1. We estimate colony reproductive success, in numbers of offspring colonies arising from a colon... more 1. We estimate colony reproductive success, in numbers of offspring colonies arising from a colony's daughter queens, of colonies of the red harvester ant, Pogonomyrmex barbatus. 2. A measure of lifetime reproductive success is essential to understand the relation of ecological factors, phenotype and fitness in a natural population. This was possible for the first time in a natural population of ant colonies using data from long-term study of a population of colonies in southeastern Arizona, for which ages of all colonies are known from census data collected since 1985. 3. Parentage analyses of microsatellite data from 5 highly polymorphic loci were used to assign offspring colonies to maternal parent colonies in a population of about 265 colonies, ages 1–28 years, sampled in 2010. 4. The estimated population growth rate R o was 1Á69 and generation time was 7Á8 years. There was considerable variation among colonies in reproductive success: of 199 possible parent colonies , only 49 (~ 25%) had offspring colonies on the site. The mean number of offspring colonies per maternal parent colony was 2Á94 and ranged from 1 to 8. A parent was identified for the queen of 146 of 247 offspring colonies. There was no evidence for reproductive senescence; fecundity was about the same throughout the 25–30 year lifespan of a colony. 5. There were no trends in the distance or direction of the dispersal of an offspring relative to its maternal parent colony. There was no relationship between the number of gynes produced by a colony in 1 year and the number of offspring colonies subsequently founded by its daughter reproductive females. 6. The results provide the first estimate of a life table for a population of ant colonies and the first estimate of the female component of colony lifetime reproductive success. The results suggest that commonly used measures of reproductive output may not be correlated with realized reproductive success. This is the starting point for future investigation asking whether variation in reproductive success is related to phenotypic variation among colonies in behavioural and ecological traits.
Collective behavior is the outcome of a network of local interactions. Here, I consider collectiv... more Collective behavior is the outcome of a network of local interactions. Here, I consider collective behavior as the result of algorithms that have evolved to operate in response to a particular environment and physiological context. I discuss how algorithms are shaped by the costs of operating under the constraints that the environment imposes, the extent to which the environment is stable, and the distribution, in space and time, of resources. I suggest that a focus on the dynamics of the environment may provide new hypotheses for elucidating the algorithms that produce the collective behavior of cellular systems.
In the 0 5 summer blockbuster Ant-Man, the character Hank Pym, a scientist who has invented a sui... more In the 0 5 summer blockbuster Ant-Man, the character Hank Pym, a scientist who has invented a suit that can shrink a person down to the size of an insect, remarks that ants can perform amazing feats, but they need a leader to tell them what to do. Pym wears a small device behind his ear that allows him to instruct the ants to act as a phalanx of attackers that helps the antsized human hero defeat an evil mastermind.
The problem of collective search is a tradeoff between searching thoroughly and covering as much ... more The problem of collective search is a tradeoff between searching thoroughly and covering as much area as possible. This tradeoff depends on the density of searchers. Solutions to the problem of collective search are currently of much interest in robotics and in the study of distributed algorithms, for example to design ways that without central control robots can use local information to perform search and rescue operations. Ant colonies operate without central control. Because they can perceive only local, mostly chemical and tactile cues, they must search collectively to find resources and to monitor the colony's environment. Examining how ants in diverse environments solve the problem of collective search can elucidate how evolution has led to diverse forms of collective behavior. An experiment on the International Space Station in January 2014 examined how ants (Tetramorium caespitum) perform collective search in microgravity. In the ISS experiment, the ants explored a small arena in which a barrier was lowered to increase the area and thus lower ant density. In microgravity, relative to ground controls, ants explored the area less thoroughly and took more convoluted paths. It appears that the difficulty of holding on to the surface interfered with the ants' ability to search collectively. Ants frequently lost contact with the surface, but showed a remarkable ability to regain contact with the surface.
