Homing pigeons (Columba livia) have been the central model of avian navigation research for many decades, but only more recently has research extended into understanding their mechanisms of orientation in the familiar area. The discovery (facilitated by GPS tracking) that pigeons gradually acquire with experience individually idiosyncratic routes home to which they remain faithful on repeated releases, even if displaced off-route, has helped uncover the fundamental role of familiar visual landmarks in the avian familiar area map. We evaluate the robustness and generality of the route-following phenomenon by examining extant studies in depth, including the single published counter-example, providing a detailed comparison of route efficiencies, flight corridor widths and fidelity. We combine this analysis with a review of inferences that can be drawn from other experimental approaches to understanding the nature of familiar area orientation in pigeons, including experiments on landmark recognition, and response to clock-shift, to build the first detailed picture of how bird orientation develops with experience of the familiar area. We articulate alternative hypotheses for how guidance might be controlled during route following, concluding that although much remains unknown, the details of route following strongly support a pilotage interpretation. Predictable patterns of efficiency increase, but limited to the local route, typical corridor widths of 100-200 m, high-fidelity pinch-points on route, attraction to landscape edges, and a robustness to clock-shift procedures, all demonstrate that birds can associatively acquire a map of their familiar area guided (at least partially) by direct visual control from memorised local landscape features.
The aerial lifestyle of central-place foraging birds allows wide-ranging movements, raising fundamental questions about their remarkable navigation and memory systems. For example, we know that pigeons (Columba livia), long-standing models for avian navigation, rely on individually distinct routes when homing from familiar sites. But it remains unknown how they cope with the task of learning several routes in parallel. Here, we examined how learning multiple routes influences homing in pigeons. We subjected groups of pigeons to different training protocols, defined by the sequence in which they were repeatedly released from three different sites, either sequentially, in rotation or randomly. We observed that pigeons from all groups successfully developed and applied memories of the different release sites (RSs), irrespective of the training protocol, and that learning several routes in parallel did not impair their capacity to quickly improve their homing efficiency over multiple releases. Our data also indicated that they coped with increasing RS uncertainty by adjusting both their initial behaviour upon release and subsequent homing efficiency. The results of our study broaden our understanding of avian route following and open new possibilities for studying learning and memory in free-flying animals.
Observations of the flight paths of pigeons navigating from familiar locations have shown that these birds are able to learn and subsequently follow habitual routes home. It has been suggested that navigation along these routes is based on the recognition of memorized visual landmarks. Previous research has identified the effect of landmarks on flight path structure, and thus the locations of potentially salient sites. Pigeons have also been observed to be particularly attracted to strong linear features in the landscape, such as roads and rivers. However, a more general understanding of the specific characteristics of the landscape that facilitate route learning has remained out of reach. In this study, we identify landscape complexity as a key predictor of the fidelity to the habitual route, and thus conclude that pigeons form route memories most strongly in regions where the landscape complexity is neither too great nor too low. Our results imply that pigeons process their visual environment on a characteristic spatial scale while navigating and can explain the different degrees of success in reproducing route learning in different geographical locations.
Hierarchical organization is widespread in the societies of humans and other animals, both in social structure and in decision-making contexts. In the case of collective motion, the majority of case studies report that dominant individuals lead group movements, in agreement with the common conflation of the terms "dominance" and "leadership." From a theoretical perspective, if social relationships influence interactions during collective motion, then social structure could also affect leadership in large, swarm-like groups, such as fish shoals and bird flocks. Here we use computer-vision-based methods and miniature GPS tracking to study, respectively, social dominance and in-flight leader-follower relations in pigeons. In both types of behavior we find hierarchically structured networks of directed interactions. However, instead of being conflated, dominance and leadership hierarchies are completely independent of each other. Although dominance is an important aspect of variation among pigeons, correlated with aggression and access to food, our results imply that the stable leadership hierarchies in the air must be based on a different set of individual competences. In addition to confirming the existence of independent and context-specific hierarchies in pigeons, we succeed in setting out a robust, scalable method for the automated analysis of dominance relationships, and thus of social structure, applicable to many species. Our results, as well as our methods, will help to incorporate the broader context of animal social organization into the study of collective behavior.
