Foraging synchrony drives resilience in human-dolphin mutualism.

Interactions between humans and nature have profound consequences, which rarely are mutually beneficial. Further, behavioral and environmental changes can turn human-wildlife cooperative interactions into conflicts, threatening their continued existence. By tracking fine-scale behavioral interactions between artisanal fishers and wild dolphins targeting migratory mullets, we reveal that foraging synchrony is key to benefiting both predators. Dolphins herd mullet schools toward the coast, increasing prey availability within the reach of the net-casting fishers, who gain higher foraging success-but only when matching the casting behavior with the dolphins' foraging cues. In turn, when dolphins approach the fishers' nets closely and cue fishers in, they dive for longer and modify their active foraging echolocation to match the time it takes for nets to sink and close over mullets-but only when fishers respond to their foraging cues appropriately. Using long-term demographic surveys, we show that cooperative foraging generates socioeconomic benefits for net-casting fishers and ca. 13% survival benefits for cooperative dolphins by minimizing spatial overlap with bycatch-prone fisheries. However, recent declines in mullet availability are threatening these short- and long-term benefits by reducing the foraging success of net-casting fishers and increasing the exposure of dolphins to bycatch in the alternative fisheries. Using a numerical model parametrized with our empirical data, we predict that environmental and behavioral changes are pushing this traditional human-dolphin cooperation toward extinction. We propose two possible conservation actions targeting fishers' behavior that could prevent the erosion of this century-old fishery, thereby safeguarding one of the last remaining cases of human-wildlife cooperation.

Social foraging and the associated benefits of group-living in Cliff Swallows decrease over 40 years.

Animals that feed socially can sometimes better locate prey, often by transferring information about food that is patchy, dense, and temporally and spatially unpredictable. Information transfer is a potential benefit of living in breeding colonies where unsuccessful foragers can more readily locate successful ones and thereby improve feeding efficiency. Most studies on social foraging have been short-term, and how long-term environmental change affects both foraging strategies and the associated benefits of coloniality is generally unknown. In the colonial Cliff Swallow (Petrochelidon pyrrhonota), we examined how social foraging, information transfer, and feeding ecology changed over a 40-year period in western Nebraska. Relative to the 1980's, Cliff Swallows in 2016-2022 were more likely to forage solitarily or in smaller groups, spent less time foraging, were more successful as solitaries, fed in more variable locations, and engaged less in information transfer at the colony site. The total mass of insects brought back to nestlings per parental visit declined over the study. The diversity of insect families captured increased over time, and some insect taxa dropped out of the diet, although the three most common insect families remained the same among the decades. Nestling Cliff Swallow body mass at 10 days of age and the number of nestlings surviving per nest declined more sharply with colony size in 2015-2022 than in 1984-1991 at sites where the confounding effects of ectoparasites were removed. Adult body mass during provisioning of nestlings was lower in more recent years, but the change did not vary with colony size. The reason(s) for the reduction in social foraging and information transfer over time are unclear, but the consequence is that colonial nesting may no longer offer the same fitness advantages for Cliff Swallows as in the 1980's. The results illustrate flexibility of foraging behavior and dynamic shifts in the potential selective pressures for group-living.

Social dynamics and individual hunting tactics of white sharks revealed by biologging.

Social foraging, where animals forage in groups, takes many forms but is less studied in marine predators as measuring social associations in the wild is challenging. We used biologging (activity, cameras and telemetry receivers) sensors to measure social associations and simultaneous behaviour, in white sharks (Carcharodon carcharias) off Guadalupe Island, Mexico. Animal-borne telemetry receivers revealed that sharks varied in the number of associations they formed and occurred most often when sharks were swimming in straight paths or when they were turning frequently. While many associations were likely random, there was evidence of some stronger associations. Sharks varied in the depths they used and their activity, with some individuals more active in shallow water while others were more active 200-300 m deep. We propose that white sharks associate with other individuals so they can inadvertently share information on the location or remains of large prey. However, there may be a wide range of individual variability in both behaviour and sociality. Biologging now enables social associations of animals to be measured, concurrent with measures of their behaviour, so that social foraging of large marine predators can be quantified in the wild.

Cognition and covariance in the producer-scrounger game.