'Division of labor' is a misleading way to describe the organization of tasks in social insect co... more 'Division of labor' is a misleading way to describe the organization of tasks in social insect colonies, because there is little evidence for persistent individual specialization in task. Instead, task allocation in social insects occurs through distributed processes whose advantages, such as resilience, differ from those of division of labor, which are mostly based on learning. The use of the phrase 'division of labor' persists for historical reasons, and tends to focus attention on differences among individuals in internal attributes. This focus distracts from the main questions of interest in current research, which require an understanding of how individuals interact with each other and their environments. These questions include how colony behavior is regulated, how the regulation of colony behavior develops over the lifetime of a colony, what are the sources of variation among colonies in the regulation of behavior, and how the collective regulation of colony behavior evolves.
We propose a distributed model of nestmate recognition, analogous to the one used by the vertebra... more We propose a distributed model of nestmate recognition, analogous to the one used by the vertebrate immune system, in which colony response results from the diverse reactions of many ants. The model describes how individual behaviour produces colony response to non-nestmates. No single ant knows the odour identity of the colony. Instead, colony identity is defined collectively by all the ants in the colony. Each ant responds to the odour of other ants by reference to its own unique decision boundary, which is a result of its experience of encounters with other ants. Each ant thus recognizes a particular set of chemical profiles as being those of non-nestmates. This model predicts, as experimental results have shown, that the outcome of behavioural assays is likely to be variable, that it depends on the number of ants tested, that response to non-nestmates changes over time and that it changes in response to the experience of individual ants. A distributed system allows a colony to identify non-nestmates without requiring that all individuals have the same complete information and helps to facilitate the tracking of changes in cuticular hydrocarbon profiles, because only a subset of ants must respond to provide an adequate response.
Harvester ant colonies adjust their foraging activity to day-today changes in food availability a... more Harvester ant colonies adjust their foraging activity to day-today changes in food availability and hour-to-hour changes in environmental conditions. This collective behavior is regulated through interactions, in the form of brief antennal contacts, between outgoing foragers and returning foragers with food. Here we consider how an ant, waiting in the entrance chamber just inside the nest entrance, uses its accumulated experience of interactions to decide whether to leave the nest to forage. Using videos of field observations, we tracked the interactions and foraging decisions of ants in the entrance chamber. Outgoing foragers tended to interact with returning foragers at higher rates than ants that returned to the deeper nest and did not forage. To provide a mechanistic framework for interpreting these results, we develop a decision model in which ants make decisions based upon a noisy accumulation of individual contacts with returning foragers. The model can reproduce core trends and realistic distributions for individual ant interaction statistics, and suggests possible mechanisms by which foraging activity may be regulated at an individual ant level.
Similar patterns of inter- action, such as network motifs and feedback loops, are used in many na... more Similar patterns of inter- action, such as network motifs and feedback loops, are used in many natural collective processes, proba- bly because they have evolved independently under similar pres- sures. Here I consider how three environmental constraints may shape the evolution of collective behavior: the patchiness of resourc- es, the operating costs of maintain- ing the interaction network that produces collective behavior, and the threat of rupture of the net- work. The ants are a large and successful taxon that have evolved in very diverse environments. Ex- amples from ants provide a starting point for examining more generally the fit between the particular pattern of interaction that regu- lates activity, and the environment in which it functions.
Collective behavior is the outcome of a network of local interactions. Here, I consider collectiv... more Collective behavior is the outcome of a network of local interactions. Here, I consider collective behavior as the result of algorithms that have evolved to operate in response to a particular environment and physiological context. I discuss how algorithms are shaped by the costs of operating under the constraints that the environment imposes, the extent to which the environment is stable, and the distribution, in space and time, of resources. I suggest that a focus on the dynamics of the environment may provide new hypotheses for elucidating the algorithms that produce the collective behavior of cellular systems.
'Division of labor' is a misleading way to describe the organization of tasks in social insect co... more 'Division of labor' is a misleading way to describe the organization of tasks in social insect colonies, because there is little evidence for persistent individual specialization in task. Instead, task allocation in social insects occurs through distributed processes whose advantages, such as resilience, differ from those of division of labor, which are mostly based on learning. The use of the phrase 'division of labor' persists for historical reasons, and tends to focus attention on differences among individuals in internal attributes. This focus distracts from the main questions of interest in current research, which require an understanding of how individuals interact with each other and their environments. These questions include how colony behavior is regulated, how the regulation of colony behavior develops over the lifetime of a colony, what are the sources of variation among colonies in the regulation of behavior, and how the collective regulation of colony behavior evolves.
