Matching Items (16)
Description
Like individual organisms, complex social groups are able to maintain predictable trajectories of growth, from initial colony foundation to mature reproductively capable units. They do so while simultaneously responding flexibly to variation in nutrient availability and intake. Leafcutter ant colonies function as tri-trophic systems, in which the ants harvest vegetation to grow a fungus that, in turn, serves as food for the colony. Fungal growth rates and colony worker production are interdependent, regulated by nutritional and behavioral feedbacks. Fungal growth and quality are directly affected by worker foraging decisions, while worker production is, in turn, dependent on the amount and condition of the fungus. In this dissertation, I first characterized the growth relationship between the workers and the fungus of the desert leafcutter ant Acromyrmex versicolor during early stages of colony development, from colony foundation by groups of queens through the beginnings of exponential growth. I found that this relationship undergoes a period of slow growth and instability when workers first emerge, and then becomes allometrically positive. I then evaluated how mass and element ratios of resources collected by the ants are translated into fungus and worker population growth, and refuse, finding that colony digestive efficiency is comparable to digestive efficiencies of other herbivorous insects and ruminants. To test how colonies behaviorally respond to perturbations of the fungus garden, I quantified activity levels and task performance of workers in colonies with either supplemented or diminished fungus gardens, and found that colonies adjusted activity and task allocation in response to the fungus garden size. Finally, to identify possible forms of nutrient limitation, I measured how colony performance was affected by changes in the relative amounts of carbohydrates, protein, and phosphorus available in the resources used to grow the fungus garden. From this experiment, I concluded that colony growth is primarily carbohydrate-limited.
ContributorsClark, Rebecca, 1981- (Author) / Fewell, Jennifer H (Thesis advisor) / Mueller, Ulrich (Committee member) / Liebig, Juergen (Committee member) / Elser, James (Committee member) / Harrison, Jon (Committee member) / Arizona State University (Publisher)
Created2011
Description
Social insect colonies exhibit striking diversity in social organization. Included in this overwhelming variation in structure are differences in colony queen number. The number of queens per colony varies both intra- and interspecifically and has major impacts on the social dynamics of a colony and the fitness of its members. To understand the evolutionary transition from single to multi-queen colonies, I examined a species which exhibits variation both in mode of colony founding and in the queen number of mature colonies. The California harvester ant Pogonomyrmex californicus exhibits both variation in the number of queens that begin a colony (metrosis) and in the number of queens in adult colonies (gyny). Throughout most of its range, colonies begin with one queen (haplometrosis) but in some populations multiple queens cooperate to initiate colonies (pleometrosis). I present results that confirm co-foundresses are unrelated. I also map the geographic occurrence of pleometrotic populations and show that the phenomenon appears to be localized in southern California and Northern Baja California. Additionally, I provide genetic evidence that pleometrosis leads to primary polygyny (polygyny developing from pleometrosis) a phenomenon which has received little attention and is poorly understood. Phylogenetic and haplotype analyses utilizing mitochondrial markers reveal that populations of both behavioral types in California are closely related and have low mitochondrial diversity. Nuclear markers however, indicate strong barriers to gene flow between focal populations. I also show that intrinsic differences in queen behavior lead to the two types of populations observed. Even though populations exhibit strong tendencies on average toward haplo- or pleometrosis, within population variation exists among queens for behaviors relevant to metrosis and gyny. These results are important in understanding the dynamics and evolutionary history of a distinct form of cooperation among unrelated social insects. They also help to understand the dynamics of intraspecific variation and the conflicting forces of local adaptation and gene flow.