The producer-scrounger game is a key element of foraging ecology in many systems. Producing and scrounging typically covary negatively, but partitioning this covariance into contributions of individual plasticity and consistent between individual differences is key to understanding population-level consequences of foraging strategies. Furthermore, little is known about the role cognition plays in the producer-scrounger game. We investigated the role of cognition in these alternative foraging tactics in wild mixed-species flocks of great tits and blue tits, using a production learning task in which we measured individuals' speed of learning to visit the single feeder in an array that would provide them with a food reward. We also quantified the proportion of individuals' feeds that were scrounges ('proportion scrounged'); scrounging was possible if individuals visited immediately after a previous rewarded visitor. Three learning experiments-initial and two reversal learning-enabled us to estimate the repeatability and covariance of each foraging behaviour. First, we examined whether individuals learned to improve their scrounging success (i.e. whether they obtained food by scrounging when there was an opportunity to do so). Second, we quantified the repeatability of proportion scrounged, and asked whether proportion scrounged affected production learning speed among individuals. Third, we used multivariate analyses to partition within- and among-individual components of covariance between proportion scrounged and production learning speed. Individuals improved their scrounging success over time. Birds with a greater proportion scrounged took longer to learn their own rewarding feeder. Although multivariate analyses showed that covariance between proportion scrounged and learning speed was driven primarily by within-individual variation, that is, by behavioural plasticity, among-individual differences also played a role for blue tits. This is the first demonstration of a cognitive trait influencing producing and scrounging in the same wild system, highlighting the importance of cognition in the use of alternative resource acquisition tactics. The results of our covariance analyses suggest the potential for genetic differences in allocation to alternative foraging tactics, which are likely species- and system-dependent. They also point to the need to control for different foraging tactics when studying individual cognition in the wild.

Limited sexual segregation in a dimorphic avian scavenger, the Andean condor.

Sexual segregation is widely reported among sexually dimorphic species and generally attributed to intraspecific competition. Prey diversity and human activities can reinforce niche segregation by increasing resource heterogeneity. Here, we explored trophic and spatial sexual segregation in the only avian scavenger that exhibits pronounced sexual size dimorphism (up to 50% difference in body mass) and a highly despotic social system, the Andean condor (Vultur gryphus). We predicted that larger and dominant males would exclude smaller and subordinate females from high-quality resources, leading to sexual segregation particularly in human-dominated landscapes showing increased prey diversity. We compared resource use between females and males across six sites in Argentina featuring a range of prey diversity via stable isotopes analysis of molted feathers (n = 141 individuals). We then focused on two sites featuring contrasting levels of prey diversity and quantified assimilated diet via stable isotopes and space use via GPS monitoring (n = 23 and 12 tagged individuals). We found no clear differences in isotopic niche space, individual variation in isotopic signature, or assimilated diet between females and males. However, there were differences in foraging locations between sexes, with females apparently using areas of fewer food resources more frequently than males. Local conditions defined the dynamics of fine-scale sexual differences in foraging sites; yet, unpredictable and ephemeral carrion resources likely prevent segregation by sexes at the landscape scale. Our study highlights complex dynamics of sexual segregation in vultures and the relevancy of analyses under multiple spatial-temporal scales to explore segregation in social species.

Consistent sociality but flexible social associations across temporal and spatial foraging contexts in a colonial breeder.

When the consequences of sociality differ depending on the state of individual animals and the experienced environment, individuals may benefit from altering their social behaviours in a context-dependent manner. Thus, to fully address the hypotheses about the role of social associations it is imperative to consider the multidimensional nature of sociality by explicitly examining social associations across multiple scales and contexts. We simultaneously recorded > 8000 associations from 85% of breeding individuals from a colony of Australasian gannets (Morus serrator) over a 2-week period, and examined gregariousness across four foraging states using multilayer social network analysis. We found that social associations varied in a context-dependent manner, highlighting that social associations are most prevalent during foraging (local enhancement) and in regions expected to provide clustered resources. We also provide evidence of individual consistency in gregariousness, but flexibility in social associates, demonstrating that individuals can adjust their social behaviours to match experienced conditions.

Competitive advantage of rare behaviours induces adaptive diversity rather than social conformity in skill learning.

Recent studies have emphasized the role of social learning and cultural transmission in promoting conformity and uniformity in animal groups, but little attention has been given to the role of negative frequency-dependent learning in impeding conformity and promoting diversity instead. Here, we show experimentally that under competitive conditions that are common in nature, social foragers (although capable of social learning) are likely to develop diversity in foraging specialization rather than uniformity. Naive house sparrows that were introduced into groups of foraging specialists did not conform to the behaviour of the specialists, but rather learned to use the alternative food-related cues, thus forming groups of complementary specialists. We further show that individuals in such groups may forage more effectively in diverse environments. Our results suggest that when the benefit from socially acquired skills diminishes through competition in a negative frequency-dependent manner, animal societies will become behaviourally diverse rather than uniform.