Many exciting studies have begun to elucidate the genetics of the morphological and physiological... more Many exciting studies have begun to elucidate the genetics of the morphological and physiological diversity of ants, but as yet few studies have investigated the genetics of ant behavior directly. Ant genomes are marked by extreme rates of gene turnover, especially in gene families related to ol-factory communication, such as the synthesis of cuticular hydrocarbons and the perception of environmental semiochemicals. Transcriptomic and epi-genetic differences are apparent between reproductive and sterile females, males and females, and workers that differ in body size. Quantitative genetic approaches suggest heritability of task performance, and population genetic studies indicate a genetic association with reproductive status in some species. Gene expression is associated with behavior including foraging, response to queens attempting to join a colony, circadian patterns of task performance, and age-related changes of task. Ant behavioral genetics needs further investigation of the feedback between individual-level physiological changes and socially mediated responses to environmental conditions.
'Division of labor' is a misleading way to describe the organization of tasks in social insect co... more 'Division of labor' is a misleading way to describe the organization of tasks in social insect colonies, because there is little evidence for persistent individual specialization in task. Instead, task allocation in social insects occurs through distributed processes whose advantages, such as resilience, differ from those of division of labor, which are mostly based on learning. The use of the phrase 'division of labor' persists for historical reasons, and tends to focus attention on differences among individuals in internal attributes. This focus distracts from the main questions of interest in current research, which require an understanding of how individuals interact with each other and their environments. These questions include how colony behavior is regulated, how the regulation of colony behavior develops over the lifetime of a colony, what are the sources of variation among colonies in the regulation of behavior, and how the collective regulation of colony behavior evolves.
Harvester ant colonies adjust their foraging activity to day-today changes in food availability a... more Harvester ant colonies adjust their foraging activity to day-today changes in food availability and hour-to-hour changes in environmental conditions. This collective behavior is regulated through interactions, in the form of brief antennal contacts, between outgoing foragers and returning foragers with food. Here we consider how an ant, waiting in the entrance chamber just inside the nest entrance, uses its accumulated experience of interactions to decide whether to leave the nest to forage. Using videos of field observations, we tracked the interactions and foraging decisions of ants in the entrance chamber. Outgoing foragers tended to interact with returning foragers at higher rates than ants that returned to the deeper nest and did not forage. To provide a mechanistic framework for interpreting these results, we develop a decision model in which ants make decisions based upon a noisy accumulation of individual contacts with returning foragers. The model can reproduce core trends and realistic distributions for individual ant interaction statistics, and suggests possible mechanisms by which foraging activity may be regulated at an individual ant level.
We propose a distributed model of nestmate recognition, analogous to the one used by the vertebra... more We propose a distributed model of nestmate recognition, analogous to the one used by the vertebrate immune system, in which colony response results from the diverse reactions of many ants. The model describes how individual behaviour produces colony response to non-nestmates. No single ant knows the odour identity of the colony. Instead, colony identity is defined collectively by all the ants in the colony. Each ant responds to the odour of other ants by reference to its own unique decision boundary, which is a result of its experience of encounters with other ants. Each ant thus recognizes a particular set of chemical profiles as being those of non-nestmates. This model predicts, as experimental results have shown, that the outcome of behavioural assays is likely to be variable, that it depends on the number of ants tested, that response to non-nestmates changes over time and that it changes in response to the experience of individual ants. A distributed system allows a colony to identify non-nestmates without requiring that all individuals have the same complete information and helps to facilitate the tracking of changes in cuticular hydrocarbon profiles, because only a subset of ants must respond to provide an adequate response.
