cognition remains debated, particularly outside vertebrates. Would individuals still rely on numerousness if less computationally demanding cues, visual features extracted at the early stage of visual processing, were available to assess quantity? In primates, individuals show a numerical bias as they tend to rely on the number of items rather than non-numerical cues, such as total area, to categorize quantities. In this study, we trained free-flying honeybees to discriminate between two and four items in conditions where numerosity covaried with the total area and perimeter (Experiment Size) or the convex hull (Experiment Space) cues, mimicking ecological contexts. Transfer tests assessed which numerical or non-numerical cues were learned and preferentially used by the bees. Bees primarily relied on numerousness over these non-numerical cues. Individual analyses revealed two consistent strategies: a "numerical bias" strategy, in which bees encoded numerical information while ignoring non-numerical cues, and a "generalist" strategy, where bees flexibly switched between cues and favored non-numerical information when cues conflicted. We further reported improved discrimination when smaller quantities appeared on the left and larger ones on the right, consistent with an oriented mental number line. Together, these findings demonstrate a spontaneous numerical bias in honeybees and reveal that individuals within the same species can adopt distinct strategies when evaluating quantity. Our findings also suggest that distantly related taxa like bees and primates may have independently evolved comparable mechanisms for quantity evaluation.

Sentinel behaviour occurs when individuals use raised positions to scan for predators while the rest of the group forages. Here, we investigated whether a colonial cooperatively breeding species that forages in large groups, the sociable weaver, Philetairus socius, displays sentinel behaviour. This behaviour has been reported in species with similar ecology, behaviour and foraging habits, (e.g. ground foraging in open habitats where aerial predators are common) and, hence, we expected that it could occur in sociable weavers. On the other hand, sentinel behaviour appears to be less common in species that live in very large groups. We used an experimental set-up consisting of an artificial feeding station and perches to assess occurrence of sentinel related behaviours: (i) perching events > 30s on an elevated position, (ii) head-movements and (iii) alarm calling. Birds were seldom observed perching while others fed, and those that did, perched for periods that were too short to be considered as sentinel behaviour (less than 5s on average). Our results suggest that this behaviour is uncommon or even absent in sociable weavers. We discuss whether other factors such as foraging in very large groups, or interspecific foraging associations might make sentinel behaviour less important in this species.

From facial expressions to gestures, animals use multiple signal modalities to express emotions and communicate. In dogs, tail movements are conspicuous behaviours associated with emotional states, but this link remains debated. We investigated canine emotional states underlying tail wagging by systematically analysing differences in tail movements in a computer-controlled task encompassing two non-social Conditions - Rewarded (positive) and Unrewarded (negative), and two Epochs (pre-response and outcome anticipation). Using pose-tracking we found that 11 out of 23 dogs did not wag their tails in at least 75% of trials, suggesting that some dogs may inherently wag less or that tail wagging is primarily a social signal. Our results showed that dogs were more likely to wag during positive anticipation; whereas in the negative condition, despite tail amplitude being more prominent, increased speeds reflected arousal rather than valence. Further work should assess tail kinematics in social contexts to test and extend these findings.

The ability of animals to innovate - solve novel problems - can shape their ecology and evolution. Here we investigate how individual traits and environmental complexity relate to successful solving of a novel problem. We presented foraging bumble bees (Bombus impatiens) with artificial flowers of not-previously-encountered shapes and recorded the bees latency to access nectar. We measured individual foraging traits across multiple trips with simple flowers that did not require innovation, and bees were foraging either in a simple or complex environment (cluttered flight arena). Bees in complex environments took longer to find and were less likely to land on novel flowers, indicating that environmental complexity may take up cognitive resources and make search more difficult. However, we did not find an effect of environmental treatment on the ability or time to access reward in novel flowers once bees had landed on them. In contrast, behavioral traits significantly predicted how quickly bees solved novel flowers. In particular, overall foraging tempo as well as routine formation, i.e. how much bees followed a fixed route on known flowers, predicted innovation - faster bees innovated faster, and bees with more repetitive foraging sequences were slower to solve the novel tasks. Overall, while the degree of evolutionary novelty in tasks or solutions is always hard to evaluate, our findings demonstrate that environment and individual traits may affect innovation in different ways. Individuals in simple environments may be more likely to detect, and individuals that are generally faster and have a lower tendency to develop fixed routines may be more likely to solve, novel tasks.