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

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

The two oxpecker species reveal the role of movement rates and foraging intensity in species coexistence.

The two Buphagus oxpecker species are specialized passerines that forage for ticks and other food particles on the body of ungulates in the African savannahs. One of their intriguing features is their ability to coexist despite sharing the same, specialized diet. Using co-occurrence data (photographs of giraffes with oxpeckers on them) and approximate Bayesian computing, we demonstrate that yellow-billed oxpeckers changed host faster than red-billed oxpeckers and appeared to displace red-billed oxpeckers from preferred giraffe body parts. Conversely, red-billed oxpeckers exhibited a fuller use of each host and displaced yellow-billed oxpeckers from distal giraffe body parts. These findings highlight that the partition of giraffe hosts in two separate niches was only part of the coexistence story in this species pair. More precisely, the oxpeckers shared the resource by exploiting it at different rates. They engaged in different trade-offs between giving-up density, patch discovery rate and competitor displacement ability. They illustrate the importance of the time frame of interactions.

Evidence of sociality in the timing and location of foraging in a colonial seabird.

Social foraging behaviours, which range from cooperative hunting to local enhancement, can result in increased prey capture and access to information, which may significantly reduce time and energy costs of acquiring prey. In colonial species, it has been proposed that the colony itself may act as a site of social information transfer and group formation. However, conclusive evidence from empirical studies is lacking. In particular, most studies in colonial species have generally focussed on behaviours either at the colony or at foraging sites in isolation, and have failed to directly connect social associations at the colony to social foraging. In this study, we simultaneously tracked 85% of a population of Australasian gannets (Morus serrator) over multiple foraging trips, to study social associations at the colony and test whether these associations influence the location of foraging sites. We found that gannets positively associate with conspecifics while departing from the colony and that co-departing gannets have more similar initial foraging patches than individuals that did not associate at the colony. These results provide strong evidence for the theory that the colony may provide a source of information that influences foraging location.

Foraging and inter-individual distances of bearded capuchin monkeys.

Extractive foraging is a skill young capuchin monkeys learn over time. A key unknown is whether unskilled individuals occupy spatial positions that increase their opportunities to learn. We observed the spatial positions of individuals in a group of capuchin monkeys in Northeastern Brazil. To improve our understanding of the relationship between learning by young capuchin monkeys and inter-individual distance, we investigated the associations between the proximity of individuals and their age, activity, and proficiency at extractive foraging. To do this, we used one form of extractive foraging, opening palm nuts, as an index of proficiency at all types of extractive foraging. Our results indicate that, in the subset of the data where dyads consisted of one proficient individual and a partner with any level of proficiency, the distance between individuals was predicted by their foraging activity (i.e., extractive foraging, other foraging, or not foraging). In those dyads, the proficiency of the partner did not significantly improve prediction of inter-individual distances, indicating that spatial proximity of proficient individuals to others does not function primarily to increase opportunities for unskilled individuals to observe extractive foraging. Dyads in which both individuals were engaged in similar foraging activities (e.g., both "extractive foraging") exhibited the shortest inter-individual distances. Proximity between individuals engaged in similar foraging activities may result from the spatial distribution of resources or from social learning mechanisms, such as local or stimulus enhancement.

Selective contribution of the telencephalic arcopallium to the social facilitation of foraging efforts in the domestic chick.

To investigate the neural basis of socio-economic behaviors in birds, we examined the effects of bilateral electrolytic lesions of arcopallium (Arco, the major descending pallial area of the avian telencephalon) and the surrounding nuclei in domestic chicks. We tested foraging effort (running distance) in an I-shaped maze with two food patches that delivered food in a biased manner according to a variable interval schedule. Normally, chicks run back and forth between the patches, and the patch use time matches the respective food delivery rate. In the paired phase, even without actual interference of food, chicks showed social facilitation of running effort compared with the single phase. Chicks with lesions in the Arco and lateral Arco showed significant reductions in social facilitation. The lesion effects of the lateral Arco were particularly selective, as it was not accompanied by changes in running distance in the single phase. Lesions of the nidopallium and nucleus taeniae of the amygdala produced no changes in foraging behavior. On the other hand, the Arco lesion did not impair social facilitation of operant peck latency. In accordance with this, anterograde tracing revealed characteristic projections from the lateral Arco to the extended amygdala, hippocampus, and septum, as well as wide areas of limbic nuclei in the hypothalamus and medial areas of the striatum including the nucleus accumbens. Pathways from the lateral Arco could enable chicks to overcome the extra effort investment of social foraging, suggesting functional and anatomical analogies to the anterior cingulate cortex and basolateral amygdala in mammals.

