Matching Items (20)
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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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One hypothesis for the small size of insects relative to vertebrates, and the existence of giant fossil insects, is that atmospheric oxygen levels have constrained body sizes because oxygen delivery would be unable to match the needs of metabolically active tissues in larger insects. This study tested whether oxygen delivery

One hypothesis for the small size of insects relative to vertebrates, and the existence of giant fossil insects, is that atmospheric oxygen levels have constrained body sizes because oxygen delivery would be unable to match the needs of metabolically active tissues in larger insects. This study tested whether oxygen delivery becomes more challenging for larger insects by measuring the oxygen-sensitivity of flight metabolic rates and behavior during hovering for 11 different species of dragonflies that range in mass by an order of magnitude. Animals were flown in 7 different oxygen concentrations ranging from 30% to 2.5% to assess the sensitivity of their behavior and flight metabolic rates to oxygen. I also assessed the oxygen-sensitivity of flight in low-density air (nitrogen replaced with helium), to increase the metabolic demands of hovering flight. Lowered atmosphere densities did induce higher metabolic rates. Flight behaviors but not flight metabolic rates were highly oxygen-sensitive. A significant interaction between oxygen and mass was found for total flight time, with larger dragonflies varying flight time more in response to atmospheric oxygen. This study provides some support for the hypothesis that larger insects are more challenged in oxygen delivery, as predicted by the oxygen limitation hypothesis for insect gigantism in the Paleozoic.
ContributorsHenry, Joanna Randyl (Author) / Harrison, Jon F. (Thesis advisor) / Kaiser, Alexander (Committee member) / Rutowski, Ronald L (Committee member) / Arizona State University (Publisher)
Created2011
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For over a century, researchers have been investigating collective cognition, in which a group of individuals together process information and act as a single cognitive unit. However, I still know little about circumstances under which groups achieve better (or worse) decisions than individuals. My dissertation research directly addressed this longstanding

For over a century, researchers have been investigating collective cognition, in which a group of individuals together process information and act as a single cognitive unit. However, I still know little about circumstances under which groups achieve better (or worse) decisions than individuals. My dissertation research directly addressed this longstanding question, using the house-hunting ant Temnothorax rugatulus as a model system. Here I applied concepts and methods developed in psychology not only to individuals but also to colonies in order to investigate differences of their cognitive abilities. This approach is inspired by the superorganism concept, which sees a tightly integrated insect society as the analog of a single organism. I combined experimental manipulations and models to elucidate the emergent processes of collective cognition. My studies show that groups can achieve superior cognition by sharing the burden of option assessment among members and by integrating information from members using positive feedback. However, the same positive feedback can lock the group into a suboptimal choice in certain circumstances. Although ants are obligately social, my results show that they can be isolated and individually tested on cognitive tasks. In the future, this novel approach will help the field of animal behavior move towards better understanding of collective cognition.
ContributorsSasaki, Takao (Author) / Pratt, Stephen C (Thesis advisor) / Amazeen, Polemnia (Committee member) / Liebig, Jürgen (Committee member) / Janssen, Marco (Committee member) / Fewell, Jennifer (Committee member) / Hölldobler, Bert (Committee member) / Arizona State University (Publisher)
Created2013
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ABSTRACT 1. Aposematic signals advertise prey distastefulness or metabolic unprofitability to potential predators and have evolved independently in many prey groups over the course of evolutionary history as a means of protection from predation. Most aposematic signals investigated to date exhibit highly chromatic patterning; however, relatives in these toxic groups

