The cooperative construction of shared dwellings is a phylogenetically-widespread evolutionary puzzle. Shared shelters are common goods – all individuals in the shelter benefit, at the expense of those individuals that contribute to the construction. The evolution of cooperation requires existing variation for selection to act upon and genetic benefits to cooperators, through inclusive fitness or direct rewards. This study focuses on two genera of mice, Mus and Peromyscus, to examine shared construction and social monogamy as potential transitions to more sophisticated forms of sociality, such as cooperative breeding. The mound-building mouse (Mus spicilegus) is named for the large mounds that groups of mice build and beneath which they overwinter. Variation in mtDNA and 14 microsatellites show limited genetic structure across the geographic range of M. spicilegus. Mice from the same mound are more genetically related than mice from different mounds, and males and females associated with a mound are equally likely to be relatives. However in spring, when breeding begins, male kin are more likely to share a territory than are female kin. One possible interpretation is that males associate with kin to minimize the costs of being cuckolded, as this study finds evidence of multiple paternity in every litter genotyped. By increasing the chances of the cuckold being a brother, a male still gains inclusive fitness benefits from paternal care to extra-pair offspring in this socially monogamous species. Behavioral experiments show that another socially monogamous mouse species, the oldfield mouse (Peromyscus polionotus), can coordinate construction with unfamiliar, unrelated conspecifics. In contrast, two other closely related Peromyscus species do not dig longer burrows in pairs than they would have as individuals. Male-female P. polionotus pairs tend to dig longer burrows than pairs of the same sex, but males within opposite sex pairs do most of the digging, particularly when paired with an unfamiliar female. Male burrowing could be the product of female choice in this monogamous species. In M. spicilegus and P. polionotus, shared parental care and construction shed light on the evolution of cooperation and conflict.

The spread of an agricultural lifestyle played a crucial role in the development of social complexity and in defining trajectories of human history. This dissertation presents the results of research into how agricultural strategies were modified during the spread of agriculture into Southwest China. By incorporating advances from the fields of plant biology and ecological niche modeling into archaeological research, this dissertation addresses how humans adapted their agricultural strategies or invented appropriate technologies to deal with the challenges presented by the myriad of ecological niches in southwest China. This dissertation uses ecological niche modeling to examine the options and constraints associated with practicing different types of agriculture in the specific ecological niches of southwest China. The predictions made by these models are then tested against archaeobotanical data from a series of sites from across the region. This approach allows one to understand how the spread of agriculture took place in its particular social and economic contexts. / Anthropology

Increasingly, individual variation is being recognized as an important influence on behavioral evolution. Sources of variation are therefore an important target for research into the development, evolution, and function of behavior. By providing information about the timescale on which individuals are responsive to their environment, patterns of within-individual variation can shed light on function of behavioral variation. Here, I wanted to understand the function of behavioral variation and the genetic and environmental sources of variation in behavior. First, I test the hypotheses that variation in begging signals nestling hunger, need, or quality. Hunger is a short-term response to food deprivation, while need and quality give long-term information about fitness benefits of gaining more food and fitness potential, respectively. Second, I test the hypotheses that variation in begging is due to genetic, permanent environment, common environmental, and maternal effects. I test these hypotheses in the begging behavior of western bluebirds (Sialia mexicana), making repeated measurements across the nestling period. I show that begging behavior is consistent across the nestling period, and that nestling begging intensity increases with food deprivation. Nestlings fed during a given parental visit beg at higher intensity than nestmates, and on average wait longer since their last meal compared to individuals who were not fed in the same visit. These results support the hypothesis that variation in nestling begging signals hunger. I also show that responsiveness to food deprivation is negatively related to condition, but this effect is not consistent across the nestling period. Finally, variation in begging is produced by a common environmental effect that is correlated through time, suggesting that begging is strongly influenced by the nest environment. Together, these results indicate that variation in begging signals short-term changes in hunger and that environmental effects dominate the production of variation in begging.