Collective decision making is a fundamental aspect of group behavior in both animals and humans, and often involves reaching a consensus on the best of n options, using empirical evidence. Although many parallels have been drawn between human and animal collective decisions, collective human behavior is rarely studied in the type of embodied scenarios that animals are often faced with. In this study, we placed human groups in a virtual setup similar to nest site selection in social animals, in which they explored a shared environment and reached a consensus based on their observations of empirical features. In groups of up to 10, participants had to reach consensus on the empirically largest of four candidate sites without verbal communication, instead using movement-based interactions in a custom-developed 3D virtual environment for online multi-participant experiments. The results showed that the speed and accuracy of consensus was importantly modulated by perceptual difficulty and information availability, but that no speed-accuracy trade-off was present. Participants attempted to reach consensus on the empirically largest site by flexibly adapting their use of social information to perceptual difficulty, their spatial position, and the time already spent supporting some option. When a minority of informed individuals were present, these individuals exercised greater independence and influenced the group to faster and more accurate consensus. These results extend previous findings on social decision making strategies in humans to nonverbal scenarios akin to those of social insects.

Social living is widely considered a key driver of cognitive evolution, yet individuals within a species and even within the same group can differ substantially in their sociability (i.e., an individuals propensity to form and maintain social bonds), which can ultimately shape the social environment they experience by influencing how they interact with, respond to, and engage in it. How such individual variation in this personality trait affects social cognition, however, remains poorly understood. To address this question, we used two selectively bred lines of Japanese quail (Coturnix japonica) that consistently differ in social motivation, a key component of sociability, which we used as a proxy for this broader trait. In these lines, S+ individuals show high social motivation, whereas S- individuals show low social motivation. We compared their sociocognitive performance across three tasks: a gaze following task, a social buffering task, and a social discrimination learning task. Our findings revealed that Japanese quail reliably followed the gaze of conspecifics, providing the first evidence of this ability in this species. However, there was no difference between lines, suggesting that basic gaze following into the distance is independent of social motivation. In contrast, line differences emerged in the other tasks. S+ quail were more sensitive and less adaptable in response to environmental changes, particularly under social isolation, although the presence of a conspecific strongly buffered these effects. S- quail, on the other hand, outperformed S+ individuals in the social discrimination learning task, rapidly exploiting available social cues to guide foraging decisions. Overall, this study demonstrates that social motivation modulates sociocognitive performance in a context-dependent manner. Rather than conferring a general cognitive advantage, divergent selection on social motivation leads to different strategies of social information use across tasks and contexts, highlighting the complex interplay between personality, social environment, and cognition.

When novel nutrient-rich food sources become available to species sharing the same natural habitat, interspecies competition may arise, yielding insights into the ecological and social dynamics of the observed species. Here, we investigated food consumption patterns, and consequent social interactions, by two sympatric species of mice in response to a novel nutrient-rich food source. By deploying, in the mices natural habitat, baited video-monitored chambers, we collected, over a 5-month period, 1805 observations of food visiting by Apodemus agrarius and Apodemus flavicollis. We also documented interspecific encounters, with 86.7% of the cases showing agonism. In these interspecies agonistic encounters, A. flavicollis was always the initiator of agonism, attacking within 2 sec in 92.3% of the cases, and being dominant over A. agrarius in 84.6%. Analysis of food visiting behavior revealed that, initially, both species preferred nocturnality. However, after the interspecies fights, A. agrarius switched its temporal preference to diurnality, leading to temporal niche segregation between the two species and a significant reduction of interspecies encounters. Moreover, A. agrarius demonstrated hour-specific avoidance of A. flavicollis, visiting significantly less in hours with A. flavicollis compared to hours without. Through temporal niche switching, A. agrarius managed to access the food source safely, without fights. In contrast, A. flavicollis remained consistently nocturnal across the entire study. Notably, our study presents the first 24h foraging actogram for free-living rodents. Moreover, while rodent interspecific competition is a well-known phenomenon, most of what we know about it comes from indirect observations. Direct observations of rodent interspecific interactions in nature are rare. Our work is the first direct (video-monitored) observation of temporal switch-inducing interspecies interactions in nature. As free-living rodents are currently considered a major model system for the study of interspecific competition, these results may offer precious insights for a better understanding of social dynamics, especially in asymmetric relationships. Furthermore, our findings highlight the significance of considering temporal dynamics in studies of interspecific interactions.