Riparian deforestation is a significant threat to freshwater riverine ecosystems and sensitive fauna that depend on clean water. Sensitive aquatic species are vulnerable to the destruction of riparian forest cover which diminishes protection from pollutants, sedimentation, and solar radiation, while also depleting dissolved oxygen. In this thesis, I explore the influence of degraded riparian forest cover and its effect on dissolved oxygen on the embryonic viability of a sensitive freshwater habitat specialist, the Eastern hellbender. Hellbenders are a large-bodied, long-lived amphibian that inhabits fast flowing, cold mountain streams in the eastern United States. Over the last several decades, hellbender populations have experienced declines that are associated with low riparian forest cover, a geriatric population age structure, and high rates of nest failure. Adult male hellbenders normally provide extensive paternal care to embryos and larvae over an 8-month period. However, researchers have recently discovered that in degraded populations, hellbender nests are failing due to whole-clutch filial cannibalism by adult males. The underlying mechanism that triggers males to eat their young remains unknown, but one possibility is that eggs are not developing properly and as a result the attending male ceases to provide care. However, the embryonic viability of clutches developing in habitats with low riparian forest cover is unknown. Given the limited research on hellbender embryonic viability, I first sought to examine whether embryo viability is associated with a forest cover gradient. To accomplish this, I inherited two years of laboratory and field data, and I conducted a final third year of data collection for the study. Over these three years of data collection, I simultaneously evaluated embryo viability in a controlled captive rearing system while classifying nest failure due to whole-clutch cannibalism of sibling embryos in the field. I found significantly lower hellbender embryo viability, faster hatching times, and higher rates of underdeveloped hatchlings in hellbender populations with degraded riparian forest cover. However, I found no relationship between embryonic viability and whole-clutch filial cannibalism. I concluded that elevated specific conductance of water (i.e., dissolved ions) associated with loss of forest cover may affect the embryonic development of hellbenders, but this hypothesis has yet to be tested. To further explore the impact of degraded riparian forest cover on hellbender embryonic viability, I designed a study to evaluate the influence of depleted dissolved oxygen on embryonic development. To accomplish this, I designed and implemented a dissolved oxygen manipulation system to rear sibling embryos in high, medium, and low dissolved oxygen concentrations. I found that hellbender embryos are vulnerable to relatively modest reductions in dissolved oxygen (i.e., 5 mg/L), comparable to those found to affect embryos of some of the most sensitive species examined to date. Nests reared in low dissolved oxygen had a lower percentage of viable hatchlings, lower hatching success, higher rates of underdeveloped hatchlings, and smaller morphometrics compared to those reared at higher dissolved oxygen concentrations. I concluded that hellbenders may be particularly susceptible to reductions in dissolved oxygen because of their high degree of specialization for well oxygenated streams, and I suggest prioritizing additional research on dissolved oxygen to advance hellbender conservation. Taken together, my research established a foundation of knowledge on the sensitivities of hellbender embryos to degraded water quality caused by low riparian forest cover. Further, my work underscored the importance of riparian forest conservation for hellbender populations and for other freshwater biodiversity. Protection of riparian forest will also be critical to build resilience in streams against the existential threat of climate change. / Master of Science / Deforestation along rivers is a significant threat to freshwater systems and sensitive animals that depend on clean water. Sensitive river species are vulnerable to the destruction of forest cover which removes protection from pollutants, run off, sun light, and exposes them to low dissolved oxygen—the amount of oxygen in the water, which is necessary for the survival of aquatic animals. In this thesis, I explore how low forest cover may impact egg quality of a sensitive freshwater animal, the eastern hellbender salamander. Hellbenders are a large-bodied, long-lived amphibian that lives in fast-moving, cold mountain streams in the eastern United States. Over the last several decades, hellbender numbers have