Dora Biro - Profile on Academia.edu (original) (raw)

Papers by Dora Biro

The field of collective animal behaviour examines how relatively simple, local interactions betwe... more The field of collective animal behaviour examines how relatively simple, local interactions between individuals in groups combine to produce global-level outcomes. Existing mathematical models and empirical work have identified candidate mechanisms for numerous collective phenomena but have typically focused on one-off or short-term performance. We argue that feedback between collective performance and learning – giving the former the capacity to become an adaptive, and potentially cumulative, process – is a currently poorly explored but crucial mechanism in understanding collective systems. We synthesise material ranging from swarm intelligence in social insects through collective movements in vertebrates to collective decision making in animal and human groups, to propose avenues for future research to identify the potential for changes in these systems to accumulate over time. What Are Collective Behaviours and How Do They Arise? Some of the most impressive biological phenomena emerge out of interactions among members of animal groups. Bird flocks, fish schools, and insect swarms perform highly coordinated collective movements that can encompass thousands of individuals, producing complex group-level patterns that are difficult to predict from the behaviour of isolated individuals only. Animal groups are also able to solve problems that are beyond the capacities of single individuals [1]; ant colonies, for example, tackle certain types of optimisation problems so effectively that they have inspired an entire field of computer science [2]. Despite the appearance of synchronised organisation, it is increasingly well understood that no central control acts on the collective as a whole; instead, the global patterns result from simple, local interactions among the group's neighbouring members – a form of biological self-organisation [3] (see Glossary). Recent years have seen a proliferation of both empirical and theoretical work on the mechanistic underpinnings of collective animal behaviour [4], with self-organisation emerging as a major principle in various contexts including collective motion [5], decision making [6] and construction [7], activity synchronisation [8], and the spontaneous emergence of leader–follower relations [9]. Nonetheless, a rigorous adaptive framework is yet to be applied to collective animal behaviour; little is known about the nature of the selective forces that act at the level of the individual behavioural rules to shape pattern formation at group level. Over shorter timescales, and crucially for this review, no major synthesis has yet examined collective behaviour from a time–depth perspective; we do not know: (i) what changes group-level organisation might undergo over the course of repeated executions of collective tasks; (ii) to what extent solutions arrived at collectively are retained (learned), either at the individual or at the collective level, with the potential to influence future interactions; or (iii) what effect changes in group composition, due to natural demographic processes, have on whether solutions are 'inherited' from previous generations.

Visually-mediated site recognition by the homing pigeon may rely on a snapshot-like mechanism

Mechanisms of visually mediated site recognition by the homing pigeon

Animal Behaviour, 2003

The recognition of familiar areas by homing pigeons, Columba livia, is now known to depend at lea... more The recognition of familiar areas by homing pigeons, Columba livia, is now known to depend at least in part on visual cues. Birds allowed a 5-min preview of the surrounding landscape prior to release home faster than those denied access to such cues, suggesting that ...

Proceedings of the National Academy of Sciences of the United States of America, 2013

Hierarchical organization is widespread in the societies of humans and other animals, both in soc... more 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.

PloS one, 2013

Burmese long-tailed macaques (Macaca fascicularis aurea) are one of a limited number of wild anim... more 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.

Journal of the Royal Society, Interface / the Royal Society, 2013

Travelling in groups gives animals opportunities to share route information by following cues fro... more Travelling in groups gives animals opportunities to share route information by following cues from each other's 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 neighbour's 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.

Proceedings. Biological sciences / The Royal Society, 2013

The Journal of experimental biology, 2013

In many species, group members obtain benefits from moving collectively, such as enhanced foragin... more In many species, group members obtain benefits from moving collectively, such as enhanced foraging efficiency or increased predator detection. In situations where the groupʼs 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.

Animal Cognition, 2009

Recent etho-archaeological studies of stone-tool use by wild chimpanzees have contributed valuabl... more 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.

Current biology : CB, 2012

Learning & Behavior, 2010

Field experiments can provide compelling demonstrations of social learning in wild populations. S... more 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.