The behavioral ecology of the American Wigeon (Anas americana) was studied over its annual cycle in western Canada between 1974 and 1978. The objective of this work was to examine the cross-seasonal interactions of the biology and behavior of the wigeon as they underlie the evolution of its mating system. The wigeon was more herbivorous than any other dabbling duck studied so far and spent a considerable amount of time feeding in upland habitats. Vegetation was important in the diet year round but particularly from late summer through the winter. Plant seeds were a rich source of protein and carbohydrate early in the spring. During breeding stages animal foods were selected by wigeons, particularly females. Wigeons usually did not feed on small crustaceans and gastropods but selected larger aquatic insect foods. Relative to other Anas soecies, the wigeon has a goose-like bill whose structure facilitates feeding on upland and aquatic vegetation. With the evolution of this specialization, efficient surface-straining of small foods has been sacrificed. To increase food digestibility, wigeons altered gizzard grit composition and organ sizes. Through these mechanisms I have demonstrated how wigeons have been able to increase the efficiency of digestion.

The behavioral ecology of the American Wigeon (Anas americana) was studied over its annual cycle in western Canada between 1974 and 1978. The objective of this work was to examine the cross-seasonal interactions of the biology and behavior of the wigeon as they underlie the evolution of its mating system. The wigeon was more herbivorous than any other dabbling duck studied so far and spent a considerable amount of time feeding in upland habitats. Vegetation was important in the diet year round but particularly from late summer through the winter. Plant seeds were a rich source of protein and carbohydrate early in the spring. During breeding stages animal foods were selected by wigeons, particularly females. Wigeons usually did not feed on small crustaceans and gastropods but selected larger aquatic insect foods. Relative to other Anas soecies, the wigeon has a goose-like bill whose structure facilitates feeding on upland and aquatic vegetation. With the evolution of this specialization, efficient surface-straining of small foods has been sacrificed. To increase food digestibility, wigeons altered gizzard grit composition and organ sizes. Through these mechanisms I have demonstrated how wigeons have been able to increase the efficiency of digestion.

No contexto da herbivoria, a interação inseto-planta é um sistema dinâmico. As plantas apresentam características que influenciam a preferência e o desempenho dos herbívoros, como por exemplo, a variação da dureza e espessura do limbo foliar, de forma dependente à espécie e à idade da estrutura considerada. Por sua vez, os insetos podem responder às barreiras das plantas por mecanismos fisiológicos e comportamentais. Há registro de nove espécies de passifloráceas utilizadas por Heliconius erato phyllis (Fabricius, 1775) para o Rio Grande do Sul. As larvas deste herbívoro, entretanto, alimentam-se preferencialmente de Passiflora misera Humbold, Bonplant & Kunth, que confere maior performance, apesar de apresentar menor quantidade de nutrientes que outras hospedeiras utilizadas (ex. Passiflora suberosa Linnaeus). Desta forma, o desempenho associado ao consumo de uma passiflorácea possivelmente relaciona-se, não só ao conteúdo nutricional desta, mas também aos mecanismos morfológicos e comportamentais envolvidos na alimentação. Neste trabalho, verificamos se a dificuldade em acessar o alimento, imposta pela dureza e espessura das folhas das hospedeiras e pelas limitações estruturais das mandíbulas de H. erato phyllis, reflete ou não alterações na utilização da planta, analisando os danos causados a estas e o comportamento das larvas deste herbívoro. Para tanto P. misera, P. suberosa, Passiflora caerulea Linnaeus, Passiflora edulis Sims e Passiflora alata Dryander foram cultivadas, sendo folhas jovens e velhas dessas passifloráceas caracterizadas quanto à dureza e espessura, tanto do limbo como da nervura central. A freqüência dos danos causados à P. misera, P. suberosa, P. caerulea e P. edulis foi avaliada, em relação à idade da folha e ao longo da ontogênese larval. Quantificou-se, também, o índice de microdureza de Vickers das mandíbulas para larvas criadas em P. misera e P. suberosa. Adicionalmente, o tempo relativo aos comportamentos (repousando, alimentando, deslocando, provando e cortando nervura) foi quantificado nas cinco passifloráceas, tanto em folhas jovens quanto nas velhas. Os dados obtidos evidenciaram a existência de variação expressiva em