rapidly declined. These declines are associated with low forest cover along streams, elderly populations, and unsuccessful nests. Adult male hellbenders normally provide care to their eggs and hatchlings for about eight months after the eggs are laid. Researchers have recently found that in streams where hellbender numbers are declining, hellbender nests are not successful because adult males are eating their eggs before the eggs can hatch. Researchers know that low forest cover triggers hellbender fathers to cannibalize their eggs, but the mechanism by which forest cover causes this trigger is unknown. Unsuccessful nests due to cannibalistic fathers in declining populations suggest that hellbender eggs might be unhealthy, making it more profitable to eat them than to care for them. However, the health of eggs in water from low forest cover is unknown and an important knowledge gap to fill to uncover what is triggering cannibalism in low forest cover populations. Given the limited research on hellbender egg health, I first sought to examine whether egg health is associated with changes in forest cover. To do this, I received two years of laboratory and field data, and I conducted a final third year of data collection for the project. Over these three years of data collection, I determined egg health by raising eggs in their own site water in the lab and simultaneously determined whether siblings in the field were eaten by their dads. I found lower hellbender egg health, shorter time to hatch, and more premature hatchlings in hellbender populations with low forest cover. However, I found no relationship between egg health and whether hellbender dads ate their eggs. I concluded that poor water quality associated with loss of forest cover may affect the health of hellbender eggs, but this has not been tested yet. To further explore the impact of low forest cover on hellbender egg health, I designed a study to determine the influence of low dissolved oxygen on egg health and growth. To do this, I designed and implemented a dissolved oxygen experiment to raise sibling eggs in high, medium, and low dissolved oxygen levels. I found that hellbender eggs are vulnerable to relatively moderate levels of reduced oxygen (i.e., 5 mg/l), which is comparable to how other highly sensitive species react. Nests raised in low dissolved oxygen had a lower number of healthy hatchlings, lower number of eggs that hatched, higher numbers of premature hatchlings, and smaller sizes compared to those raised in higher dissolved oxygen levels. I concluded that hellbenders may be particularly vulnerable to low dissolved oxygen because they have adapted to live in well oxygenated streams, and I emphasize the importance of studying dissolved oxygen to inform hellbender conservation. Taken together, my research created a foundation of knowledge on the vulnerabilities of hellbender eggs to poor water quality because of low forest cover. Filling these knowledge gaps is important, because it informs policy makers and managers on how to best protect hellbenders and other stream animals from declining. Further, my work emphasized the importance of forest conservation for hellbenders and for other sensitive freshwater animals. Forested streams will also be important to build protection against other human disturbances, such as climate change.

<p>Natural and sexual selection and sexual conflict are forces shaping the evolution of reproductive behaviour, while constrained by factors like environment, physiology and life-history trade-offs. Parental care is costly both in terms of time and energy. In fish, filial cannibalism is a strategy for caring males to compensate for some of the energy loss. Human impact like eutrophication also alters the basics for animals living in that environment. It is fundamental to any species to adjust its behaviour to a changing environment. Studying sand goby males, I found trade-offs both regarding parental care allocations, like ventilation vs. predator defence, and investment in present vs. future reproductive success. Paternal sand gobies exposed to water with low oxygen levels increased their fanning effort but did not compensate by eating more eggs, even though an increased current parental effort should affect future reproductive success negatively. Investigating if patterns of filial cannibalism change with time of season, I found no correlation. Theory predicts that it should pay more to eat eggs early, when future mating potential is higher than late in the season. However, as early hatching fry are likely to gain higher fitness through larger size the next breeding season, this may provide an opposing selection pressure. In species with male care the only way a female can affect the level of post-spawning care is by choosing a good mate. A female preference to spawn in nests that already contain eggs of other females has been interpreted as a means to avoid filial cannibalism through a dilution effect or to decrease the costs of search time. Yet, in my study females did not avoid filial cannibalism by preferring large clutches to small ones. Oxygenation of the eggs might be a key factor, since both large and small females preferred spawning in nests with small clutches. Thus, as in most animals, trade-offs clearly govern the reproductive behaviour of sand gobies.</p>