ContributorsOverson, Rick P (Author) / Gadau, Jürgen (Thesis advisor) / Fewell, Jennifer H (Committee member) / Hölldobler, Bert (Committee member) / Johnson, Robert A. (Committee member) / Liebig, Jürgen (Committee member) / Arizona State University (Publisher)
Created2011
Description
Division of labor, whereby different group members perform different functions, is a fundamental attribute of sociality. It appears across social systems, from simple cooperative groups to complex eusocial colonies. A core challenge in sociobiology is to explain how patterns of collective organization are generated. Theoretical models propose that division of labor self-organizes, or emerges, from interactions among group members and the environment; division of labor is also predicted to scale positively with group size. I empirically investigated the emergence and scaling of division of labor in evolutionarily incipient groups of sweat bees and in eusocial colonies of harvester ants. To test whether division of labor is an emergent property of group living during early social evolution, I created de novo communal groups of the normally solitary sweat bee Lasioglossum (Ctenonomia) NDA-1. A division of labor repeatedly arose between nest excavation and guarding tasks; results were consistent with hypothesized effects of spatial organization and intrinsic behavioral variability. Moreover, an experimental increase in group size spontaneously promoted higher task specialization and division of labor. Next, I examined the influence of colony size on division of labor in larger, more integrated colonies of the harvester ant Pogonomyrmex californicus. Division of labor scaled positively with colony size in two contexts: during early colony ontogeny, as colonies grew from tens to hundreds of workers, and among same-aged colonies that varied naturally in size. However, manipulation of colony size did not elicit a short-term response, suggesting that the scaling of division of labor in P. californicus colonies is a product of functional integration and underlying developmental processes, rather than a purely emergent epiphenomenon. This research provides novel insights into the organization of work in insect societies, and raises broader questions about the role of size in sociobiology.
ContributorsHolbrook, Carter Tate (Author) / Fewell, Jennifer H (Thesis advisor) / Gadau, Jürgen (Committee member) / Harrison, Jon F. (Committee member) / Hölldobler, Berthold (Committee member) / Johnson, Robert A. (Committee member) / Arizona State University (Publisher)
Created2011
Description
A notable feature of advanced eusocial insect groups is a division of labor within the sterile worker caste. However, the physiological aspects underlying the differentiation of behavioral phenotypes are poorly understood in one of the most successful social taxa, the ants. By starting to understand the foundations on which social behaviors are built, it also becomes possible to better evaluate hypothetical explanations regarding the mechanisms behind the evolution of insect eusociality, such as the argument that the reproductive regulatory infrastructure of solitary ancestors was co-opted and modified to produce distinct castes. This dissertation provides new information regarding the internal factors that could underlie the division of labor observed in both founding queens and workers of Pogonomyrmex californicus ants, and shows that changes in task performance are correlated with differences in reproductive physiology in both castes. In queens and workers, foraging behavior is linked to elevated levels of the reproductively-associated juvenile hormone (JH), and, in workers, this behavioral change is accompanied by depressed levels of ecdysteroid hormones. In both castes, the transition to foraging is also associated with reduced ovarian activity. Further investigation shows that queens remain behaviorally plastic, even after worker emergence, but the association between JH and behavioral bias remains the same, suggesting that this hormone is an important component of behavioral development in these ants. In addition to these reproductive factors, treatment with an inhibitor of the nutrient-sensing pathway Target of Rapamycin (TOR) also causes queens to become biased towards foraging, suggesting an additional sensory component that could play an important role in division of labor. Overall, this work provides novel identification of the possible regulators behind ant division of labor, and suggests how reproductive physiology could play an important role in the evolution and regulation of non-reproductive social behaviors.
ContributorsDolezal, Adam G (Author) / Amdam, Gro V (Thesis advisor) / Brent, Colin S. (Committee member) / Gadau, Juergen (Committee member) / Hoelldobler, Bert (Committee member) / Liebig, Juergen (Committee member) / Arizona State University (Publisher)
Created2012
Description
The repression of reproductive competition and the enforcement of altruism are key components to the success of animal societies. Eusocial insects are defined by having a reproductive division of labor, in which reproduction is relegated to one or few individuals while the rest of the group members maintain the colony and help raise offspring. However, workers have retained the ability to reproduce in most insect societies. In the social Hymenoptera, due to haplodiploidy, workers can lay unfertilized male destined eggs without mating. Potential conflict between workers and queens can arise over male production, and policing behaviors performed by nestmate workers and queens are a means of repressing worker reproduction. This work describes the means and results of the regulation of worker reproduction in the ant species Aphaenogaster cockerelli. Through manipulative laboratory studies on mature colonies, the lack of egg policing and the presence of physical policing by both workers and queens of this species are described. Through chemical analysis and artificial chemical treatments, the role of cuticular hydrocarbons as indicators of fertility status and the informational basis of policing in this species is demonstrated. An additional queen-specific chemical signal in the Dufour's gland is discovered to be used to direct nestmate aggression towards reproductive competitors. Finally, the level of actual worker-derived males in field colonies is measured. Together, these studies demonstrate the effectiveness of policing behaviors on the suppression of worker reproduction in a social insect species, and provide an example of how punishment and the threat of punishment is a powerful force in maintaining cooperative societies.