In many species, group members obtain benefits from moving collectively, such as enhanced foraging efficiency or increased predator detection. In situations where the groups decision involves integrating individual preferences, group cohesion can lead to more accurate outcomes than solitary decisions. In homing pigeons, a classic model in avian orientation studies, individuals learn habitual routes home, but whether and how co-navigating birds acquire and share route-based information is unknown. Using miniature GPS loggers, we examined these questions by first training pairs (the smallest possible flocks) of pigeons together, and then releasing them with other pairs that had received separate pair-training. Our results show that, much like solitary individuals, pairs of birds are able to establish idiosyncratic routes that they recapitulate together faithfully. Also, when homing with other pairs they exhibit a transition from a compromise- to a leadership-like mechanism of conflict resolution as a function of the degree of disagreement (distance separating the two preferred routes) between the two pairs, although pairs tolerate a greater range of disagreements prior to the transition than do single birds. We conclude that through shared experiences during past decision-making, pairs of individuals can become units so closely coordinated that their behaviour resembles that of single birds. This has implications for the behaviour of larger groups, within which certain individuals have closer social affiliations or share a history of previous associations.
Tool use is a vital component of the human behavioural repertoire. The benefits of tool use have often been assumed to be self-evident: by extending control over our environment, we have increased energetic returns and buffered ourselves from potentially harmful influences. In recent decades, however, the study of tool use in both humans and non-human animals has expanded the way we think about the role of tools in the natural world. This Theme Issue is aimed at bringing together this developing body of knowledge, gathered across multiple species and from multiple research perspectives, to chart the wider evolutionary context of this phylogenetically rare behaviour.
Travelling in groups gives animals opportunities to share route information by following cues from each others movement. The outcome of group navigation will depend on how individuals respond to each other within a flock, school, swarm or herd. Despite the abundance of modelling studies, only recently have researchers developed techniques to determine the interaction rules among real animals. Here, we use high-resolution GPS (global positioning system) tracking to study these interactions in pairs of pigeons flying home from a familiar site. Momentary changes in velocity indicate alignment with the neighbours direction, as well as attraction or avoidance depending on distance. Responses were stronger when the neighbour was in front. From the flocking behaviour, we develop a model to predict features of group navigation. Specifically, we show that the interactions between pigeons stabilize a side-by-side configuration, promoting bidirectional information transfer and reducing the risk of separation. However, if one bird gets in front it will lead directional choices. Our model further predicts, and observations confirm, that a faster bird (as measured from solo flights) will fly slightly in front and thus dominate the choice of homing route. Our results explain how group decisions emerge from individual differences in homing flight behaviour.
Burmese long-tailed macaques (Macaca fascicularis aurea) are one of a limited number of wild animal species to use stone tools, with their tool use focused on pounding shelled marine invertebrates foraged from intertidal habitats. These monkeys exhibit two main styles of tool use: axe hammering of oysters, and pound hammering of unattached encased foods. In this study, we examined macroscopic use-wear patterns on a sample of 60 wild macaque stone tools from Piak Nam Yai Island, Thailand, that had been collected following behavioural observation, in order to (i) quantify the wear patterns in terms of the types and distribution of use-damage on the stones, and (ii) develop a Use-Action Index (UAI) to differentiate axe hammers from pound hammers by wear patterns alone. We used the intensity of crushing damage on differing surface zones of the stones, as well as stone weight, to produce a UAI that had 92% concordance when compared to how the stones had been used by macaques, as observed independently prior to collection. Our study is the first to demonstrate that quantitative archaeological use-wear techniques can accurately reconstruct the behavioural histories of non-human primate stone tools.
The sun has long been thought to guide bird navigation as the second step in a two-stage process, in which determining position using a map is followed by course setting using a compass, both over unfamiliar and familiar terrain. The animals endogenous clock time-compensates the solar compass for the suns apparent movement throughout the day, and this allows predictable deflections in orientation to test for the compass influence using clock-shift manipulations. To examine the influence of the solar compass during a highly familiar navigational task, 24 clock-shifted homing pigeons were precision-tracked from a release site close to and in sight of their final goal, the colony loft. The resulting trajectories displayed significant partial deflection from the loft direction as predicted by either fast or slow clock-shift treatments. The partial deflection was also found to be stable along the entire trajectory indicating regular updating of orientation via input from the solar compass throughout the final approach flight to the loft. Our results demonstrate that time-compensated solar cues are deeply embedded in the way birds orient during homing flight, are accessed throughout the journey and on a remarkably fine-grained scale, and may be combined effectively simultaneously with direct guidance from familiar landmarks, even when birds are flying towards a directly visible goal.