Proceedings. Biological sciences / The Royal Society, 2005

The mechanisms used by homing pigeons (Columba livia) to navigate homeward from distant sites hav... more The mechanisms used by homing pigeons (Columba livia) to navigate homeward from distant sites have been well studied, yet the mechanisms underlying navigation within, and mapping of, the local familiar area have been largely neglected. In the local area pigeons pote ntially have access to a powerful navigational aid-a memorized landscape map. Current opinion suggests that landmarks are used only to recognize a familiar start position and that the goalward route is then achieved solely using compass orientation. We used highresolution global positioning system (GPS) loggers to track homing pigeons as they became progressively familiar with a local homing task. Here, we demonstrate that birds develop highly stereotyped yet individually distinctive routes over the landscape, which remain substantially inefficient. Precise aerial route recapitulation implies close control by localized geocentric cues. Magnetic cues are unlikely to have been used, since recapitulation remains despite magnetic disruption treatment, and olfactory cues would have been positionally unstable under the variable wind conditions, making visual landmarks the most likely cues used.

PloS one, 2013

The sun has long been thought to guide bird navigation as the second step in a two-stage process,... more 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 animal's endogenous clock time-compensates the solar compass for the sun's 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.

Animal cognition, 2011

Social animals have much to gain from observing and responding appropriately to the actions of th... more 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 model’s 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.

Theory in Biosciences, 2008

Moving animal groups provide some of the most intriguing and difficult to characterise examples o... more Moving animal groups provide some of the most intriguing and difficult to characterise examples of collective behaviour. We review some recent (and not so recent) empirical research on the motion of animal groups, including fish, locusts and homing pigeons. An important concept which unifies our understanding of these groups is that of transfer of directional information. Individuals which change their direction of travel in response to the direction taken by their near neighbours can quickly transfer information about the presence of a predatory threat or food source. We show that such information transfer is optimised when the density of individuals in a group is close to that at which a phase transition occurs between random and ordered motion. Similarly, we show that even relatively small differences in information possessed by group members can lead to strong collectivelevel decisions for one of two options. By combining the use of self-propelled particle and social force models of collective motion with thinking about the evolution of flocking we aim to better understand how complexity arises within these groups.

Nature, 2010

Animals that travel together in groups display a variety of fascinating motion patterns thought t... more 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 1-3 . 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 4-7 and are virtually non-existent for long-distance group motion within a natural setting because of the associated technological difficulties 8 . 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 analyzed using a variety of correlation functions inspired by approaches common in statistical physics. We find a welldefined 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 organisation 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 leaderfollower relationships are consistently manifested.

The field of collective animal behaviour examines how relatively simple, local interactions betwe... more The field of collective animal behaviour examines how relatively simple, local interactions between individuals in groups combine to produce global-level outcomes. Existing mathematical models and empirical work have identified candidate mechanisms for numerous collective phenomena but have typically focused on one-off or short-term performance. We argue that feedback between collective performance and learning – giving the former the capacity to become an adaptive, and potentially cumulative, process – is a currently poorly explored but crucial mechanism in understanding collective systems. We synthesise material ranging from swarm intelligence in social insects through collective movements in vertebrates to collective decision making in animal and human groups, to propose avenues for future research to identify the potential for changes in these systems to accumulate over time. What Are Collective Behaviours and How Do They Arise? Some of the most impressive biological phenomena emerge out of interactions among members of animal groups. Bird flocks, fish schools, and insect swarms perform highly coordinated collective movements that can encompass thousands of individuals, producing complex group-level patterns that are difficult to predict from the behaviour of isolated individuals only. Animal groups are also able to solve problems that are beyond the capacities of single individuals [1]; ant colonies, for example, tackle certain types of optimisation problems so effectively that they have inspired an entire field of computer science [2]. Despite the appearance of synchronised organisation, it is increasingly well understood that no central control acts on the collective as a whole; instead, the global patterns result from simple, local interactions among the group's neighbouring members – a form of biological self-organisation [3] (see Glossary). Recent years have seen a proliferation of both empirical and theoretical work on the mechanistic underpinnings of collective animal behaviour [4], with self-organisation emerging as a major principle in various contexts including collective motion [5], decision making [6] and construction [7], activity synchronisation [8], and the spontaneous emergence of leader–follower relations [9]. Nonetheless, a rigorous adaptive framework is yet to be applied to collective animal behaviour; little is known about the nature of the selective forces that act at the level of the individual behavioural rules to shape pattern formation at group level. Over shorter timescales, and crucially for this review, no major synthesis has yet examined collective behaviour from a time–depth perspective; we do not know: (i) what changes group-level organisation might undergo over the course of repeated executions of collective tasks; (ii) to what extent solutions arrived at collectively are retained (learned), either at the individual or at the collective level, with the potential to influence future interactions; or (iii) what effect changes in group composition, due to natural demographic processes, have on whether solutions are 'inherited' from previous generations.

