Seasonality influences life history through its effect on the availability of essential resources, with birds timing breeding to occur during peak food availability. Due to density-dependence, investment in breeding is determined largely by the seasonality of food availability, with an increased investment being traded-off against adult survival. A bird's mass acts as an index of a species' foraging environment, because a bird bases its foraging decisions on a trade-off between the risk of predation and the risk of starvation. Under constant predation risk a bird increases its mass as insurance against increased foraging unpredictability. In tropical savannahs day length and temperature remains relatively constant, and there is not a season of increased density-dependent mortality which acts across all species. Thus species have evolved a broad range of life history traits under the same environmental conditions, although how a species experiences seasonality depends largely on its foraging niche. This thesis shows that most savannah species varied their mass across the year, having a reduced mass in the non-breeding season which suggests that foraging remained predictable. Independent of gonad or egg growth they then increased their mass as they started to breed, with the timing of breeding coinciding with peak food availability. Across species in the same foraging niche mass acts as an index of breeding investment, with females increasing their mass more than males. While across species in different foraging niches an increased mass response was associated with higher adult survival, probably because breeding strategy and subsequently adult survival are governed by food limitation. This thesis shows that birds adaptively manage their mass during breeding and that mass is not a result of energetic stress, thus under constant predation risk a bird's mass is a result of foraging predictability as a function of competition for available food and investment in breeding.

Animal community structures, life histories and individual foraging behaviour are all an outcome of a trade-off between competition for resources (and thus the risk of starvation) and survival (and thus the risk of predation). The relative importance of these factors however, differs between ecosystems, and especially when comparing temperate to tropical ones, we usually find marked differences. The seasonality of tropical ecosystems is much reduced compared to temperate ones, and weather conditions are less extreme. Accordingly tropical systems are characterised by higher species diversity, and different life history traits have been found between temperate and tropical birds. However, how the different environmental factors interact, and how predation and starvation risk vary to cause these differences still remains largely unknown. We studied the feeding behaviour of several granivorous Estrildid finches in scrub savannah habitat in central Nigeria to test how they respond to varying degrees of starvation and predation risk. During field observations and aviary experiments we investigated whether there is seasonal variation in the birds’ foraging behaviour correlating with the abundance of grass seeds and tested how they respond to different group sizes and differing distances from cover (representing a difference in predation risk). Further we also carried out field observation on the natural feeding behaviour of several closely related sympartic Estrildid finches to investigate inter-specific and seasonal differences in competition and microhabitat choice to see if this could explain their coexistence. Finally we studied habitat choice, movement behaviour and breeding biology of the potentially threatened endemic Rock Firefinch (Lagonosticta sanguinodorsalis) between the wet and the dry season via radio-tracking to establish its habitat requirements and gain the first information in its life history traits and population trends. We found little seasonal variation in the species’ foraging behaviour, and parameters that varied did not do so in a consistent manner. Thus, we found little evidence for a seasonal change in the risk of starvation. However, the abundance of several bird species varied widely between seasons and species leaving during periods of food shortage might have released competition for remaining resources. Birds did not show a strong response in their feeding behaviour with respect to cover in either intake rate or timing of feeding. However, intake rate increased with group size, which we believe to be due to scramble competition rather than risk dilution. We therefore conclude that predation did not shape the foraging behaviour of tropical granivorous passerines as markedly as that of temperate ones. Rock Firefinches were found to breed between the late rainy and the early dry season. They selected inselberg habitat, where most nests were found between rocky boulders. During the dry season, when water sources in inselberg habitat had dried out, they had to fly distances of up to 700 m to the gallery forest to get water and this led to the inclusion of more scrub savannah and gallery forest within their home ranges. Daily egg survival was 0.89 ± 0.03 calculated after the Mayfield analysis and most failing nests were depredated probably mainly by lizards. We suggest that in addition to nest predation, water availability might limit breeding time and thus reproductive output of Rock Firefinches. Predation risk did not seem to be of high importance in shaping the birds’ feeding behaviour because there was no seasonal variation in the risk of starvation. We found some suggestive evidence that competition might be important and it is likely that bird populations constantly stay close to carrying capacity. In contrast to temperate regions the need to conserve water might be of higher importance in shaping the birds’ feeding behaviour. High adult survival rates might be due to reduced seasonality in the risk of starvation thereby leading to reduced predation risk on adult birds. High nest predation might also be of higher importance in shaping the life history traits of tropical passerines, but at present this suggestion remains speculative. The results fit into the general framework that there is a trade-off between starvation and predation risk, and in the absence of starvation risk for some species in tropical areas, predation risk is also relatively unimportant.

