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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

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
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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.

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
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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

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
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Description
At the heart of every eusocial insect colony is a reproductive division of labor. This division can emerge through dominance interactions at the adult stage or through the production of distinct queen and worker castes at the larval stage. In both cases, this division depends on plasticity within an individual

At the heart of every eusocial insect colony is a reproductive division of labor. This division can emerge through dominance interactions at the adult stage or through the production of distinct queen and worker castes at the larval stage. In both cases, this division depends on plasticity within an individual to develop reproductive characteristics or serve as a worker. In order to gain insight into the evolution of reproductive plasticity in the social insects, I investigated caste determination and dominance in the ant Harpegnathos saltator, a species that retains a number of ancestral characteristics. Treatment of worker larvae with a juvenile hormone (JH) analog induced late-instar larvae to develop as queens. At the colony level, workers must have a mechanism to regulate larval development to prevent queens from developing out of season. I identified a new behavior in H. saltator where workers bite larvae to inhibit queen determination. Workers could identify larval caste based on a chemical signal specific to queen-destined larvae, and the production of this signal was directly linked to increased JH levels. This association provides a connection between the physiological factors that induce queen development and the production of a caste-specific larval signal. In addition to caste determination at the larval stage, adult workers of H. saltator compete to establish a reproductive hierarchy. Unlike other social insects, dominance in H. saltator was not related to differences in JH or ecdysteroid levels. Instead, changes in brain levels of biogenic amines, particularly dopamine, were correlated with dominance and reproductive status. Receptor genes for dopamine were expressed in both the brain and ovaries of H. saltator, and this suggests that dopamine may coordinate changes in behavior at the neurological level with ovarian status. Together, these studies build on our understanding of reproductive plasticity in social insects and provide insight into the evolution of a reproductive division of labor.
ContributorsPenick, Clint A (Author) / Liebig, Jürgen (Thesis advisor) / Brent, Colin (Committee member) / Gadau, Jürgen (Committee member) / Hölldobler, Bert (Committee member) / Rutowski, Ron (Committee member) / Arizona State University (Publisher)
Created2012
Description
Evolution is a powerful process that acts on features as organisms adapt to fill a variety of niches. It is visible in the emergence of the beak in the fossil record, through a number of small changes over time. To explain and convey these changes to a general audience, I

Evolution is a powerful process that acts on features as organisms adapt to fill a variety of niches. It is visible in the emergence of the beak in the fossil record, through a number of small changes over time. To explain and convey these changes to a general audience, I produced an art book combining my review of bird beak evolution with art. The intent was to present evolution in an informative, visual, and engaging manner that a general audience would be able to understand.
ContributorsWalls, Sarah Camille (Author) / Collins, James (Thesis director) / Hodgen, Heidi (Committee member) / School of Life Sciences (Contributor, Contributor) / School of Art (Contributor) / Barrett, The Honors College (Contributor)
Created2020-05
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Description
Dominance behavior can regulate a division of labor in a group, such as that between reproductive and non-reproductive individuals. Manipulations of insect societies in a controlled environment can reveal how dominance behavior is regulated. Here, I examined how morphological caste, fecundity, group size, and age influence the expression of

Dominance behavior can regulate a division of labor in a group, such as that between reproductive and non-reproductive individuals. Manipulations of insect societies in a controlled environment can reveal how dominance behavior is regulated. Here, I examined how morphological caste, fecundity, group size, and age influence the expression of dominance behavior using the ponerine ant Harpegnathos saltator. All H. saltator females have the ability to reproduce. Only those with a queen morphology that enables dispersal, however, show putative sex pheromones. In contrast, those with a worker morphology normally express dominance behavior. To evaluate how worker-like dominance behavior and associated traits could be expressed in queens, I removed the wings from alate gynes, those with a queen morphology who had not yet mated or left the nest, making them dealate. Compared to gynes with attached wings, dealates frequently performed dominance behavior. In addition, only the dealates demonstrated worker-like ovarian activity in the presence of reproductive individuals, whereas gynes with wings produced sex pheromones exclusively. Therefore, the attachment of wings determines a gyne’s expression of worker-like dominance behavior and physiology. When the queen dies, workers establish a reproductive hierarchy among themselves by performing a combination of dominance behaviors. To understand how reproductive status depends on these interactions as well as a worker’s age, I measured the frequency of dominance behaviors in groups of different size composed of young and old workers. The number of workers who expressed dominance scaled with the size of the group, but younger ones were more likely to express dominance behavior and eventually become reproductive. Therefore, the predisposition of age integrates with a self-organized process to form this reproductive hierarchy. A social insect’s fecundity and fertility signal depends on social context because fecundity increases with colony size. To evaluate how a socially dependent signal regulates dominance behavior, I manipulated a reproductive worker’s social context. Reproductive workers with reduced fecundity and a less prominent fertility signal expressed more dominance behavior than those with a stronger fertility signal and higher fecundity. Therefore, dominance behavior reinforces rank to compensate for a weak signal, indicating how social context can feed back to influence the maintenance of dominance. Mechanisms that regulate H. saltator’s reproductive hierarchy can inform how the reproductive division of labor is regulated in other groups of animals.
ContributorsPyenson, Benjamin (Author) / Liebig, Jürgen (Thesis advisor) / Hölldobler, Bert (Committee member) / Fewell, Jennifer (Committee member) / Pratt, Stephen (Committee member) / Kang, Yun (Committee member) / Arizona State University (Publisher)
Created2022
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Description
Evolution is a key feature of undergraduate biology education: the AmericanAssociation for the Advancement of Science (AAAS) has identified evolution as one of the five core concepts of biology, and it is relevant to a wide array of biology-related careers. If biology instructors want students to use evolution to address scientific challenges post-graduation,