ABSTRACT 1. Aposematic signals advertise prey distastefulness or metabolic unprofitability to potential predators and have evolved independently in many prey groups over the course of evolutionary history as a means of protection from predation. Most aposematic signals investigated to date exhibit highly chromatic patterning; however, relatives in these toxic groups with patterns of very low chroma have been largely overlooked. 2. We propose that bright displays with low chroma arose in toxic prey species because they were more effective at deterring predation than were their chromatic counterparts, especially when viewed in relatively low light environments such as forest understories. 3. We analyzed the reflectance and radiance of color patches on the wings of 90 tropical butterfly species that belong to groups with documented toxicity that vary in their habitat preferences to test this prediction: Warning signal chroma and perceived chromaticity are expected to be higher and brightness lower in species that fly in open environments when compared to those that fly in forested environments. 4. Analyses of the reflectance and radiance of warning color patches and predator visual modeling support this prediction. Moreover, phylogenetic tests, which correct for statistical non-independence due to phylogenetic relatedness of test species, also support the hypothesis of an evolutionary correlation between perceived chromaticity of aposematic signals and the flight habits of the butterflies that exhibit these signals.
ContributorsDouglas, Jonathan Marion (Author) / Rutowski, Ronald L (Thesis advisor) / Gadau, Juergen (Committee member) / McGraw, Kevin J. (Committee member) / Arizona State University (Publisher)
Created2013
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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
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Colorful ornaments in animals often serve as sexually selected signals of quality. While pigment-based colors are well-studied in these regards, structural colors that result from the interaction of light with photonic nanostructures are comparatively understudied in terms of their consequences in social contexts, their costs of production, and even the

Colorful ornaments in animals often serve as sexually selected signals of quality. While pigment-based colors are well-studied in these regards, structural colors that result from the interaction of light with photonic nanostructures are comparatively understudied in terms of their consequences in social contexts, their costs of production, and even the best way to measure them. Iridescent colors are some of the most brilliant and conspicuous colors in nature, and I studied the measurement, condition-dependence, and signaling role of iridescence in Anna's hummingbirds (Calypte anna). While most animal colors are easily quantified using well-established spectrophotometric techniques, the unique characteristics of iridescent colors present challenges to measurement and opportunities to quantify novel color metrics. I designed and tested an apparatus for careful control and measurement of viewing geometry and highly repeatable measurements. These measurements could be used to accurately characterize individual variation in iridescent Anna's hummingbirds to examine their condition-dependence and signaling role. Next, I examined the literature published to date for evidence of condition-dependence of structural colors in birds. Using meta-analyses, I found that structural colors of all three types - white, ultra-violet/blue, and iridescence - are significantly condition-dependent, meaning that they can convey information about quality to conspecifics. I then investigated whether iridescent colors were condition-dependent in Anna's hummingbirds both in a field correlational study and in an experimental study. Throughout the year, I found that iridescent feathers in both male and female Anna's hummingbirds become less brilliant as they age. Color was not correlated with body condition in any age/sex group. However, iridescent coloration in male Anna's hummingbirds was significantly affected by experimental protein in the diet during feather growth, indicating that iridescent color may signal diet quality. Finally, I examined how iridescent colors were used to mediate social competitions in male and female Anna's hummingbirds. Surprisingly, males that were less colorful won significantly more contests than more colorful males, and colorful males received more aggression. Less colorful males may be attempting to drive away colorful neighbors that may be preferred mates. Female iridescent ornament size and color was highly variable, but did not influence contest outcomes or aggression.
ContributorsMeadows, Melissa (Author) / McGraw, Kevin J. (Thesis advisor) / Rutowski, Ronald L (Committee member) / Sabo, John L (Committee member) / Alcock, John (Committee member) / Deviche, Pierre (Committee member) / Arizona State University (Publisher)
Created2012
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Differences between males and females can evolve through a variety of mechanisms, including sexual and ecological selection. Because coloration is evolutionarily labile, sexually dichromatic species are good models for understanding the evolution of sex differences. While many jumping spiders exhibit diverse and brilliant coloration, they have been notably absent from