A observação de gradientes latitudinais em aspectos da história de vida de aves tem motivado o estudo da evolução e variabilidade das histórias de vida nestes organismos. Um exemplo bem documentado é a variação no tamanho da ninhada, onde aves de latitudes menores tendem a ter ninhadas menores do que os seus homólogos de latitudes altas. Uma hipótese que visa explicar esta variação propõe que a sobrevivência em latitudes tropicais é maior para compensar o tamanho da ninhada menor e evitar a extinção das populações. Esta explicação tem tido grande aceitação e apoio por parte de alguns estudos, mas tem sido questionada por outros que não encontraram taxas de sobrevivência mais elevadas em aves tropicais. De modo implícito, todos estes estudos basearam seus resultados na sobrevivência de indivíduos adultos. As populações com o tamanho da ninhada menor não poderiam crescer da mesma maneira que as populações com ninhadas maiores; portanto, se justifica acreditar que algo deve mudar com a latitude para manter o balanço em tamanho populacional. Na busca por explicações alternativas para a persistência das populações de aves tropicais com relativamente pequenos tamanhos de ninhada, surge outra hipótese que propõe que, se não houver diferenças na sobrevivência de indivíduos adultos entre latitudes, o aspecto fundamental que varia é a sobrevivência juvenil, com sobrevivência maior para os juvenis das zonas tropicais em comparação com os juvenis das zonas temperadas. No entanto, atualmente há pouca evidência que suporta esta conclusão. Os resultados contrastantes desses estudos sugerem a falta de um consenso geral sobre a hipótese de que as aves tropicais têm taxas de sobrevivência mais elevadas do que as aves de regiões temperadas, motivando a formulação de hipóteses alternativas e convidando novos testes de hipótese. Neste estudo, pretendemos a) avaliar o efeito da idade sobre a sobrevivência em aves tropicais, estimando as probabilidades anuais de sobrevivência aparentes idade-específicas para um conjunto de aves passeriformes de sub-bosque na Amazônia central brasileira; e b) contribuir para o debate sobre o gradiente latitudinal na sobrevivência de adultos, comparando nossas estimativas com estimativas de outras latitudes. Para estimar a sobrevivência idade-específica ajustamos aos nossos dados um modelo Cormack-Jolly-Seber (CJS) hierárquico para n espécies, que trata os parâmetros espécie-específicos como efeitos aleatórios, que são estimados e que descrevem todo o conjunto de espécies; para comparação de métodos, ajustamos uma versão de efeitos fixos do modelo. Para a determinação da idade das aves usamos o sistema WRP. Apresentamos uma nova variante do modelo CJS com um parâmetro de mistura para a sobrevivência de aves de idade incerta no momento da primeira captura. Encontramos efeito forte da idade na sobrevivência, com probabilidades de sobrevivência menor para os jovens do que para os adultos; evidência de efeito latitude sobre a sobrevivência, que suporta a hipótese amplamente aceita de variação na sobrevivência com a latitude; e discutimos diferenças metodológicas interessantes entre modelo de efeitos aleatórios e fixos relacionados com a precisão das estimativas e o âmbito de inferência, que nos levam a concluir que os modelos de efeitos aleatórios são os mais adequados para a nossa análise. Concluímos que não é necessário invocar uma hipótese alternativa de maior sobrevivência juvenil nos trópicos a fim de explicar o gradiente latitudinal no tamanho da ninhada. / The observation of latitudinal gradients in bird life history traits has motivated the study of avian life history evolution and variability. A well-documented example is the variation in clutch size, where lower latitude birds tend to have smaller clutches than their higher latitude counterparts. A hypothesis that explains this variation proposes that survival in tropical latitudes is higher to compensate for smaller clutch size and prevent population extinctions. This explanation has had a wide acceptance and support by some studies, but has been questioned by others who have not found such higher survival rates in tropical birds. In an implicit manner, all these studies have based their results on adult survival. Populations with smaller clutch size would not be able to grow as well as populations with larger clutches; therefore one is justified to believe that something else must change with latitude. In