relação à idade da folha, quanto à dureza e espessura para as espécies de passifloráceas utilizadas pelas larvas de H. erato phyllis. O tipo e a freqüência do dano (corte do limbo, raspagem e corte da nervura central) foram influenciados pela variação desses parâmetros físicos, sendo que a raspagem do limbo ocorreu quando larvas de primeiro ínstar foram criadas em folhas velhas. Houve influência ontogenética no incremento de dureza das mandíbulas, porém não foram detectados indícios de que a planta hospedeira influencie no aumento deste parâmetro nessas estruturas. As larvas empregaram grande parte do tempo em repouso, independente da espécie de hospedeira e, na maioria dos casos, dedicaram mais tempo à alimentação em P. misera. Larvas observadas em P. alata dedicaram mais tempo ao repouso e menos tempo à alimentação realizando de uma a duas refeições com pequena duração, a cada seis horas. Esta discrepância no tempo destinado a todos os comportamentos das larvas em P. alata se deve, provavelmente, a fatores químicos inerentes a esta passiflorácea. Lagartas de primeiro ínstar em folhas velhas de P. suberosa e P. caerulea despenderam maior tempo no deslocamento, demonstrando que a procura por um sítio de alimentação favorável é de fundamental importância para esta fase do desenvolvimento, evidenciando assim que a dureza foliar constitui fator limitante para ínstares iniciais, neste heliconíneo. / The plant-insect interaction is a dynamic system. Variation in plant morphological traits such as hardness and thickness of the leaves can influence preference and performance of herbivores, depending upon the species and age of the plant structure taking into. On the other hand, insects can respond to these barriers through physiological and behavioral mechanisms. Nine species of passion vine are used by Heliconius erato phyllis (Fabricius, 1775) in Rio Grande do Sul State. However, their larvae feed preferentially on Passiflora misera Humbold, Bonplant & Kunth, which provide greater performance, despite having a lower amount of nutrients when compared to other host plants (e.g. Passiflora suberosa Linnaeus). Thus, the performance related to consumption of a particular passion vine possibly relates not only to their nutritional value, but also to morphological and behavioral mechanisms involved on feeding by these butterfly larvae. In this study, we investigate whether the difficulty in accessing food, imposed by the hardness and thickness of host leaves when in association to structural limitations on the mandibles of H. erato phyllis larvae, lead in turn to changes in host plant use. We examined the plant hosts damages and also larval behavior of this herbivore on five cultivated passion vine: P. misera, P. suberosa, Passiflora caerulea Linnaeus, Passiflora edulis Sims and Passiflora alata Dryander. Young and old leaves of these passion vine species were characterized in terms of hardness and thickness for the blade and the midrib. The frequency of differences on damage caused to P. misera, P. suberosa, P. caerulea and P. edulis was evaluated in relation to leaf age throughout larval ontogeny. Vickers hardness index was quantified for mandibles dissected from larvae reared on P. misera and P. suberosa. Additionally, the time spent in different behaviors (resting, feeding, walking, tasting and vein cutting) was quantified for the five passion vine, while feeding on both young and old leaves. The data showed the existence of expressive variation in hardness and thickness according to leaf age for the passion vine species used by H. erato phyllis larvae. The type (cutting blade, scraping and vein cutting) and frequency of damages were influenced by the variation of such leaf parameters. Scraping occurred when first instar were reared on old leaves. There was an ontogenetic influence on mandibular hardness, which increases with larval age. However, there was no evidence that the host plant influences the hardness in these structures throughout ontogeny. Independently of the host tanken into account, larvae employed most of the time resting. In several cases, they devoted more time feeding in P. misera. In P. alata, they spent more time resting and less time feeding, having up to only two meals of short duration each, at every six hours. This discrepancy in time is probably due to the presence of deterrent chemicals present in this passion vine. In old leaves of P. suberosa and P. caerulea, first instar larvae spent more time moving, thus showing that the search for an suitable food site is extremely important for this stage of development. Thus, we demonstrate that leaf hardness is a limiting factor for the early larval instars in this heliconian butterfly.