In some species with uniparental care, when the parental individual deserts the offspring or dies, its mate may adopt the parental responsibilities, a behaviour known as flexible compensation of uniparental care. However, for most of the species in which this behavioural response has been reported in the literature, there was no thorough investigation of its effects upon offspring fitness to support it. In the Neotropical harvestman Serracutisoma proximum, a species with female uniparental care, harem-owning males stand on unattended clutches when the egg-tending females desert or die. Here, we investigate if this species constitutes an actual example of flexible compensation of uniparental care by evaluating the effects of ecological (namely climatic conditions and mate availability) and life-history factors (namely male attractiveness, clutch size and filial cannibalism) on males\' behaviour and, ultimately, on offspring survival. We expect unfavourable climatic conditions, as well as low mate availability and male attractiveness, to negatively affect the exhibition of compensatory behaviours, but a positive effect of clutch size. Moreover, we expect an increase in the extent of filial cannibalism to be a strategy that alleviates the costs of compensation. However, males\' behaviour towards unattended offspring has no effect on offspring survival, and other results were inconsistent with our predictions except for climatic conditions affecting the extent of both the presumed behaviour of standing on unattended clutches and filial cannibalism. Our results, therefore, do not support the existence of flexible compensation of uniparental care in S. proximum. Rather than a parental activity, we argue that males&rsquo; behaviour corresponds to a mating tactic that increases fertilization and mating success with the returning egg-tending females or newcomer mates foraging on the unattended offspring. This study demonstrates that for presumed cases of flexible compensation of uniparental care it is necessary to evaluate the effects on offspring fitness since not always an apparent parental behaviour is what it seems to be / Em algumas espécies com cuidado uniparental, quando o individuo parental deserta a prole ou morre, seu parceiro pode adotar as responsabilidades parentais, um comportamento conhecido como compensação flexível do cuidado uniparental. No entanto, para a maioria das espécies em que esta resposta comportamental tem sido relatada na literatura, não houve uma investigação completa de seus efeitos sobre a aptidão da prole para apoiá-lo. No opilião Neotropical Serracutisoma proximum, uma espécie com cuidado uniparental da fêmea, os machos donos de harem ficam sobre desovas desatendidas quando as fêmeas que deveriam cuidar dos ovos desertam ou morrem. Aqui, investigamos se esta espécie constitui um exemplo real de compensação flexível do cuidado uniparental, avaliando os efeitos de fatores ecológicos (condições climáticas e disponibilidade de parceiras) e de história de vida (atratividade do macho, tamanho da desova e canibalismo filial) sobre o comportamento dos machos e, em última instância, sobre a sobrevivência da prole. Esperamos que condições climáticas desfavoráveis, assim como baixa disponibilidade de parceiras e atratividade do macho, afetem negativamente a exibição de comportamentos compensatórios, mas que o tamanho da desova tenha um efeito positivo. Além disso, esperamos que um aumento na intensidade do canibalismo filial seja uma estratégia que alivie os custos da compensação. Entretanto, o comportamento dos machos em relação à prole desatendida não tem efeito sobre a sobrevivência da prole, e os outros resultados foram inconsistentes com nossas previsões, exceto pelas condições climáticas que afetam tanto a intensidade do comportamento dos machos sobre as desovas quanto o canibalismo filial. Nossos resultados, portanto, não apoiam a existência de compensação flexível do cuidado uniparental em S. proximum. Ao invés de uma atividade parental, argumentamos que o comportamento dos machos corresponde a uma tática de acasalamento que aumenta o sucesso de fertilização e de acasalamento com as fêmeas que retornam às suas desovas ou com novas fêmeas que se alimentam dos ovos desprotegidos. Este estudo demonstra que, para os casos em que se pressupõe compensação flexível de cuidado uniparental, é necessário avaliar os efeitos sobre a aptidão prole, uma vez que nem sempre um aparente comportamento parental é o que parece ser