the search for alternative explanations to the persistence of tropical bird populations with relatively small clutch sizes it has also been proposed that, if there were no differences in adult survival among latitudes, the fundamental trait that varies is juvenile survival, with higher survival rates for tropical juveniles birds than for temperate ones. However, currently there is little evidence that supports this conclusion. The contrasting results of those studies suggest a lack of a general consensus about the hypothesis that tropical birds have higher survival rates than birds of temperate regions, motivating the formulation of alternative hypotheses, and inviting further tests of the hypothesis. In our study we aim to a) assess the effect of age on survival in a tropical bird community, estimating age-specific annual apparent survival probabilities for a set of passerine understory birds from the central Brazilian Amazon; and b) contribute to the debate about the latitudinal gradient in adult survival by comparing our adult survival estimates to estimates of temperate-zone adult survival probabilities. To estimate the age-specific survival we fit to our data a hierarchical multispecies Cormack-Jolly-Seber (CJS) model for n species, that treats species-specific parameters as random effects that are estimated and that describe the whole assemblage of species; for comparison of methods, we also fit a fixed-effects version of the model. To age birds we use the cycle-based WRP system. We introduce a novel variant of CJS model with a mixture component for the survival of birds of uncertain age at the time of banding. We found strong effect of age on survival, with juveniles surviving less than adults; evidence of latitude effect on survival, that supports the widely accepted hypothesis of variation on survival with latitude; and methodological differences between random and fixed effects model related to precision of estimates and scope of inference, that lead us to conclude that random-effects models are more appropriate for our analysis. We conclude that there is no reason for an alternative latitudinal trend in juvenile survival to account for the general trend in clutch size.

A observação de gradientes latitudinais em aspectos da história de vida de aves tem motivado o estudo da evolução e variabilidade das histórias de vida nestes organismos. Um exemplo bem documentado é a variação no tamanho da ninhada, onde aves de latitudes menores tendem a ter ninhadas menores do que os seus homólogos de latitudes altas. Uma hipótese que visa explicar esta variação propõe que a sobrevivência em latitudes tropicais é maior para compensar o tamanho da ninhada menor e evitar a extinção das populações. Esta explicação tem tido grande aceitação e apoio por parte de alguns estudos, mas tem sido questionada por outros que não encontraram taxas de sobrevivência mais elevadas em aves tropicais. De modo implícito, todos estes estudos basearam seus resultados na sobrevivência de indivíduos adultos. As populações com o tamanho da ninhada menor não poderiam crescer da mesma maneira que as populações com ninhadas maiores; portanto, se justifica acreditar que algo deve mudar com a latitude para manter o balanço em tamanho populacional. Na busca por explicações alternativas para a persistência das populações de aves tropicais com relativamente pequenos tamanhos de ninhada, surge outra hipótese que propõe que, se não houver diferenças na sobrevivência de indivíduos adultos entre latitudes, o aspecto fundamental que varia é a sobrevivência juvenil, com sobrevivência maior para os juvenis das zonas tropicais em comparação com os juvenis das zonas temperadas. No entanto, atualmente há pouca evidência que suporta esta conclusão. Os resultados contrastantes desses estudos sugerem a falta de um consenso geral sobre a hipótese de que as aves tropicais têm taxas de sobrevivência mais elevadas do que as aves de regiões temperadas, motivando a formulação de hipóteses alternativas e convidando novos testes de hipótese. Neste estudo, pretendemos a) avaliar o efeito da idade sobre a sobrevivência em aves tropicais, estimando as probabilidades anuais de sobrevivência aparentes idade-específicas para um conjunto de aves passeriformes de sub-bosque na Amazônia central brasileira; e b) contribuir para o debate sobre o gradiente latitudinal na sobrevivência de adultos, comparando nossas estimativas com