No contexto da herbivoria, a interação inseto-planta é um sistema dinâmico. As plantas apresentam características que influenciam a preferência e o desempenho dos herbívoros, como por exemplo, a variação da dureza e espessura do limbo foliar, de forma dependente à espécie e à idade da estrutura considerada. Por sua vez, os insetos podem responder às barreiras das plantas por mecanismos fisiológicos e comportamentais. Há registro de nove espécies de passifloráceas utilizadas por Heliconius erato phyllis (Fabricius, 1775) para o Rio Grande do Sul. As larvas deste herbívoro, entretanto, alimentam-se preferencialmente de Passiflora misera Humbold, Bonplant & Kunth, que confere maior performance, apesar de apresentar menor quantidade de nutrientes que outras hospedeiras utilizadas (ex. Passiflora suberosa Linnaeus). Desta forma, o desempenho associado ao consumo de uma passiflorácea possivelmente relaciona-se, não só ao conteúdo nutricional desta, mas também aos mecanismos morfológicos e comportamentais envolvidos na alimentação. Neste trabalho, verificamos se a dificuldade em acessar o alimento, imposta pela dureza e espessura das folhas das hospedeiras e pelas limitações estruturais das mandíbulas de H. erato phyllis, reflete ou não alterações na utilização da planta, analisando os danos causados a estas e o comportamento das larvas deste herbívoro. Para tanto P. misera, P. suberosa, Passiflora caerulea Linnaeus, Passiflora edulis Sims e Passiflora alata Dryander foram cultivadas, sendo folhas jovens e velhas dessas passifloráceas caracterizadas quanto à dureza e espessura, tanto do limbo como da nervura central. A freqüência dos danos causados à P. misera, P. suberosa, P. caerulea e P. edulis foi avaliada, em relação à idade da folha e ao longo da ontogênese larval. Quantificou-se, também, o índice de microdureza de Vickers das mandíbulas para larvas criadas em P. misera e P. suberosa. Adicionalmente, o tempo relativo aos comportamentos (repousando, alimentando, deslocando, provando e cortando nervura) foi quantificado nas cinco passifloráceas, tanto em folhas jovens quanto nas velhas. Os dados obtidos evidenciaram a existência de variação expressiva em relação à idade da folha, quanto à dureza e espessura para as espécies de passifloráceas utilizadas pelas larvas de H. erato phyllis. O tipo e a freqüência do dano (corte do limbo, raspagem e corte da nervura central) foram influenciados pela variação desses parâmetros físicos, sendo que a raspagem do limbo ocorreu quando larvas de primeiro ínstar foram criadas em folhas velhas. Houve influência ontogenética no incremento de dureza das mandíbulas, porém não foram detectados indícios de que a planta hospedeira influencie no aumento deste parâmetro nessas estruturas. As larvas empregaram grande parte do tempo em repouso, independente da espécie de hospedeira e, na maioria dos casos, dedicaram mais tempo à alimentação em P. misera. Larvas observadas em P. alata dedicaram mais tempo ao repouso e menos tempo à alimentação realizando de uma a duas refeições com pequena duração, a cada seis horas. Esta discrepância no tempo destinado a todos os comportamentos das larvas em P. alata se deve, provavelmente, a fatores químicos inerentes a esta passiflorácea. Lagartas de primeiro ínstar em folhas velhas de P. suberosa e P. caerulea despenderam maior tempo no deslocamento, demonstrando que a procura por um sítio de alimentação favorável é de fundamental importância para esta fase do desenvolvimento, evidenciando assim que a dureza foliar constitui fator limitante para ínstares iniciais, neste heliconíneo. / The plant-insect interaction is a dynamic system. Variation in plant morphological traits such as hardness and thickness of the leaves can influence preference and performance of herbivores, depending upon the species and age of the plant structure taking into. On the other hand, insects can respond to these barriers through physiological and behavioral mechanisms. Nine species of passion vine are used by Heliconius erato phyllis (Fabricius, 1775) in Rio Grande do Sul State. However, their larvae feed preferentially on Passiflora misera Humbold, Bonplant & Kunth, which provide greater performance, despite having a lower amount of nutrients when compared to other host plants (e.g. Passiflora suberosa Linnaeus). Thus, the performance related to consumption of a particular passion vine possibly relates not only to their nutritional value, but also to morphological and behavioral mechanisms involved on feeding by these butterfly larvae. In this study, we investigate whether the difficulty in accessing food, imposed by the hardness and thickness of host leaves when in association to structural limitations on the mandibles of H. erato phyllis larvae, lead in turn to changes in host plant use. We examined the plant hosts damages