Natural and sexual selection and sexual conflict are forces shaping the evolution of reproductive behaviour, while constrained by factors like environment, physiology and life-history trade-offs. Parental care is costly both in terms of time and energy. In fish, filial cannibalism is a strategy for caring males to compensate for some of the energy loss. Human impact like eutrophication also alters the basics for animals living in that environment. It is fundamental to any species to adjust its behaviour to a changing environment. Studying sand goby males, I found trade-offs both regarding parental care allocations, like ventilation vs. predator defence, and investment in present vs. future reproductive success. Paternal sand gobies exposed to water with low oxygen levels increased their fanning effort but did not compensate by eating more eggs, even though an increased current parental effort should affect future reproductive success negatively. Investigating if patterns of filial cannibalism change with time of season, I found no correlation. Theory predicts that it should pay more to eat eggs early, when future mating potential is higher than late in the season. However, as early hatching fry are likely to gain higher fitness through larger size the next breeding season, this may provide an opposing selection pressure. In species with male care the only way a female can affect the level of post-spawning care is by choosing a good mate. A female preference to spawn in nests that already contain eggs of other females has been interpreted as a means to avoid filial cannibalism through a dilution effect or to decrease the costs of search time. Yet, in my study females did not avoid filial cannibalism by preferring large clutches to small ones. Oxygenation of the eggs might be a key factor, since both large and small females preferred spawning in nests with small clutches. Thus, as in most animals, trade-offs clearly govern the reproductive behaviour of sand gobies.

In some species with uniparental care, when the parental individual deserts the offspring or dies, its mate may adopt the parental responsibilities, a behaviour known as flexible compensation of uniparental care. However, for most of the species in which this behavioural response has been reported in the literature, there was no thorough investigation of its effects upon offspring fitness to support it. In the Neotropical harvestman Serracutisoma proximum, a species with female uniparental care, harem-owning males stand on unattended clutches when the egg-tending females desert or die. Here, we investigate if this species constitutes an actual example of flexible compensation of uniparental care by evaluating the effects of ecological (namely climatic conditions and mate availability) and life-history factors (namely male attractiveness, clutch size and filial cannibalism) on males\' behaviour and, ultimately, on offspring survival. We expect unfavourable climatic conditions, as well as low mate availability and male attractiveness, to negatively affect the exhibition of compensatory behaviours, but a positive effect of clutch size. Moreover, we expect an increase in the extent of filial cannibalism to be a strategy that alleviates the costs of compensation. However, males\' behaviour towards unattended offspring has no effect on offspring survival, and other results were inconsistent with our predictions except for climatic conditions affecting the extent of both the presumed behaviour of standing on unattended clutches and filial cannibalism. Our results, therefore, do not support the existence of flexible compensation of uniparental care in S. proximum. Rather than a parental activity, we argue that males&rsquo; behaviour corresponds to a mating tactic that increases fertilization and mating success with the returning egg-tending females or newcomer mates foraging on the unattended offspring. This study demonstrates that for presumed cases of flexible compensation of uniparental care it is necessary to evaluate the effects on offspring fitness since not always an apparent parental behaviour is what it seems to be / Em algumas espécies com cuidado uniparental, quando o individuo parental deserta a prole ou morre, seu parceiro pode adotar as responsabilidades parentais, um comportamento conhecido como compensação flexível do cuidado uniparental. No entanto, para