estimativas de outras latitudes. Para estimar a sobrevivência idade-específica ajustamos aos nossos dados um modelo Cormack-Jolly-Seber (CJS) hierárquico para n espécies, que trata os parâmetros espécie-específicos como efeitos aleatórios, que são estimados e que descrevem todo o conjunto de espécies; para comparação de métodos, ajustamos uma versão de efeitos fixos do modelo. Para a determinação da idade das aves usamos o sistema WRP. Apresentamos uma nova variante do modelo CJS com um parâmetro de mistura para a sobrevivência de aves de idade incerta no momento da primeira captura. Encontramos efeito forte da idade na sobrevivência, com probabilidades de sobrevivência menor para os jovens do que para os adultos; evidência de efeito latitude sobre a sobrevivência, que suporta a hipótese amplamente aceita de variação na sobrevivência com a latitude; e discutimos diferenças metodológicas interessantes entre modelo de efeitos aleatórios e fixos relacionados com a precisão das estimativas e o âmbito de inferência, que nos levam a concluir que os modelos de efeitos aleatórios são os mais adequados para a nossa análise. Concluímos que não é necessário invocar uma hipótese alternativa de maior sobrevivência juvenil nos trópicos a fim de explicar o gradiente latitudinal no tamanho da ninhada. / The observation of latitudinal gradients in bird life history traits has motivated the study of avian life history evolution and variability. A well-documented example is the variation in clutch size, where lower latitude birds tend to have smaller clutches than their higher latitude counterparts. A hypothesis that explains this variation proposes that survival in tropical latitudes is higher to compensate for smaller clutch size and prevent population extinctions. This explanation has had a wide acceptance and support by some studies, but has been questioned by others who have not found such higher survival rates in tropical birds. In an implicit manner, all these studies have based their results on adult survival. Populations with smaller clutch size would not be able to grow as well as populations with larger clutches; therefore one is justified to believe that something else must change with latitude. In the search for alternative explanations to the persistence of tropical bird populations with relatively small clutch sizes it has also been proposed that, if there were no differences in adult survival among latitudes, the fundamental trait that varies is juvenile survival, with higher survival rates for tropical juveniles birds than for temperate ones. However, currently there is little evidence that supports this conclusion. The contrasting results of those studies suggest a lack of a general consensus about the hypothesis that tropical birds have higher survival rates than birds of temperate regions, motivating the formulation of alternative hypotheses, and inviting further tests of the hypothesis. In our study we aim to a) assess the effect of age on survival in a tropical bird community, estimating age-specific annual apparent survival probabilities for a set of passerine understory birds from the central Brazilian Amazon; and b) contribute to the debate about the latitudinal gradient in adult survival by comparing our adult survival estimates to estimates of temperate-zone adult survival probabilities. To estimate the age-specific survival we fit to our data a hierarchical multispecies Cormack-Jolly-Seber (CJS) model for n species, that treats species-specific parameters as random effects that are estimated and that describe the whole assemblage of species; for comparison of methods, we also fit a fixed-effects version of the model. To age birds we use the cycle-based WRP system. We introduce a novel variant of CJS model with a mixture component for the survival of birds of uncertain age at the time of banding. We found strong effect of age on survival, with juveniles surviving less than adults; evidence of latitude effect on survival, that supports the widely accepted hypothesis of variation on survival with latitude; and methodological differences between random and fixed effects model related to precision of estimates and scope of inference, that lead us to conclude that random-effects models are more appropriate for our analysis. We conclude that there is no reason for an alternative latitudinal trend in juvenile survival to account for the general trend in clutch size.