Evolution is a key feature of undergraduate biology education: the AmericanAssociation for the Advancement of Science (AAAS) has identified evolution as one of the five core concepts of biology, and it is relevant to a wide array of biology-related careers. If biology instructors want students to use evolution to address scientific challenges post-graduation, students need to be able to apply evolutionary principles to real-life situations, and accept that the theory of evolution is the best scientific explanation for the unity and diversity of life on Earth. In order to help students progress on both fronts, biology education researchers need surveys that measure evolution acceptance and assessments that measure students’ ability to apply evolutionary concepts. This dissertation improves the measurement of student understanding and acceptance of evolution by (1) developing a novel Evolutionary Medicine Assessment that measures students’ ability to apply the core principles of Evolutionary Medicine to a variety of health-related scenarios, (2) reevaluating existing measures of student evolution acceptance by using student interviews to assess response process validity, and (3) correcting the validity issues identified on the most widely-used measure of evolution acceptance - the Measure of Acceptance of the Theory of Evolution (MATE) - by developing and validating a revised version of this survey: the MATE 2.0.
ContributorsMisheva, Anastasia Taya (Author) / Brownell, Sara (Thesis advisor) / Barnes, Elizabeth (Committee member) / Collins, James (Committee member) / Cooper, Katelyn (Committee member) / Sterner, Beckett (Committee member) / Arizona State University (Publisher)
Created2023
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Description
Eusocial insect colonies have often been imagined as “superorganisms” exhibiting tight homeostasis at the colony level. However, colonies lack the tight spatial and organizational integration that many multicellular, unitary organisms exhibit. Precise regulation requires rapid feedback, which is often not possible when nestmates are distributed across space, making decisions asynchronously.

Eusocial insect colonies have often been imagined as “superorganisms” exhibiting tight homeostasis at the colony level. However, colonies lack the tight spatial and organizational integration that many multicellular, unitary organisms exhibit. Precise regulation requires rapid feedback, which is often not possible when nestmates are distributed across space, making decisions asynchronously. Thus, one should expect poorer regulation in superorganisms than unitary organisms.Here, I investigate aspects of regulation in collective foraging behaviors that involve both slow and rapid feedback processes. In Chapter 2, I examine a tightly coupled system with near-instantaneous signaling: teams of weaver ants cooperating to transport massive prey items back to their nest. I discover that over an extreme range of scenarios—even up vertical surfaces—the efficiency per transporter remains constant. My results suggest that weaver ant colonies are maximizing their total intake rate by regulating the allocation of transporters among loads. This is an exception that “proves the rule;” the ant teams are recapitulating the physical integration of unitary organisms. Next, I focus on a process with greater informational constraints, with loose temporal and spatial integration. In Chapter 3, I measure the ability of solitarily foraging Ectatomma ruidum colonies to balance their collection of protein and carbohydrates given different nutritional environments. Previous research has found that ant species can precisely collect a near-constant ratio between these two macronutrients, but I discover these studies were using flawed statistical approaches. By developing a quantitative measure of regulatory effect size, I show that colonies of E. ruidum are relatively insensitive to small differences in food source nutritional content, contrary to previously published claims. In Chapter 4, I design an automated, micro-RFID ant tracking system to investigate how the foraging behavior of individuals integrates into colony-level nutrient collection. I discover that spatial fidelity to food resources, not individual specialization on particular nutrient types, best predicts individual forager behavior. These findings contradict previously published experiments that did not use rigorous quantitative measures of specialization and confounded the effects of task type and resource location.
ContributorsBurchill, Andrew Taylor (Author) / Pavlic, Theodore P (Thesis advisor) / Pratt, Stephen C (Thesis advisor) / Hölldobler, Bert (Committee member) / Cease, Arianne (Committee member) / Berman, Spring (Committee member) / Arizona State University (Publisher)
Created2022
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Description
Insecticide resistance is a continuing issue that negatively affects both public health and agriculture and allows vector-borne diseases to spread throughout the globe. To improve resistance management strategies (RMS), robust susceptibility bioassays need to be performed in order to fill the gap of the relationship between resistant and susceptible genotype