Social animals have much to gain from observing and responding appropriately to the actions of their conspecific group members. This can in turn lead to the learning of novel behavior patterns (social learning) or to foraging, ranging, or social behavioral choices copied from fellow group members, which do not necessarily result in long-term learning, but at the time represent adaptive responses to environmental cues (public information use). In the current study, we developed a novel system for the study of public information use under fully automated conditions. We modified a classic single-subject laboratory paradigm--matching-to-sample (MTS)--and examined chimpanzees ability to interpret and utilize cues provided by the behavior of a conspecific to solve the task. In Experiment 1, two subjects took turns on an identity MTS task, with one subject (the model) performing the first half of the trial and the other subject (the observer) completing the trial using the models actions as discriminative cues. Both subjects performed above chance from the first session onwards. In Experiment 2, the subjects were tested on a symbolic version of the same MTS task, with one subject showing spontaneous transfer. Our study establishes a novel method for examining public information use within a highly controlled and automated setting.
How social-living animals make collective decisions is currently the subject of intense scientific interest, with increasing focus on the role of individual variation within the group. Previously, we demonstrated that during paired flight in homing pigeons, a fully transitive leadership hierarchy emerges as birds are forced to choose between their own and their partners habitual routes. This stable hierarchy suggests a role for individual differences mediating leadership decisions within homing pigeon pairs. What these differences are, however, has remained elusive. Using novel quantitative techniques to analyse habitual route structure, we show here that leadership can be predicted from prior route-following fidelity. Birds that are more faithful to their own route when homing alone are more likely to emerge as leaders when homing socially. We discuss how this fidelity may relate to the leadership phenomenon, and propose that leadership may emerge from the interplay between individual route confidence and the dynamics of paired flight.
Experiments with robins wearing frosted goggles have revealed a tantalising relationship between object vision and magnetoreception, shedding further light on the close interconnection between the two senses in migratory birds.
Pigeons home along idiosyncratic habitual routes from familiar locations. It has been suggested that memorized visual landmarks underpin this route learning. However, the inability to experimentally alter the landscape on large scales has hindered the discovery of the particular features to which birds attend. Here, we present a method for objectively classifying the most informative regions of animal paths. We apply this method to flight trajectories from homing pigeons to identify probable locations of salient visual landmarks. We construct and apply a Gaussian process model of flight trajectory generation for pigeons trained to home from specific release sites. The model shows increasing predictive power as the birds become familiar with the sites, mirroring the animals learning process. We subsequently find that the most informative elements of the flight trajectories coincide with landscape features that have previously been suggested as important components of the homing task.
Field experiments can provide compelling demonstrations of social learning in wild populations. Social learning has been experimentally demonstrated in at least 23 field experiments, in 20 species, covering a range of contexts, such as foraging preferences and techniques, habitat choice, and predator avoidance. We review experimental approaches taken in the field and with wild animals brought into captivity and note how these approaches can be extended. Relocating individuals, introducing trained individual demonstrators or novel behaviors into a population, or providing demonstrator-manipulated artifacts can establish whether and how a particular act can be socially transmitted in the wild and can help elucidate the benefits of social learning. The type, strength, and consistency of presented social information can be varied, and the provision of conditions favoring the performance of an act can both establish individual discovery rates and help determine whether social information is needed for acquisition. By blocking particular avenues of social transmission or removing key individuals, routes of transmission in wild populations can be investigated. Manipulation of conditions proposed to favor social learning can test mathematical models of the evolution of social learning. We illustrate how field experiments are a viable, vital, and informative approach to the study of social learning.
The forests surrounding Bossou, Guinea, are home to a small, semi-isolated chimpanzee community studied for over three decades . In 1992, Matsuzawa  reported the death of a 2.5-year-old chimpanzee (Jokro) at Bossou from a respiratory illness. The infants mother (Jire) carried the corpse, mummified in the weeks following death, for at least 27 days. She exhibited extensive care of the body, grooming it regularly, sharing her day- and night-nests with it, and showing distress whenever they became separated. The carrying of infants corpses has been reported from a number of primate species, both in captivity and the wild [3-7] - albeit usually lasting a few days only - suggesting a phylogenetic continuity for a behavior that is poignant testament to the close mother-infant bond which extends across different primate taxa. In this report we recount two further infant deaths at Bossou, observed over a decade after the original episode but with striking similarities.
Animals that travel together in groups display a variety of fascinating motion patterns thought to be the result of delicate local interactions among group members. Although the most informative way of investigating and interpreting collective movement phenomena would be afforded by the collection of high-resolution spatiotemporal data from moving individuals, such data are scarce and are virtually non-existent for long-distance group motion within a natural setting because of the associated technological difficulties. Here we present results of experiments in which track logs of homing pigeons flying in flocks of up to 10 individuals have been obtained by high-resolution lightweight GPS devices and analysed using a variety of correlation functions inspired by approaches common in statistical physics. We find a well-defined hierarchy among flock members from data concerning leading roles in pairwise interactions, defined on the basis of characteristic delay times between birds directional choices. The average spatial position of a pigeon within the flock strongly correlates with its place in the hierarchy, and birds respond more quickly to conspecifics perceived primarily through the left eye-both results revealing differential roles for birds that assume different positions with respect to flock-mates. From an evolutionary perspective, our results suggest that hierarchical organization of group flight may be more efficient than an egalitarian one, at least for those flock sizes that permit regular pairwise interactions among group members, during which leader-follower relationships are consistently manifested.