A observação de gradientes latitudinais em aspectos da história de vida de aves tem motivado o estudo da evolução e variabilidade das histórias de vida nestes organismos. Um exemplo bem documentado é a variação no tamanho da ninhada, onde aves de latitudes menores tendem a ter ninhadas menores do que os seus homólogos de latitudes altas. Uma hipótese que visa explicar esta variação propõe que a sobrevivência em latitudes tropicais é maior para compensar o tamanho da ninhada menor e evitar a extinção das populações. Esta explicação tem tido grande aceitação e apoio por parte de alguns estudos, mas tem sido questionada por outros que não encontraram taxas de sobrevivência mais elevadas em aves tropicais. De modo implícito, todos estes estudos basearam seus resultados na sobrevivência de indivíduos adultos. As populações com o tamanho da ninhada menor não poderiam crescer da mesma maneira que as populações com ninhadas maiores; portanto, se justifica acreditar que algo deve mudar com a latitude para manter o balanço em tamanho populacional. Na busca por explicações alternativas para a persistência das populações de aves tropicais com relativamente pequenos tamanhos de ninhada, surge outra hipótese que propõe que, se não houver diferenças na sobrevivência de indivíduos adultos entre latitudes, o aspecto fundamental que varia é a sobrevivência juvenil, com sobrevivência maior para os juvenis das zonas tropicais em comparação com os juvenis das zonas temperadas. No entanto, atualmente há pouca evidência que suporta esta conclusão. Os resultados contrastantes desses estudos sugerem a falta de um consenso geral sobre a hipótese de que as aves tropicais têm taxas de sobrevivência mais elevadas do que as aves de regiões temperadas, motivando a formulação de hipóteses alternativas e convidando novos testes de hipótese. Neste estudo, pretendemos a) avaliar o efeito da idade sobre a sobrevivência em aves tropicais, estimando as probabilidades anuais de sobrevivência aparentes idade-específicas para um conjunto de aves passeriformes de sub-bosque na Amazônia central brasileira; e b) contribuir para o debate sobre o gradiente latitudinal na sobrevivência de adultos, comparando nossas estimativas com estimativas de outras latitudes. Para estimar a sobrevivência idade-específica ajustamos aos nossos dados um modelo Cormack-Jolly-Seber (CJS) hierárquico para n espécies, que trata os parâmetros espécie-específicos como efeitos aleatórios, que são estimados e que descrevem todo o conjunto de espécies; para comparação de métodos, ajustamos uma versão de efeitos fixos do modelo. Para a determinação da idade das aves usamos o sistema WRP. Apresentamos uma nova variante do modelo CJS com um parâmetro de mistura para a sobrevivência de aves de idade incerta no momento da primeira captura. Encontramos efeito forte da idade na sobrevivência, com probabilidades de sobrevivência menor para os jovens do que para os adultos; evidência de efeito latitude sobre a sobrevivência, que suporta a hipótese amplamente aceita de variação na sobrevivência com a latitude; e discutimos diferenças metodológicas interessantes entre modelo de efeitos aleatórios e fixos relacionados com a precisão das estimativas e o âmbito de inferência, que nos levam a concluir que os modelos de efeitos aleatórios são os mais adequados para a nossa análise. Concluímos que não é necessário invocar uma hipótese alternativa de maior sobrevivência juvenil nos trópicos a fim de explicar o gradiente latitudinal no tamanho da ninhada. / The observation of latitudinal gradients in bird life history traits has motivated the study of avian life history evolution and variability. A well-documented example is the variation in clutch size, where lower latitude birds tend to have smaller clutches than their higher latitude counterparts. A hypothesis that explains this variation proposes that survival in tropical latitudes is higher to compensate for smaller clutch size and prevent population extinctions. This explanation has had a wide acceptance and support by some studies, but has been questioned by others who have not found such higher survival rates in tropical birds. In an implicit manner, all these studies have based their results on adult survival. Populations with smaller clutch size would not be able to grow as well as populations with larger clutches; therefore one is justified to believe that something else must change with latitude. In the search for alternative explanations to the persistence of tropical bird populations