Differences between males and females can evolve through a variety of mechanisms, including sexual and ecological selection. Because coloration is evolutionarily labile, sexually dichromatic species are good models for understanding the evolution of sex differences. While many jumping spiders exhibit diverse and brilliant coloration, they have been notably absent from such studies. In the genus Habronattus, females are drab and cryptic while males are brilliantly colored, displaying some of these colors to females during elaborate courtship dances. Here I test multiple hypotheses for the control and function of male color. In the field, I found that Habronattus males indiscriminately court any female they encounter (including other species), so I first examined the role that colors play in species recognition. I manipulated male colors in H. pyrrithrix and found that while they are not required for species recognition, the presence of red facial coloration improves courtship success, but only if males are courting in the sun. Because light environment affects transmission of color signals, the multi-colored displays of males may facilitate communication in variable and unpredictable environments. Because these colors can be costly to produce and maintain, they also have the potential to signal reliable information about male quality to potential female mates. I found that both red facial and green leg coloration is condition dependent in H. pyrrithrix and thus has the potential to signal quality. Yet, surprisingly, this variation in male color does not appear to be important to females. Males of many Habronattus species also exhibit conspicuous markings on the dorsal surface of their abdomens that are not present in females and are oriented away from females during courtship. In the field, I found that these markings are paired with increased leg-waving behavior in a way that resembles the pattern and behavior of wasps; this may provide protection by exploiting the aversions of predators. My data also suggest that different activity levels between the sexes have placed different selection pressures on their dorsal color patterns. Overall, these findings challenge some of the traditional ways that we think about color signaling and provide novel insights into the evolution of animal coloration.
ContributorsTaylor, Lisa Anne (Author) / McGraw, Kevin J. (Thesis advisor) / Clark, David L. (Committee member) / Johnson, James C. (Committee member) / Alcock, John (Committee member) / Rutowski, Ronald L (Committee member) / Arizona State University (Publisher)
Created2012
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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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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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The control, function, and evolution of sleep in animals has received little attention compared to many other fitness-relevant animal behaviors. Though natural selection has largely been thought of as the driving evolutionary force shaping sleep biology, sexual and social selection may also have transformative effects on sleep quantity and quality

The control, function, and evolution of sleep in animals has received little attention compared to many other fitness-relevant animal behaviors. Though natural selection has largely been thought of as the driving evolutionary force shaping sleep biology, sexual and social selection may also have transformative effects on sleep quantity and quality in animals. An overarching hypothesis is that increased levels of investment into inter-sexual choice and intra-sexual competition will reduce sleep. An alternative hypothesis is that sexual ornamentation (e.g. avian plumage coloration and song) may have evolved to communicate sleep health and may therefore be positively related to sleep investment. In this dissertation, I studied how sleep is related to components of sexual and social selection in animals (mostly in birds). I first reviewed the literature for empirical examples of how social and sexual selection drive animal sleep patterns and found support for this relationship in some common types of inter-individual interactions (e.g. mating, intra-sexual competition, parent-offspring interactions, group interactions); I also provided new ideas and hypotheses for future research. I then tested associations between sleep behavior with expression of ornaments (song and plumage coloration), using the house finch (Haemorhous mexicanus) as a model system. For both color and song, I found support for the hypothesis that individuals with exaggerated ornaments slept deeper and longer, suggesting that sleep is a critical resource for ornament elaboration and/or may be communicated by both types of sexual signal. Following this, I tested the phylogenetic association between sleep and social/sexual selection as well as other life-history traits across birds. I found that more territorial bird species sleep less, that polygynous birds sleep more than monogamous and polygynandrous birds, and that birds migrating longer distances sleep less and have less REM sleep. Finally, in the interest of applying basic knowledge about sleep biology to current global problems, I found support for the hypothesis that house finches from city environments have developed resilience to artificial light pollution at night. Altogether, I found that social, sexual, and life-history traits are indeed important and overlooked drivers of sleep behavior from multiple levels of analysis.
ContributorsHutton, Pierce (Author) / McGraw, Kevin J (Thesis advisor) / Rutowski, Ronald L (Committee member) / Deviche, Pierre J (Committee member) / Sweazea, Karen L (Committee member) / Lesku, John A (Committee member) / Arizona State University (Publisher)
Created2021