a maioria das espécies em que esta resposta comportamental tem sido relatada na literatura, não houve uma investigação completa de seus efeitos sobre a aptidão da prole para apoiá-lo. No opilião Neotropical Serracutisoma proximum, uma espécie com cuidado uniparental da fêmea, os machos donos de harem ficam sobre desovas desatendidas quando as fêmeas que deveriam cuidar dos ovos desertam ou morrem. Aqui, investigamos se esta espécie constitui um exemplo real de compensação flexível do cuidado uniparental, avaliando os efeitos de fatores ecológicos (condições climáticas e disponibilidade de parceiras) e de história de vida (atratividade do macho, tamanho da desova e canibalismo filial) sobre o comportamento dos machos e, em última instância, sobre a sobrevivência da prole. Esperamos que condições climáticas desfavoráveis, assim como baixa disponibilidade de parceiras e atratividade do macho, afetem negativamente a exibição de comportamentos compensatórios, mas que o tamanho da desova tenha um efeito positivo. Além disso, esperamos que um aumento na intensidade do canibalismo filial seja uma estratégia que alivie os custos da compensação. Entretanto, o comportamento dos machos em relação à prole desatendida não tem efeito sobre a sobrevivência da prole, e os outros resultados foram inconsistentes com nossas previsões, exceto pelas condições climáticas que afetam tanto a intensidade do comportamento dos machos sobre as desovas quanto o canibalismo filial. Nossos resultados, portanto, não apoiam a existência de compensação flexível do cuidado uniparental em S. proximum. Ao invés de uma atividade parental, argumentamos que o comportamento dos machos corresponde a uma tática de acasalamento que aumenta o sucesso de fertilização e de acasalamento com as fêmeas que retornam às suas desovas ou com novas fêmeas que se alimentam dos ovos desprotegidos. Este estudo demonstra que, para os casos em que se pressupõe compensação flexível de cuidado uniparental, é necessário avaliar os efeitos sobre a aptidão prole, uma vez que nem sempre um aparente comportamento parental é o que parece ser

Cavidades naturais constituem um recurso reprodutivo monopolizável, cuja disponibilidade determina a intensidade da competição macho-macho que, por sua vez, pode influenciar os custos do cuidado paternal, pois quanto maior o risco de perda do recurso, menor deve ser a freqüência de forrageio dos machos. Fêmeas do opilião Magnispina neptunus utilizam cavidades naturais como sítios de nidificação, que são monopolizados por machos em um sistema de poliginia por defesa de recursos. Após a oviposição, as fêmeas abandonam os ninhos e todo o cuidado parental é exercido pelos machos. O objetivo deste trabalho foi avaliar o efeito da disponibilidade de ninhos sobre a competição intrassexual e os custos do cuidado paternal em M. neptunus. Dois grupos experimentais foram estabelecidos em laboratório: alta (8 ninhos) e baixa (4 ninhos) disponibilidade de sítios reprodutivos. Cada grupo era composto por sete terrários contendo 12 machos e 12 fêmeas. Apesar da redução dos sítios de nidificação ter reduzido o número de machos detentores de ninhos, o tamanho dos indivíduos não exerceu nenhum efeito sobre o sucesso de monopolização de ninhos. Aparentemente, o pequeno diâmetro de entrada dos ninhos experimentais reduz a importância do tamanho dos machos sobre sua probabilidade de monopolizar um sítio de nidificação. Apenas a residência prévia parece determinar o resultado das disputas entre machos e, portanto, os custos de abandonar os ninhos devem ser altos. De fato, machos guardiões permanecem quase todo o tempo dentro dos seus ninhos em ambos os grupos experimentais, o que aumenta os custos do cuidado. Como conseqüência, o canibalismo filial é freqüente, pois os custos em termos de redução no tamanho da desova são menores do que os benefícios de manter a posse do ninho e dos ovos, que sabidamente aumentam a atratividade dos machos e suas chances de obterem novas desovas / Natural cavities are a reproductive resource that can be monopolized, and their availability determines the intensity of male-male competition, which in turn may influence the costs of paternal care since the higher the risk of loosing the resource, the low should be male foraging frequency. Females of the harvestman Magnispina neptunus use natural cavities as nest site, which are monopolized by males in typical resource defense polygyny mating system. After oviposition, females leave the nests and all parental activities are in charge of males. The goal of this study was to evaluate the effect of nest site availability on the intrasexual competition and the costs of paternal care in M. neptunus. Two experimental groups were established in the laboratory: high (8 nests) and low (4 nests) nest site availability. Each group was composed of seven terraria containing 12 males and 12 females. Although the reduction of in the number of nests sites has indeed reduced the number of males holding nests, male size did not influence the probability of acquiring and maintaining a nest. Apparently, the small diameter of entrance in the experimental nests reduces the importance of male size on their resource holding power. Only previous residence seems to influence the output of agonistic interactions between two males and, therefore, the costs of leaving the nest to forage are likely to be high. In fact, guarding males remain almost all the time inside their nests, which increases the costs of paternal care. As a consequence, filial cannibalism is frequent in both experimental groups because the costs in terms of reduction in clutch size are likely to lower than the benefits of holding a nest containing eggs, which is known to increase male attractiveness and his chance of acquiring additional eggs

Cavidades naturais constituem um recurso reprodutivo monopolizável, cuja disponibilidade determina a intensidade da competição macho-macho que, por sua vez, pode influenciar os custos do cuidado paternal, pois quanto maior o risco de perda do recurso, menor deve ser a freqüência de forrageio dos machos. Fêmeas do opilião Magnispina neptunus utilizam cavidades naturais como sítios de nidificação, que são monopolizados por machos em um sistema de poliginia por defesa de recursos. Após a oviposição, as fêmeas abandonam os ninhos e todo o cuidado parental é exercido pelos machos. O objetivo deste trabalho foi avaliar o efeito da disponibilidade de ninhos sobre a competição intrassexual e os custos do cuidado paternal em M. neptunus. Dois grupos experimentais foram estabelecidos em laboratório: alta (8 ninhos) e baixa (4 ninhos) disponibilidade de sítios reprodutivos. Cada grupo era composto por sete terrários contendo 12 machos e 12 fêmeas. Apesar da redução dos sítios de nidificação ter reduzido o número de machos detentores de ninhos, o tamanho dos indivíduos não exerceu nenhum efeito sobre o sucesso de monopolização de ninhos. Aparentemente, o pequeno diâmetro de entrada dos ninhos experimentais reduz a importância do tamanho dos machos sobre sua probabilidade de monopolizar um sítio de nidificação. Apenas a residência prévia parece determinar o resultado das disputas entre machos e, portanto, os custos de abandonar os ninhos devem ser altos. De fato, machos guardiões permanecem quase todo o tempo dentro dos seus ninhos em ambos os grupos experimentais, o que aumenta os custos do cuidado. Como conseqüência, o canibalismo filial é freqüente, pois os custos em termos de redução no tamanho da desova são menores do que os benefícios de manter a posse do ninho e dos ovos, que sabidamente aumentam a atratividade dos machos e suas chances de obterem novas desovas / Natural cavities are a reproductive resource that can be monopolized, and their availability determines the intensity of male-male competition, which in turn may influence the costs of paternal care since the higher the risk of loosing the resource, the low should be male foraging frequency. Females of the harvestman Magnispina neptunus use natural cavities as nest site, which are monopolized by males in typical resource defense polygyny mating system. After oviposition, females leave the nests and all parental activities are in charge of males. The goal of this study was to evaluate the effect of nest site availability on the intrasexual competition and the costs of paternal care in M. neptunus. Two experimental groups were established in the laboratory: high (8 nests) and low (4 nests) nest site availability. Each group was composed of seven terraria containing 12 males and 12 females. Although the reduction of in the number of nests sites has indeed reduced the number of males holding nests, male size did not influence the probability of acquiring and maintaining a nest. Apparently, the small diameter of entrance in the experimental nests reduces the importance of male size on their resource holding power. Only previous residence seems to influence the output of agonistic interactions between two males and, therefore, the costs of leaving the nest to forage are likely to be high. In fact, guarding males remain almost all the time inside their nests, which increases the costs of paternal care. As a consequence, filial cannibalism is frequent in both experimental groups because the costs in terms of reduction in clutch size are likely to lower than the benefits of holding a nest containing eggs, which is known to increase male attractiveness and his chance of acquiring additional eggs