ContributorsSmith, Adrian A. (Author) / Liebig, Juergen (Thesis advisor) / Hoelldobler, Bert (Thesis advisor) / Gadau, Juergen (Committee member) / Johnson, Robert A. (Committee member) / Pratt, Stephen (Committee member) / Arizona State University (Publisher)
Created2011
Description
For interspecific mutualisms, the behavior of one partner can influence the fitness of the other, especially in the case of symbiotic mutualisms where partners live in close physical association for much of their lives. Behavioral effects on fitness may be particularly important if either species in these long-term relationships displays personality. Animal personality is defined as repeatable individual differences in behavior, and how correlations among these consistent traits are structured is termed behavioral syndromes. Animal personality has been broadly documented across the animal kingdom but is poorly understood in the context of mutualisms. My dissertation focuses on the structure, causes, and consequences of collective personality in Azteca constructor colonies that live in Cecropia trees, one of the most successful and prominent mutualisms of the neotropics. These pioneer plants provide hollow internodes for nesting and nutrient-rich food bodies; in return, the ants provide protection from herbivores and encroaching vines. I first explored the structure of the behavioral syndrome by testing the consistency and correlation of colony-level behavioral traits under natural conditions in the field. Traits were both consistent within colonies and correlated among colonies revealing a behavioral syndrome along a docile-aggressive axis. Host plants of more active, aggressive colonies had less leaf damage, suggesting a link between a colony personality and host plant health. I then studied how aspects of colony sociometry are intertwined with their host plants by assessing the relationship among plant growth, colony growth, colony structure, ant morphology, and colony personality. Colony personality was independent of host plant measures like tree size, age, volume. Finally, I tested how colony personality influenced by soil nutrients by assessing personality in the field and transferring colonies to plants the greenhouse under different soil nutrient treatments. Personality was correlated with soil nutrients in the field but was not influenced by soil nutrient treatment in the greenhouse. This suggests that soil nutrients interact with other factors in the environment to structure personality. This dissertation demonstrates that colony personality is an ecologically relevant phenomenon and an important consideration for mutualism dynamics.
ContributorsMarting, Peter (Author) / Pratt, Stephen C (Thesis advisor) / Wcislo, William T (Committee member) / Hoelldobler, Bert (Committee member) / Fewell, Jennifer H (Committee member) / Gadau, Juergen (Committee member) / Arizona State University (Publisher)
Created2018
Description
An important component of insect social structure is the number of queens that cohabitate in a colony. Queen number is highly variable between and within species. It can begin at colony initiation when often unrelated queens form cooperative social groups, a strategy known as primary polygyny. The non-kin cooperative groups formed by primary polygyny have profound effects on the social dynamics and inclusive fitness benefits within a colony. Despite this, the evolution of non-kin queen cooperation has been relatively overlooked in considerations of the evolution of cooperative sociality. To date, studies examining the costs and benefits of primary polygyny have focused primarily on the advantages of multiple queens during colony founding and early growth, but the impact of their presence extends to colony maturity and reproduction.
In this dissertation, I evaluate the ecological drivers and fitness consequences of non-kin queen cooperation, by comparing the reproduction of mature single-queen versus polygynous harvester ant (Pogonomyrmex californicus) colonies in the field. I captured and quantified the total number and biomass of reproductives across multiple mating seasons, comparing between populations that vary in the proportion of single queen versus polygynous colonies, to assess the fitness outcomes of queen cooperation. Colonies in a mainly polygynous site had lower reproductive investment than those in sites with predominantly single-queen colonies. The site dominated by polygyny had higher colony density and displayed evidence of resource limitation, pressures that may drive the evolution of queen cooperation.