The problem of collective search is a tradeoff between searching thoroughly and covering as much ... more The problem of collective search is a tradeoff between searching thoroughly and covering as much area as possible. This tradeoff depends on the density of searchers. Solutions to the problem of collective search are currently of much interest in robotics and in the study of distributed algorithms, for example to design ways that without central control robots can use local information to perform search and rescue operations. Ant colonies operate without central control. Because they can perceive only local, mostly chemical and tactile cues, they must search collectively to find resources and to monitor the colony's environment. Examining how ants in diverse environments solve the problem of collective search can elucidate how evolution has led to diverse forms of collective behavior. An experiment on the International Space Station in January 2014 examined how ants (Tetramorium caespitum) perform collective search in microgravity. In the ISS experiment, the ants explored a small arena in which a barrier was lowered to increase the area and thus lower ant density. In microgravity, relative to ground controls, ants explored the area less thoroughly and took more convoluted paths. It appears that the difficulty of holding on to the surface interfered with the ants' ability to search collectively. Ants frequently lost contact with the surface, but showed a remarkable ability to regain contact with the surface.
This field study examines the social behavior of five sympatric species of desert seed-eating ant... more This field study examines the social behavior of five sympatric species of desert seed-eating ants (Pogonomyrmex barbatus, P. rugosus, P. maricopa, P. desertorum, P. californicus). The species differed significantly in measures of activity rhythms in various colony tasks, use of space around the nest yard, and reaction to disturbance. Species differences were related to the typical size of a colony's outside work force.
Invasive species, where successful, can devastate native communities. We studied the dynamics of ... more Invasive species, where successful, can devastate native communities. We studied the dynamics of the invasive Argentine ant, Linepithema humile, for 7 years in Jasper Ridge, a biological preserve in northern California. We monitored the distributions at the hectare scale of native ant taxa and L. humile in the spring and fall from 1993 to 1999. We also studied the invasion dynamics at a finer resolution by searching for ants in 1-m 2 plots. Our results are similar at both scales. The distributions of several native species are not random with regard to L. humile; the distributions of several epigeic species with similar habitat affinities overlap much less frequently than expected with the distribution of L. humile. We found that season had a significant influence on the distributions of L. humile and several native taxa. Over the 7-year period, L. humile has increased its range size in Jasper Ridge largely at the expense of native taxa, but there is seasonal and yearly variation in this rate of increase. Studies of invasions in progress which sample across seasons and years may help to predict the spread and effects of invasive species.
In the seed-eating ant Pogonomyrmex barbatus, the return of successful foragers stimulates inacti... more In the seed-eating ant Pogonomyrmex barbatus, the return of successful foragers stimulates inactive foragers to leave the nest. The rate at which successful foragers return to the nest depends on food availability; the more food available, the more quickly foragers will find it and bring it back. Field experiments examined how quickly a colony can adjust to a decline in the rate of forager return, and thus to a decline in food availability, by slowing down foraging activity. In response to a brief, 3-to 5-min reduction in the forager return rate, foraging activity usually decreased within 2-3 min and then recovered within 5 min. This indicates that whether an inactive forager leaves the nest on its next trip depends on its very recent experience of the rate of forager return. On some days, colonies responded more to a change in forager return rate. The rapid colony response to fluctuations in forager return rate, enabling colonies to act as risk-averse foragers, may arise from the limited interval over which an ant can track its encounters with returning foragers.
Colonies of the harvester ant, Pogonomyrmex barbatus, adjust the direction and length of foraging... more Colonies of the harvester ant, Pogonomyrmex barbatus, adjust the direction and length of foraging trails in response to the foraging behavior of their conspecific neighbors. In the absence of any interaction with its neighbor, a mature colony expands its foraging range at a rate of 0.85 + 0.15 m per day. Exclusion experiments show that if a colony is prevented from using its foraging trails, the neighbors of that colony will enter its foraging range within 10 days. Exclusion experiments were performed with three age classes of colonies: young (1 year old), intermediate (3-4 years old), and old (5 years old or more). Colonies 3 4 years old are most likely to expand foraging ranges, and to retain newly-gained areas.