Wild birds respond to flockmate loss by increasing their social network associations to others.

Understanding the consequences of losing individuals from wild populations is a current and pressing issue, yet how such loss influences the social behaviour of the remaining animals is largely unexplored. Through combining the automated tracking of winter flocks of over 500 wild great tits (Parus major) with removal experiments, we assessed how individuals' social network positions responded to the loss of their social associates. We found that the extent of flockmate loss that individuals experienced correlated positively with subsequent increases in the number of their social associations, the average strength of their bonds and their overall connectedness within the social network (defined as summed edge weights). Increased social connectivity was not driven by general disturbance or changes in foraging behaviour, but by modifications to fine-scale social network connections in response to losing their associates. Therefore, the reduction in social connectedness expected by individual loss may be mitigated by increases in social associations between remaining individuals. Given that these findings demonstrate rapid adjustment of social network associations in response to the loss of previous social ties, future research should examine the generality of the compensatory adjustment of social relations in ways that maintain the structure of social organization.

Stable producer-scrounger dynamics in wild birds: sociability and learning speed covary with scrounging behaviour.

There has been extensive game-theoretic modelling of conditions leading to equilibria of producer-scrounger dichotomies in groups. However there is a surprising paucity of experimental evidence in wild populations. Here, we examine producer-scrounger games in five subpopulations of birds feeding at a socially learnt foraging task. Over four weeks, a bimodal distribution of producers and scroungers emerged in all areas, with pronounced and consistent individual tactic specialization persisting over 3 years. Tactics were unrelated to exploratory personality, but correlated with latency to contact and learn the foraging task, with the late arrivers and slower learners more likely to adopt the scrounging role. Additionally, the social environment was also important: at the broad scale, larger subpopulations with a higher social density contained proportionally more scroungers, while within subpopulations scroungers tended to be central in the social network and be observed in larger foraging flocks. This study thus provides a rare example of a stable, dimorphic distribution of producer-scrounger tactics in a wild population. It further gives support across multiple scales for a major prediction of social foraging theory; that the frequency of scroungers increases with group size.

Producers and scroungers: feeding-type composition changes with group size in a socially foraging spider.

In groups of socially foraging animals, feeding behaviour may change with group size in response to varying cost-benefit trade-offs. Numerous studies have described group-size effects on group-average feeding behaviour, particularly emphasizing an increase in scrounging incidence for larger groups, where individuals (scroungers) feed from the food sources others (producers) discovered. However, individual variation in feeding behaviour remains unconsidered in the vast majority of these studies even though theoretical models predict individuals to specialize in feeding tactic and anticipate higher scrounger-type frequencies in larger groups. We combined group-level and individual-level analyses of group-size effects on social foraging in the subsocial spider Australomisidia ergandros Lending novel experimental support to model predictions, we found that individuals specialize in feeding tactic and that higher scrounging and lower producing incidence in larger groups were mediated through shifts in the ratio of feeding types. Further, feeding-type specialization was not explained by innate individual differences in hunting ability as all feeding types were equally efficient in prey capture when foraging alone. Context adaptivity of feeding behaviour might allow this subsocial species to succeed under varying socioecological conditions.

The local enhancement conundrum: in search of the adaptive value of a social learning mechanism.

Social learning mechanisms are widely thought to vary in their degree of complexity as well as in their prevalence in the natural world. While learning the properties of a stimulus that generalize to similar stimuli at other locations (stimulus enhancement) prima facie appears more useful to an animal than learning about a specific stimulus at a specific location (local enhancement), empirical evidence suggests that the latter is much more widespread in nature. Simulating populations engaged in a producer-scrounger game, we sought to deploy mathematical models to identify the adaptive benefits of reliance on local enhancement and/or stimulus enhancement, and the alternative conditions favoring their evolution. Surprisingly, we found that while stimulus enhancement readily evolves, local enhancement is advantageous only under highly restricted conditions: when generalization of information was made unreliable or when error in social learning was high. Our results generate a conundrum over how seemingly conflicting empirical and theoretical findings can be reconciled. Perhaps the prevalence of local enhancement in nature is due to stimulus enhancement costs independent of the learning task itself (e.g. predation risk), perhaps natural habitats are often characterized by unreliable yet highly rewarding payoffs, or perhaps local enhancement occurs less frequently, and stimulus enhancement more frequently, than widely believed.