I also used microsatellite markers to examine how polygynous queens share worker and reproductive production with nest-mate queens. The majority of queens fairly contribute to worker production and equally share reproductive output. However, there is a low frequency of queens that under-produce workers and over-produce reproductive offspring. This suggests that cheating by reproducing queens is possible, but uncommon. Competitive pressure from neighboring colonies could reduce the success of colonies that contain cheaters and maintain a low frequency of this phenotype in the population.
In this dissertation, I evaluate the ecological drivers and fitness consequences of non-kin queen cooperation, by comparing the reproduction of mature single-queen versus polygynous harvester ant (Pogonomyrmex californicus) colonies in the field. I captured and quantified the total number and biomass of reproductives across multiple mating seasons, comparing between populations that vary in the proportion of single queen versus polygynous colonies, to assess the fitness outcomes of queen cooperation. Colonies in a mainly polygynous site had lower reproductive investment than those in sites with predominantly single-queen colonies. The site dominated by polygyny had higher colony density and displayed evidence of resource limitation, pressures that may drive the evolution of queen cooperation.
I also used microsatellite markers to examine how polygynous queens share worker and reproductive production with nest-mate queens. The majority of queens fairly contribute to worker production and equally share reproductive output. However, there is a low frequency of queens that under-produce workers and over-produce reproductive offspring. This suggests that cheating by reproducing queens is possible, but uncommon. Competitive pressure from neighboring colonies could reduce the success of colonies that contain cheaters and maintain a low frequency of this phenotype in the population.
ContributorsHaney, Brian R (Author) / Fewell, Jennifer H (Thesis advisor) / Cole, Blaine J. (Committee member) / Gadau, Juergen (Committee member) / Hoelldobler, Bert (Committee member) / Rutowski, Ron L (Committee member) / Arizona State University (Publisher)
Created2017
Description
A key factor in the success of social animals is their organization of work. Mathematical models have been instrumental in unraveling how simple, individual-based rules can generate collective patterns via self-organization. However, existing models offer limited insights into how these patterns are shaped by behavioral differences within groups, in part because they focus on analyzing specific rules rather than general mechanisms that can explain behavior at the individual-level. My work argues for a more principled approach that focuses on the question of how individuals make decisions in costly environments.
In Chapters 2 and 3, I demonstrate how this approach provides novel insights into factors that shape the flexibility and robustness of task organization in harvester ant colonies (Pogonomyrmex barbatus). My results show that the degree to which colonies can respond to work in fluctuating environments depends on how individuals weigh the costs of activity and update their behavior in response to social information. In Chapter 4, I introduce a mathematical framework to study the emergence of collective organization in heterogenous groups. My approach, which is based on the theory of multi-agent systems, focuses on myopic agents whose behavior emerges out of an independent valuation of alternative choices in a given work environment. The product of this dynamic is an equilibrium organization in which agents perform different tasks (or abstain from work) with an analytically defined set of threshold probabilities. The framework is minimally developed, but can be extended to include other factors known to affect task decisions including individual experience and social facilitation. This research contributes a novel approach to developing (and analyzing) models of task organization that can be applied in a broader range of contexts where animals cooperate.
In Chapters 2 and 3, I demonstrate how this approach provides novel insights into factors that shape the flexibility and robustness of task organization in harvester ant colonies (Pogonomyrmex barbatus). My results show that the degree to which colonies can respond to work in fluctuating environments depends on how individuals weigh the costs of activity and update their behavior in response to social information. In Chapter 4, I introduce a mathematical framework to study the emergence of collective organization in heterogenous groups. My approach, which is based on the theory of multi-agent systems, focuses on myopic agents whose behavior emerges out of an independent valuation of alternative choices in a given work environment. The product of this dynamic is an equilibrium organization in which agents perform different tasks (or abstain from work) with an analytically defined set of threshold probabilities. The framework is minimally developed, but can be extended to include other factors known to affect task decisions including individual experience and social facilitation. This research contributes a novel approach to developing (and analyzing) models of task organization that can be applied in a broader range of contexts where animals cooperate.