1. We estimate colony reproductive success, in numbers of offspring colonies arising from a colon... more 1. We estimate colony reproductive success, in numbers of offspring colonies arising from a colony's daughter queens, of colonies of the red harvester ant, Pogonomyrmex barbatus. 2. A measure of lifetime reproductive success is essential to understand the relation of ecological factors, phenotype and fitness in a natural population. This was possible for the first time in a natural population of ant colonies using data from long-term study of a population of colonies in southeastern Arizona, for which ages of all colonies are known from census data collected since 1985. 3. Parentage analyses of microsatellite data from 5 highly polymorphic loci were used to assign offspring colonies to maternal parent colonies in a population of about 265 colonies, ages 1–28 years, sampled in 2010. 4. The estimated population growth rate R o was 1Á69 and generation time was 7Á8 years. There was considerable variation among colonies in reproductive success: of 199 possible parent colonies , only 49 (~ 25%) had offspring colonies on the site. The mean number of offspring colonies per maternal parent colony was 2Á94 and ranged from 1 to 8. A parent was identified for the queen of 146 of 247 offspring colonies. There was no evidence for reproductive senescence; fecundity was about the same throughout the 25–30 year lifespan of a colony. 5. There were no trends in the distance or direction of the dispersal of an offspring relative to its maternal parent colony. There was no relationship between the number of gynes produced by a colony in 1 year and the number of offspring colonies subsequently founded by its daughter reproductive females. 6. The results provide the first estimate of a life table for a population of ant colonies and the first estimate of the female component of colony lifetime reproductive success. The results suggest that commonly used measures of reproductive output may not be correlated with realized reproductive success. This is the starting point for future investigation asking whether variation in reproductive success is related to phenotypic variation among colonies in behavioural and ecological traits.
Collective behavior is the outcome of a network of local interactions. Here, I consider collectiv... more Collective behavior is the outcome of a network of local interactions. Here, I consider collective behavior as the result of algorithms that have evolved to operate in response to a particular environment and physiological context. I discuss how algorithms are shaped by the costs of operating under the constraints that the environment imposes, the extent to which the environment is stable, and the distribution, in space and time, of resources. I suggest that a focus on the dynamics of the environment may provide new hypotheses for elucidating the algorithms that produce the collective behavior of cellular systems.
In the 0 5 summer blockbuster Ant-Man, the character Hank Pym, a scientist who has invented a sui... more In the 0 5 summer blockbuster Ant-Man, the character Hank Pym, a scientist who has invented a suit that can shrink a person down to the size of an insect, remarks that ants can perform amazing feats, but they need a leader to tell them what to do. Pym wears a small device behind his ear that allows him to instruct the ants to act as a phalanx of attackers that helps the antsized human hero defeat an evil mastermind.
The problem of collective search is a tradeoff between searching thoroughly and covering as much ... more The problem of collective search is a tradeoff between searching thoroughly and covering as much area as possible. This tradeoff depends on the density of searchers. Solutions to the problem of collective search are currently of much interest in robotics and in the study of distributed algorithms, for example to design ways that without central control robots can use local information to perform search and rescue operations. Ant colonies operate without central control. Because they can perceive only local, mostly chemical and tactile cues, they must search collectively to find resources and to monitor the colony's environment. Examining how ants in diverse environments solve the problem of collective search can elucidate how evolution has led to diverse forms of collective behavior. An experiment on the International Space Station in January 2014 examined how ants (Tetramorium caespitum) perform collective search in microgravity. In the ISS experiment, the ants explored a small arena in which a barrier was lowered to increase the area and thus lower ant density. In microgravity, relative to ground controls, ants explored the area less thoroughly and took more convoluted paths. It appears that the difficulty of holding on to the surface interfered with the ants' ability to search collectively. Ants frequently lost contact with the surface, but showed a remarkable ability to regain contact with the surface.