with relatively small clutch sizes it has also been proposed that, if there were no differences in adult survival among latitudes, the fundamental trait that varies is juvenile survival, with higher survival rates for tropical juveniles birds than for temperate ones. However, currently there is little evidence that supports this conclusion. The contrasting results of those studies suggest a lack of a general consensus about the hypothesis that tropical birds have higher survival rates than birds of temperate regions, motivating the formulation of alternative hypotheses, and inviting further tests of the hypothesis. In our study we aim to a) assess the effect of age on survival in a tropical bird community, estimating age-specific annual apparent survival probabilities for a set of passerine understory birds from the central Brazilian Amazon; and b) contribute to the debate about the latitudinal gradient in adult survival by comparing our adult survival estimates to estimates of temperate-zone adult survival probabilities. To estimate the age-specific survival we fit to our data a hierarchical multispecies Cormack-Jolly-Seber (CJS) model for n species, that treats species-specific parameters as random effects that are estimated and that describe the whole assemblage of species; for comparison of methods, we also fit a fixed-effects version of the model. To age birds we use the cycle-based WRP system. We introduce a novel variant of CJS model with a mixture component for the survival of birds of uncertain age at the time of banding. We found strong effect of age on survival, with juveniles surviving less than adults; evidence of latitude effect on survival, that supports the widely accepted hypothesis of variation on survival with latitude; and methodological differences between random and fixed effects model related to precision of estimates and scope of inference, that lead us to conclude that random-effects models are more appropriate for our analysis. We conclude that there is no reason for an alternative latitudinal trend in juvenile survival to account for the general trend in clutch size.

Graduation date: 2006 / I examined the avian biogeography of the islands of Lago Gatun, Panama, in an effort to better understand the effects of forest fragmentation in this biodiverse region, and specifically to understand the importance of fragment isolation and the mechanism behind its effects on tropical bird distribution. I combined exhaustive surveys of 29 islands with an experimental release program. Analyses of these data were conducted in a hierarchical fashion to evaluate evidence for avian dispersal limitation at multiple scales. First, I assessed the potential roles of area and isolation in determining avian species richness across this series of fragments, and how these relationships may differ for edge and forest dependent birds. Next, I analyzed community structure and species distributions to determine the relative contributions of island area and isolation in creating patterns of species-specific insular occurrence. Lastly, I assessed the results of the experimental investigation to evaluate the hypothesis that dispersal limitation explains the insular distribution patterns of several tropical forest birds. Isolation is a significant predictor of avian species richness, but only after accounting for the stronger effect of area. Species-isolation relationships are different in this archipelago for birds that rely on forest and edge habitats, respectively. Species-specific distributions are significantly nested when islands are ordered by area, and by isolation once the effect of area is considered. Occurrence of most forest species is sensitive to area and isolation of the archipelago. Examination of guild structure suggests that multiple mechanisms are responsible for these occurrence patterns. There are distinct species-specific differences in ability to cross small gaps, and species which are better able to cross these gaps are more widely distributed across the archipelago than those species that negotiate the same barriers poorly. Species that performed uniformly well in release experiments were much less likely to have suffered insular extinction in the preceding 25 years than those species that showed moderate to poor experimental performance. There is strong evidence of a morphological basis for the patterns. The cumulative evidence from these analyses is the most comprehensive evidence to date of local-scale dispersal limitation in volant birds.