Recent etho-archaeological studies of stone-tool use by wild chimpanzees have contributed valuable data towards elucidating the variables that influenced the emergence and development of the first lithic industries among Plio-Pleistocene hominins. Such data help to identify potential behaviours entailed in the first percussive technologies that are invisible in archaeological records. The long-term research site of Bossou in Guinea features a unique chimpanzee community whose members systematically use portable stones as hammers and anvils to crack open nuts in natural as well as in field experimental settings. Here we present the first analysis of repeated reuse of the same tool-composites in wild chimpanzees. Data collected over 5 years of experimental nut-cracking sessions at an "outdoor laboratory" site were assessed for the existence of systematic patterns in the selection of tool-composites, at group and at individual levels. Chimpanzees combined certain stones as hammer and anvil more often than expected by chance, even when taking into account preferences for individual stones by themselves. This may reflect an ability to recognise the nut-cracker as a single tool (composed of two elements, but functional only as a whole), as well as discrimination of tool quality-effectiveness. Through repeatedly combining the same pairs of stones--whether due to preferences for particular composites or for the two elements independently--tool-users may amplify use-wear traces and increase the likelihood of fracturing the stones, and thus of detaching pieces by battering.
Chimpanzees (Pan troglodytes) are known to make and use a variety of tools, activities which require them to employ their hands in a skilful manner. The learning process underlying the acquisition of tool-using skills, and the degree of laterality evident in both immature and mature performers are investigated here focusing on the use of leaves for drinking water by members of the Bossou chimpanzee community in Guinea, West Africa. In addition, comparisons are drawn between the present findings and our previous data on the cracking of oil-palm nuts (Elaeis guineensis) using stone tools by members of the same community. The use of leaves for drinking water emerges approximately 2 years earlier than nut cracking, at around the age of 1.5 years, although the manufacture of leaf tools begins only at 3.5 years of age. In addition, in clear contrast with nut cracking, the majority of chimpanzees are ambidextrous in their use of leaves, with only certain individuals showing a bias for one hand. We discuss possible explanations for the earlier emergence and increased ambidextrousness that characterises leaf-tool use in comparison with other forms of tool use by wild chimpanzees. In summary, our results provide the first detailed description of the acquisition process underlying leaf-tool use along with the accompanying patterns of handedness, while also being the first to provide comparisons of the development of different forms of tool use within the same wild chimpanzee population.
For animals that travel in groups, the directional choices of conspecifics are potentially a rich source of information for spatial learning. In this study, we investigate how the opportunity to follow a locally experienced demonstrator affects route learning by pigeons over repeated homing flights. This test of social influences on navigation takes advantage of the individually distinctive routes that pigeons establish when trained alone. We found that pigeons learn routes just as effectively while flying with a partner as control pigeons do while flying alone. However, rather than learning the exact route of the demonstrator, the paired routes shifted over repeated flights, which suggests that the birds with less local experience also took an active role in the navigational task. The efficiency of the original routes was a key factor in how far they shifted, with less efficient routes undergoing the greatest changes. In this context, inefficient routes are unlikely to be maintained through repeated rounds of social transmission, and instead more efficient routes are achieved because of the interaction between social learning and information pooling.
We report on the development of a novel shared touch-panel apparatus for examining a diverse range of topics in great ape social cognition and interaction. Our apparatus-named the Arena System-is composed of a single multitouch monitor that spans across two separate testing booths, so that individuals situated in each booth have tactile access to half of the monitor and visual access to the whole monitor. Additional components of the system include a smart-film barrier able to restrict visual access between the booths, as well as two automated feeding devices that dispense food rewards to the subjects. The touch-panel, smart-film, and feeders are controlled by a PC that is also responsible for running the experimental tasks. We present data from a pilot behavioral game theory study with two chimpanzees in order to illustrate the efficacy of our method, and we suggest applications for a range of topics including animal social learning, coordination, and behavioral economics. The system enables fully automated experimental procedures, which means that no human participation is needed to run the tasks. The novel use of a touch-panel in a social setting allows for a finer degree of data resolution than do the traditional experimental apparatuses used in prior studies on great ape social interaction.
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