'Division of labor' is a misleading way to describe the organization of tasks in social insect co... more 'Division of labor' is a misleading way to describe the organization of tasks in social insect colonies, because there is little evidence for persistent individual specialization in task. Instead, task allocation in social insects occurs through distributed processes whose advantages, such as resilience, differ from those of division of labor, which are mostly based on learning. The use of the phrase 'division of labor' persists for historical reasons, and tends to focus attention on differences among individuals in internal attributes. This focus distracts from the main questions of interest in current research, which require an understanding of how individuals interact with each other and their environments. These questions include how colony behavior is regulated, how the regulation of colony behavior develops over the lifetime of a colony, what are the sources of variation among colonies in the regulation of behavior, and how the collective regulation of colony behavior evolves.
We propose a distributed model of nestmate recognition, analogous to the one used by the vertebra... more We propose a distributed model of nestmate recognition, analogous to the one used by the vertebrate immune system, in which colony response results from the diverse reactions of many ants. The model describes how individual behaviour produces colony response to non-nestmates. No single ant knows the odour identity of the colony. Instead, colony identity is defined collectively by all the ants in the colony. Each ant responds to the odour of other ants by reference to its own unique decision boundary, which is a result of its experience of encounters with other ants. Each ant thus recognizes a particular set of chemical profiles as being those of non-nestmates. This model predicts, as experimental results have shown, that the outcome of behavioural assays is likely to be variable, that it depends on the number of ants tested, that response to non-nestmates changes over time and that it changes in response to the experience of individual ants. A distributed system allows a colony to identify non-nestmates without requiring that all individuals have the same complete information and helps to facilitate the tracking of changes in cuticular hydrocarbon profiles, because only a subset of ants must respond to provide an adequate response.
Harvester ant colonies adjust their foraging activity to day-today changes in food availability a... more Harvester ant colonies adjust their foraging activity to day-today changes in food availability and hour-to-hour changes in environmental conditions. This collective behavior is regulated through interactions, in the form of brief antennal contacts, between outgoing foragers and returning foragers with food. Here we consider how an ant, waiting in the entrance chamber just inside the nest entrance, uses its accumulated experience of interactions to decide whether to leave the nest to forage. Using videos of field observations, we tracked the interactions and foraging decisions of ants in the entrance chamber. Outgoing foragers tended to interact with returning foragers at higher rates than ants that returned to the deeper nest and did not forage. To provide a mechanistic framework for interpreting these results, we develop a decision model in which ants make decisions based upon a noisy accumulation of individual contacts with returning foragers. The model can reproduce core trends and realistic distributions for individual ant interaction statistics, and suggests possible mechanisms by which foraging activity may be regulated at an individual ant level.
Similar patterns of inter- action, such as network motifs and feedback loops, are used in many na... more Similar patterns of inter- action, such as network motifs and feedback loops, are used in many natural collective processes, proba- bly because they have evolved independently under similar pres- sures. Here I consider how three environmental constraints may shape the evolution of collective behavior: the patchiness of resourc- es, the operating costs of maintain- ing the interaction network that produces collective behavior, and the threat of rupture of the net- work. The ants are a large and successful taxon that have evolved in very diverse environments. Ex- amples from ants provide a starting point for examining more generally the fit between the particular pattern of interaction that regu- lates activity, and the environment in which it functions.
Collective behavior is the outcome of a network of local interactions. Here, I consider collectiv... more Collective behavior is the outcome of a network of local interactions. Here, I consider collective behavior as the result of algorithms that have evolved to operate in response to a particular environment and physiological context. I discuss how algorithms are shaped by the costs of operating under the constraints that the environment imposes, the extent to which the environment is stable, and the distribution, in space and time, of resources. I suggest that a focus on the dynamics of the environment may provide new hypotheses for elucidating the algorithms that produce the collective behavior of cellular systems.
'Division of labor' is a misleading way to describe the organization of tasks in social insect co... more 'Division of labor' is a misleading way to describe the organization of tasks in social insect colonies, because there is little evidence for persistent individual specialization in task. Instead, task allocation in social insects occurs through distributed processes whose advantages, such as resilience, differ from those of division of labor, which are mostly based on learning. The use of the phrase 'division of labor' persists for historical reasons, and tends to focus attention on differences among individuals in internal attributes. This focus distracts from the main questions of interest in current research, which require an understanding of how individuals interact with each other and their environments. These questions include how colony behavior is regulated, how the regulation of colony behavior develops over the lifetime of a colony, what are the sources of variation among colonies in the regulation of behavior, and how the collective regulation of colony behavior evolves.