Why are ravens smart? Exploring the social intelligence hypothesis.

Ravens and other corvids are renowned for their 'intelligence'. For long, this reputation has been based primarily on anecdotes but in the last decades experimental evidence for impressive cognitive skills has accumulated within and across species. While we begin to understand the building blocks of corvid cognition, the question remains why these birds have evolved such skills. Focusing on Northern Ravens Corvus corax, I here try to tackle this question by relating current hypotheses on brain evolution to recent empirical data on challenges faced in the birds' daily life. Results show that foraging ravens meet several assumptions for applying social intelligence: (1) they meet repeatedly at foraging sites, albeit individuals have different site preferences and vary in grouping dynamics; (1) foraging groups are structured by dominance rank hierarchies and social bonds; (3) individual ravens memorize former group members and their relationship valence over years, deduce third-party relationships and use their social knowledge in daily life by supporting others in conflicts and intervening in others' affiliations. Hence, ravens' socio-cognitive skills may be strongly shaped by the 'complex' social environment experienced as non-breeders.

Exploring the size of Andean condor foraging groups along an altitudinal and latitudinal gradient in the Tropical Andes: Ecological and conservation implications.

Patterns of variation in the size of vulture foraging groups, and their ecological causes and consequences, remain little explored despite strong links with the carrion recycling service that this key functional group provides. We documented the group size-frequency pattern of Andean condors Vultur gryphus gathered to feed on 42 equine carcasses experimentally placed in Bolivia, between 2012 and 2019, along an elevation range of 1,300-4,500 m asl. Based on the location (altitude and latitude) of the foraging groups, we examined the relationship between their size and species' population parameters (size and trend), habitat conditions, and livestock carcass availability and predictability. Condors utilized a high frequency (93%) of carcasses forming groups that ranged from 1 to 80 individuals (mean = 25, median = 18) and shaped a "lazy-J curve" typical pattern of size-frequency distribution whereby few groups (5, 12%) were large (> 55 individuals) and most (21, 50%) were relatively small (<19 individuals). Group size related to altitude in that most larger groups formed at lower sites (below c. 3,000 m asl), likely following an altitudinal gradient whereby larger groups are more likely to form around larger carcasses (i.e., cattle), which are more likely to occur at lower elevations. Regardless of population size, group size could be an adaptive response of condors via local enhancement for improving individual scavenging efficiency. Many information gaps on this topic still exist, thus we provide a set of questions to address them, especially amidst the unrestricted impacts of human activities that condition vulture survival globally.

Social foragers adopt a riskier foraging mode in the centre of their groups.

Foraging in groups provides many benefits that are not necessarily experienced the same way by all individuals. I explore the possibility that foraging mode, the way individuals exploit resources, varies as a function of spatial position in the group, reflecting commonly occurring spatial differences in predation risk. I show that semipalmated sandpipers (Calidris pusilla), a social foraging avian species, tended to adopt a riskier foraging mode in the central, more protected areas of their groups. Central birds effectively used the more peripheral group members as sentinels, allowing them to exploit a wider range of resources within the same group at the same time. This finding provides a novel benefit of living in groups, which may have a broad relevance given that social foraging species often exploit a large array of resources.

Increased initial task difficulty drives social foragers to develop sub-optimal conformity instead of adaptive diversity.

The extent to which animal societies exhibit social conformity as opposed to behavioural diversity is commonly attributed to adaptive learning strategies. Less attention is given to the possibility that the relative difficulty of learning a task socially as opposed to individually can be critical for social learning dynamics. Here we show that by raising initial task difficulty, house sparrows previously shown to exhibit adaptive social diversity become predominantly conformists. The task we used required opening feeding well covers (easier to learn socially) and to choose the covers with the rewarding cues (easy to learn individually). We replicated a previous study where sparrows exhibited adaptive diversity, but did not pre-train the naive sparrows to open covers, making the task initially more difficult. In sharp contrast to the previous study results, most sparrows continued to conform to the demonstrated cue even after experiencing greater success with the alternative rewarding cue for which competition was less intense. Thus, our study shows that a task's cognitive demands, such as the initial dependency on social demonstration, can change the entire learning dynamics, causing social animals to exhibit sub-optimal social conformity rather than adaptive diversity under otherwise identical conditions.