Insecticide resistance is a continuing issue that negatively affects both public health and agriculture and allows vector-borne diseases to spread throughout the globe. To improve resistance management strategies (RMS), robust susceptibility bioassays need to be performed in order to fill the gap of the relationship between resistant and susceptible genotype and phenotype, and a deeper knowledge of how bioassay data relates to vector control success or failure is imperative. A bioassay method that is infrequently used but yields robust results is the topical application bioassay, where the insect is directly treated with a constant volume and concentration of an insecticide via a syringe. To bring more attention to this method, my colleagues and I published a paper in the Journal of Visualized Experiments where the optimized protocol of the topical application bioassay for mosquitoes and fruit flies is described, and the strengths and limitations to the method are explained. To further investigate insecticide susceptibility tests, I set up my individual project where I used Aedes aegypti mosquitoes to compare the topical application bioassay to the commonly used Centers for Disease Control and Prevention (CDC) bottle bioassay and World Health Organization (WHO) tube test. The objective of this study was to test which method exhibited the most variability in mortality results, which would guide the choice of assay to determine the link between resistant and susceptible genotype and phenotype. The results showed that the topical application method did indeed exhibit the least amount of variation, followed by the CDC bottle bioassay (WHO data is currently being collected). This suggests that the topical application bioassay could be a useful tool in insecticide resistance surveillance studies, and, depending on the goal, may be better than the CDC and WHO tube tests for assessing resistance levels at a given site. This study challenges the value of the widely used CDC and WHO assays and provides a discussion on the importance of technical and practical resistance assays. This will help vector control specialists to collect accurate surveillance data that will inform effective RMS.
ContributorsAlthoff, Rachel (Author) / Huijben, Silvie (Thesis advisor) / Harris, Robin (Committee member) / Collins, James (Committee member) / Arizona State University (Publisher)
Created2022
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Description
Guided by Tinto’s Theory of College Student Departure, I conducted a set of five studies to identify factors that influence students’ social integration in college science active learning classes. These studies were conducted in large-enrollment college science courses and some were specifically conducted in undergraduate active learning biology courses.

Guided by Tinto’s Theory of College Student Departure, I conducted a set of five studies to identify factors that influence students’ social integration in college science active learning classes. These studies were conducted in large-enrollment college science courses and some were specifically conducted in undergraduate active learning biology courses. Using qualitative and quantitative methodologies, I identified how students’ identities, such as their gender and LGBTQIA identity, and students’ perceptions of their own intelligence influence their experience in active learning science classes and consequently their social integration in college. I also determined factors of active learning classrooms and instructor behaviors that can affect whether students experience positive or negative social integration in the context of active learning. I found that students’ hidden identities, such as the LGBTQIA identity, are more relevant in active learning classes where students work together and that the increased relevance of one’s identity can have a positive and negative impact on their social integration. I also found that students’ identities can predict their academic self-concept, or their perception of their intelligence as it compares to others’ intelligence in biology, which in turn predicts their participation in small group-discussion. While many students express a fear of negative evaluation, or dread being evaluated negatively by others when speaking out in active learning classes, I identified that how instructors structure group work can cause students to feel more or less integrated into the college science classroom. Lastly, I identified tools that instructors can use, such as name tents and humor, which can positive affect students’ social integration into the college science classroom. In sum, I highlight inequities in students’ experiences in active learning science classrooms and the mechanisms that underlie some of these inequities. I hope this work can be used to create more inclusive undergraduate active learning science courses.
ContributorsCooper, Katelyn M (Author) / Brownell, Sara E (Thesis advisor) / Stout, Valerie (Committee member) / Collins, James (Committee member) / Orchinik, Miles (Committee member) / Zheng, Yi (Committee member) / Arizona State University (Publisher)
Created2018