Visually-mediated site recognition by the homing pigeon may rely on a snapshot-like mechanism

Mechanisms of visually mediated site recognition by the homing pigeon

Animal Behaviour, 2003

The recognition of familiar areas by homing pigeons, Columba livia, is now known to depend at lea... more The recognition of familiar areas by homing pigeons, Columba livia, is now known to depend at least in part on visual cues. Birds allowed a 5-min preview of the surrounding landscape prior to release home faster than those denied access to such cues, suggesting that ...

Proceedings of the National Academy of Sciences of the United States of America, 2013

Hierarchical organization is widespread in the societies of humans and other animals, both in soc... more 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.

PloS one, 2013

Burmese long-tailed macaques (Macaca fascicularis aurea) are one of a limited number of wild anim... more 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.

Journal of the Royal Society, Interface / the Royal Society, 2013

Travelling in groups gives animals opportunities to share route information by following cues fro... more Travelling in groups gives animals opportunities to share route information by following cues from each other's 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 neighbour's 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.

Proceedings. Biological sciences / The Royal Society, 2013

The Journal of experimental biology, 2013

In many species, group members obtain benefits from moving collectively, such as enhanced foragin... more In many species, group members obtain benefits from moving collectively, such as enhanced foraging efficiency or increased predator detection. In situations where the groupʼs 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.

Animal Cognition, 2009

Recent etho-archaeological studies of stone-tool use by wild chimpanzees have contributed valuabl... more 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.

Current biology : CB, 2012

Learning & Behavior, 2010

Field experiments can provide compelling demonstrations of social learning in wild populations. S... more 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.

Proceedings. Biological sciences / The Royal Society, 2005

The mechanisms used by homing pigeons (Columba livia) to navigate homeward from distant sites hav... more The mechanisms used by homing pigeons (Columba livia) to navigate homeward from distant sites have been well studied, yet the mechanisms underlying navigation within, and mapping of, the local familiar area have been largely neglected. In the local area pigeons pote ntially have access to a powerful navigational aid-a memorized landscape map. Current opinion suggests that landmarks are used only to recognize a familiar start position and that the goalward route is then achieved solely using compass orientation. We used highresolution global positioning system (GPS) loggers to track homing pigeons as they became progressively familiar with a local homing task. Here, we demonstrate that birds develop highly stereotyped yet individually distinctive routes over the landscape, which remain substantially inefficient. Precise aerial route recapitulation implies close control by localized geocentric cues. Magnetic cues are unlikely to have been used, since recapitulation remains despite magnetic disruption treatment, and olfactory cues would have been positionally unstable under the variable wind conditions, making visual landmarks the most likely cues used.

PloS one, 2013

The sun has long been thought to guide bird navigation as the second step in a two-stage process,... more 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 animal's endogenous clock time-compensates the solar compass for the sun's 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.

Animal cognition, 2011

Social animals have much to gain from observing and responding appropriately to the actions of th... more 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 model’s 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.

Theory in Biosciences, 2008

Moving animal groups provide some of the most intriguing and difficult to characterise examples o... more Moving animal groups provide some of the most intriguing and difficult to characterise examples of collective behaviour. We review some recent (and not so recent) empirical research on the motion of animal groups, including fish, locusts and homing pigeons. An important concept which unifies our understanding of these groups is that of transfer of directional information. Individuals which change their direction of travel in response to the direction taken by their near neighbours can quickly transfer information about the presence of a predatory threat or food source. We show that such information transfer is optimised when the density of individuals in a group is close to that at which a phase transition occurs between random and ordered motion. Similarly, we show that even relatively small differences in information possessed by group members can lead to strong collectivelevel decisions for one of two options. By combining the use of self-propelled particle and social force models of collective motion with thinking about the evolution of flocking we aim to better understand how complexity arises within these groups.

Nature, 2010

Animals that travel together in groups display a variety of fascinating motion patterns thought t... more 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 1-3 . 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 4-7 and are virtually non-existent for long-distance group motion within a natural setting because of the associated technological difficulties 8 . 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 analyzed using a variety of correlation functions inspired by approaches common in statistical physics. We find a welldefined 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 organisation 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 leaderfollower relationships are consistently manifested.