ContributorsUdiani, Oyita (Author) / Kang, Yun (Thesis advisor) / Fewell, Jennifer H (Thesis advisor) / Janssen, Marcus A (Committee member) / Castillo-Chavez, Carlos (Committee member) / Arizona State University (Publisher)
Created2016
Description
Understanding how and why animals choose what to eat is one of the fundamental goals of nutritional and behavioral biology. This question can be scaled to animals that live in social groups, including eusocial insects. One of the factors that plays an important role in foraging decisions is the prevalence of specific nutrients and their relative balance. This dissertation explores the role of relative nutrient content in the food selection decisions of a species that is eusocial and also agricultural, the desert leafcutter ant Acromyrmex versicolor. A dietary choice assay, in which the relative amount of protein and carbohydrates in the available diets was varied, demonstrated that A. versicolor colonies regulate relative collection of protein and carbohydrates. Tracking the foraging behavior of individual workers revelaed that foragers vary in their relative collection of experimental diets and in their foraging frequency, but that there is no relationship between these key factors of foraging behavior. The high proportion of carbohydrates preferred by lab colonies suggests that they forage to nutritionally support the fungus rather than brood and workers. To test this, the relative amounts of 1) fungus, and 2) brood (larvae) was manipulated and foraging response was measured. Changing the amount of brood had no effect on foraging. Although decreasing the size of fungus gardens did not change relative P:C collection, it produced significant increases in caloric intake, supporting the assertion that the fungus is the main driver of colony nutrient regulation. The nutritional content of naturally harvested forage material collected from field colonies was measured, as was recruitment to experimental diets with varying relative macronutrient content. Field results confirmed a strong colony preference for high carbohydrate diets. They also indicated that this species may, at times, be limited in its ability to collect sufficiently high levels of carbohydrates to meet optimal intake. This dissertation provides important insights about fundamental aspects of leafcutter ant biology and extends our understanding of the role of relative nutrient content in foraging decisions to systems that span multiple trophic levels.
ContributorsSmith, Nathan Edward (Author) / Fewell, Jennifer H (Thesis advisor) / Harrison, Jon F (Committee member) / Pavlic, Ted (Committee member) / Cease, Arianne (Committee member) / Hoelldobler, Bert (Committee member) / Arizona State University (Publisher)
Created2023
Description
Social insect groups, such as bees, termites, and ants, epitomize the emergence of group-level behaviors from the aggregated actions and interactions of individuals. Ants have the unique advantage that whole colonies can be observed in artificial, laboratory nests, and each individual's behavior can be continuously tracked using imaging software. In this dissertation, I study two group behaviors: (1) the spread of alarm signals from three agitated ants to a group of 61 quiescent nestmates, and (2) the reduction in per-capita energy use as colonies scale in size from tens of ants to thousands. For my first experiment, I track the motion of Pogonomyrmex californicus ants using an overhead camera, and I analyze how propagation of an initial alarm stimulus affects their walking speeds. I then build an agent-based model that simulates two-dimensional ant motion and the spread of the alarmed state. I find that implementing a simple set of rules for motion and alarm signal transmission reproduces the empirically observed speed dynamics. For the second experiment, I simulate social insect colony workers that collectively complete a set of tasks. By assuming that task switching is energetically costly, my model recovers a metabolic rate scaling pattern, known as hypometric metabolic scaling. This relationship, which predicts an organism's metabolic rate from its mass, is observed across a diverse set of social insect groups and animal species. The results suggest an explicit link between the degree of workers' task specialization and whole-colony energy use.
ContributorsLin, Michael Robert (Author) / Milner, Fabio A (Thesis advisor, Committee member) / Fewell, Jennifer H (Thesis advisor, Committee member) / Lampert, Adam (Committee member) / Arizona State University (Publisher)
Created2021