Many exciting studies have begun to elucidate the genetics of the morphological and physiological... more Many exciting studies have begun to elucidate the genetics of the morphological and physiological diversity of ants, but as yet few studies have investigated the genetics of ant behavior directly. Ant genomes are marked by extreme rates of gene turnover, especially in gene families related to ol-factory communication, such as the synthesis of cuticular hydrocarbons and the perception of environmental semiochemicals. Transcriptomic and epi-genetic differences are apparent between reproductive and sterile females, males and females, and workers that differ in body size. Quantitative genetic approaches suggest heritability of task performance, and population genetic studies indicate a genetic association with reproductive status in some species. Gene expression is associated with behavior including foraging, response to queens attempting to join a colony, circadian patterns of task performance, and age-related changes of task. Ant behavioral genetics needs further investigation of the feedback between individual-level physiological changes and socially mediated responses to environmental conditions.
'Division of labor' is a misleading way to describe the organization of tasks in social insect co... more 'Division of labor' is a misleading way to describe the organization of tasks in social insect colonies, because there is little evidence for persistent individual specialization in task. Instead, task allocation in social insects occurs through distributed processes whose advantages, such as resilience, differ from those of division of labor, which are mostly based on learning. The use of the phrase 'division of labor' persists for historical reasons, and tends to focus attention on differences among individuals in internal attributes. This focus distracts from the main questions of interest in current research, which require an understanding of how individuals interact with each other and their environments. These questions include how colony behavior is regulated, how the regulation of colony behavior develops over the lifetime of a colony, what are the sources of variation among colonies in the regulation of behavior, and how the collective regulation of colony behavior evolves.
Harvester ant colonies adjust their foraging activity to day-today changes in food availability a... more Harvester ant colonies adjust their foraging activity to day-today changes in food availability and hour-to-hour changes in environmental conditions. This collective behavior is regulated through interactions, in the form of brief antennal contacts, between outgoing foragers and returning foragers with food. Here we consider how an ant, waiting in the entrance chamber just inside the nest entrance, uses its accumulated experience of interactions to decide whether to leave the nest to forage. Using videos of field observations, we tracked the interactions and foraging decisions of ants in the entrance chamber. Outgoing foragers tended to interact with returning foragers at higher rates than ants that returned to the deeper nest and did not forage. To provide a mechanistic framework for interpreting these results, we develop a decision model in which ants make decisions based upon a noisy accumulation of individual contacts with returning foragers. The model can reproduce core trends and realistic distributions for individual ant interaction statistics, and suggests possible mechanisms by which foraging activity may be regulated at an individual ant level.
We propose a distributed model of nestmate recognition, analogous to the one used by the vertebra... more We propose a distributed model of nestmate recognition, analogous to the one used by the vertebrate immune system, in which colony response results from the diverse reactions of many ants. The model describes how individual behaviour produces colony response to non-nestmates. No single ant knows the odour identity of the colony. Instead, colony identity is defined collectively by all the ants in the colony. Each ant responds to the odour of other ants by reference to its own unique decision boundary, which is a result of its experience of encounters with other ants. Each ant thus recognizes a particular set of chemical profiles as being those of non-nestmates. This model predicts, as experimental results have shown, that the outcome of behavioural assays is likely to be variable, that it depends on the number of ants tested, that response to non-nestmates changes over time and that it changes in response to the experience of individual ants. A distributed system allows a colony to identify non-nestmates without requiring that all individuals have the same complete information and helps to facilitate the tracking of changes in cuticular hydrocarbon profiles, because only a subset of ants must respond to provide an adequate response.