and also larval behavior of this herbivore on five cultivated passion vine: P. misera, P. suberosa, Passiflora caerulea Linnaeus, Passiflora edulis Sims and Passiflora alata Dryander. Young and old leaves of these passion vine species were characterized in terms of hardness and thickness for the blade and the midrib. The frequency of differences on damage caused to P. misera, P. suberosa, P. caerulea and P. edulis was evaluated in relation to leaf age throughout larval ontogeny. Vickers hardness index was quantified for mandibles dissected from larvae reared on P. misera and P. suberosa. Additionally, the time spent in different behaviors (resting, feeding, walking, tasting and vein cutting) was quantified for the five passion vine, while feeding on both young and old leaves. The data showed the existence of expressive variation in hardness and thickness according to leaf age for the passion vine species used by H. erato phyllis larvae. The type (cutting blade, scraping and vein cutting) and frequency of damages were influenced by the variation of such leaf parameters. Scraping occurred when first instar were reared on old leaves. There was an ontogenetic influence on mandibular hardness, which increases with larval age. However, there was no evidence that the host plant influences the hardness in these structures throughout ontogeny. Independently of the host tanken into account, larvae employed most of the time resting. In several cases, they devoted more time feeding in P. misera. In P. alata, they spent more time resting and less time feeding, having up to only two meals of short duration each, at every six hours. This discrepancy in time is probably due to the presence of deterrent chemicals present in this passion vine. In old leaves of P. suberosa and P. caerulea, first instar larvae spent more time moving, thus showing that the search for an suitable food site is extremely important for this stage of development. Thus, we demonstrate that leaf hardness is a limiting factor for the early larval instars in this heliconian butterfly.

abstract: Warning coloration deters predators from attacking prey that are defended, usually by being distasteful, toxic, or otherwise costly for predators to pursue and consume. Predators may have an innate response to warning colors or learn to associate them with a defense through trial and error. In general, predators should select for warning signals that are easy to learn and recognize. Previous research demonstrates long-wavelength colors (e.g. red and yellow) are effective because they are readily detected and learned. However, a number of defended animals display short-wavelength coloration (e.g. blue and violet), such as the pipevine swallowtail butterfly (Battus philenor). The role of blue coloration in warning signals had not previously been explicitly tested. My research showed in laboratory experiments that curve-billed thrashers (Toxostoma curvirostre) and Gambel's quail (Callipepla gambelii) can learn and recognize the iridescent blue of B. philenor as a warning signal and that it is innately avoided. I tested the attack rates of these colors in the field and blue was not as effective as orange. I concluded that blue colors may function as warning signals, but the effectiveness is likely dependent on the context and predator. Blue colors are often iridescent in nature and the effect of iridescence on warning signal function was unknown. I reared B. philenor larvae under varied food deprivation treatments. Iridescent colors did not have more variation than pigment-based colors under these conditions; variation which could affect predator learning. Learning could also be affected by changes in appearance, as iridescent colors change in both hue and brightness as the angle of illuminating light and viewer change in relation to the color surface. Iridescent colors can also be much brighter than pigment-based colors and iridescent animals can statically display different hues. I tested these potential effects on warning signal learning by domestic chickens (Gallus gallus domesticus) and found that variation due to the directionality of iridescence and a brighter warning signal did not influence learning. However, blue-violet was learned more readily than blue-green. These experiments revealed that the directionality of iridescent coloration does not likely negatively affect its potential effectiveness as a warning signal. / Dissertation/Thesis / Doctoral Dissertation Biology 2015