The problem of collective search is a tradeoff between searching thoroughly and covering as much ... more The problem of collective search is a tradeoff between searching thoroughly and covering as much area as possible. This tradeoff depends on the density of searchers. Solutions to the problem of collective search are currently of much interest in robotics and in the study of distributed algorithms, for example to design ways that without central control robots can use local information to perform search and rescue operations. Ant colonies operate without central control. Because they can perceive only local, mostly chemical and tactile cues, they must search collectively to find resources and to monitor the colony's environment. Examining how ants in diverse environments solve the problem of collective search can elucidate how evolution has led to diverse forms of collective behavior. An experiment on the International Space Station in January 2014 examined how ants (Tetramorium caespitum) perform collective search in microgravity. In the ISS experiment, the ants explored a small arena in which a barrier was lowered to increase the area and thus lower ant density. In microgravity, relative to ground controls, ants explored the area less thoroughly and took more convoluted paths. It appears that the difficulty of holding on to the surface interfered with the ants' ability to search collectively. Ants frequently lost contact with the surface, but showed a remarkable ability to regain contact with the surface.
This field study examines the social behavior of five sympatric species of desert seed-eating ant... more This field study examines the social behavior of five sympatric species of desert seed-eating ants (Pogonomyrmex barbatus, P. rugosus, P. maricopa, P. desertorum, P. californicus). The species differed significantly in measures of activity rhythms in various colony tasks, use of space around the nest yard, and reaction to disturbance. Species differences were related to the typical size of a colony's outside work force.
Invasive species, where successful, can devastate native communities. We studied the dynamics of ... more Invasive species, where successful, can devastate native communities. We studied the dynamics of the invasive Argentine ant, Linepithema humile, for 7 years in Jasper Ridge, a biological preserve in northern California. We monitored the distributions at the hectare scale of native ant taxa and L. humile in the spring and fall from 1993 to 1999. We also studied the invasion dynamics at a finer resolution by searching for ants in 1-m 2 plots. Our results are similar at both scales. The distributions of several native species are not random with regard to L. humile; the distributions of several epigeic species with similar habitat affinities overlap much less frequently than expected with the distribution of L. humile. We found that season had a significant influence on the distributions of L. humile and several native taxa. Over the 7-year period, L. humile has increased its range size in Jasper Ridge largely at the expense of native taxa, but there is seasonal and yearly variation in this rate of increase. Studies of invasions in progress which sample across seasons and years may help to predict the spread and effects of invasive species.
In the seed-eating ant Pogonomyrmex barbatus, the return of successful foragers stimulates inacti... more In the seed-eating ant Pogonomyrmex barbatus, the return of successful foragers stimulates inactive foragers to leave the nest. The rate at which successful foragers return to the nest depends on food availability; the more food available, the more quickly foragers will find it and bring it back. Field experiments examined how quickly a colony can adjust to a decline in the rate of forager return, and thus to a decline in food availability, by slowing down foraging activity. In response to a brief, 3-to 5-min reduction in the forager return rate, foraging activity usually decreased within 2-3 min and then recovered within 5 min. This indicates that whether an inactive forager leaves the nest on its next trip depends on its very recent experience of the rate of forager return. On some days, colonies responded more to a change in forager return rate. The rapid colony response to fluctuations in forager return rate, enabling colonies to act as risk-averse foragers, may arise from the limited interval over which an ant can track its encounters with returning foragers.
Colonies of the harvester ant, Pogonomyrmex barbatus, adjust the direction and length of foraging... more Colonies of the harvester ant, Pogonomyrmex barbatus, adjust the direction and length of foraging trails in response to the foraging behavior of their conspecific neighbors. In the absence of any interaction with its neighbor, a mature colony expands its foraging range at a rate of 0.85 + 0.15 m per day. Exclusion experiments show that if a colony is prevented from using its foraging trails, the neighbors of that colony will enter its foraging range within 10 days. Exclusion experiments were performed with three age classes of colonies: young (1 year old), intermediate (3-4 years old), and old (5 years old or more). Colonies 3 4 years old are most likely to expand foraging ranges, and to retain newly-gained areas.
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