abstract: Despite nearly five decades of archaeological research in the Romanian Carpathian basin and adjacent areas, how human foragers organized their stone artifact technologies under varying environmental conditions remains poorly understood. Some broad generalizations have been made; most work in the region is concerned primarily with descriptive and definitional issues rather than efforts to explain past human behavior or human-environmental interactions. Modern research directed towards understanding human adaptation to different environments remains in its infancy. Grounded in the powerful conceptual framework of evolutionary ecology and utilizing recent methodological advances, this work has shown that shifts in land-use strategies changes the opportunities for social and biological interaction among Late Pleistocene hominins in western Eurasia, bringing with it a plethora of important consequences for cultural and biological evolution. I employ, in my Dissertation, theoretical and methodological advances derived from human behavioral ecology (HBE) and lithic technology organization to show how variability in lithic technology can explain differences in technoeconomic choices and land-use strategies of Late Pleistocene foragers in Romanian Carpathians Basin and adjacent areas. Set against the backdrop of paleoenvironmental change, the principal questions I addressed are whether or not technological variation at the beginning of the Upper Paleolithic can account for fundamental changes at its end. The analysis of the Middle and Upper Paleolithic strata, from six archaeological sites, shows that the lithic industries were different not because of biocultural differences in technological organization, landuse strategies, and organizational flexibility. Instead the evidence suggests that technoeconomic strategies, the intensity of artifact curation and how foragers used the land appear to have been more closely related to changing environmental conditions, task-specific activities, and duration of occupation. This agrees well with the results of studies conducted in other areas and with those predicted from theoretically-derived models based on evolutionary ecology. My results lead to the conclusion that human landuse effectively changes the environment of selection for hominins and their lithic technologies, an important component of the interface between humans and the natural world. Foragers move across the landscape in comparable ways in very different ecological settings, cross-cutting both biological morphotypes and prehistorian-defined analytical units. / Dissertation/Thesis / Doctoral Dissertation Anthropology 2015

abstract: Why do many animals possess multiple classes of photoreceptors that vary in the wavelengths of light to which they are sensitive? Multiple spectral photoreceptor classes are a requirement for true color vision. However, animals may have unconventional vision, in which multiple spectral channels broaden the range of wavelengths that can be detected, or in which they use only a subset of receptors for specific behaviors. Branchiopod crustaceans are of interest for the study of unconventional color vision because they express multiple visual pigments in their compound eyes, have a simple repertoire of visually guided behavior, inhabit unique and highly variable light environments, and possess secondary neural simplifications. I first tested the behavioral responses of two representative species of branchiopods from separate orders, Streptocephalus mackini Anostracans (fairy shrimp), and Triops longicaudatus Notostracans (tadpole shrimp). I found that they maintain vertical position in the water column over a broad range of intensities and wavelengths, and respond behaviorally even at intensities below those of starlight. Accordingly, light intensities of their habitats at shallow depths tend to be dimmer than terrestrial habitats under starlight. Using models of how their compound eyes and the first neuropil of their optic lobe process visual cues, I infer that both orders of branchiopods use spatial summation from multiple compound eye ommatidia to respond at low intensities. Then, to understand if branchiopods use unconventional vision to guide these behaviors, I took electroretinographic recordings (ERGs) from their compound eyes and used models of spectral absorptance for a multimodel selection approach to make inferences about the number of photoreceptor classes in their eyes. I infer that both species have four spectral classes of photoreceptors that contribute to their ERGs, suggesting unconventional vision guides the described behavior. I extended the same modeling approach to other organisms, finding that the model inferences align with the empirically determined number of photoreceptor classes for this diverse set of organisms. This dissertation expands the conceptual framework of color vision research, indicating unconventional vision is more widespread than previously considered, and explains why some organisms have more spectral classes than would be expected from their behavioral repertoire. / Dissertation/Thesis / Doctoral Dissertation Biology 2016