Macroecology is the study of relationships between organisms and the environment at large spatial and temporal scales. This field of research examines patterns in species abundance, distribution and diversity. Understanding patterns in species distribution and richness can contribute significantly to our knowledge of community assembly and macroecological patterns, as well as to the effective conservation of threatened species and habitats. Although there have been a plethora of studies on birds in India over the years, there is a critical need to accurately delineate species distributions and understand patterns of richness. The focus of this study was to understand the factors (abiotic and biotic) that influence the distribution and composition of bird species in the Western Ghats, as well as to explore patterns in their geographic range sizes. The objectives of this study were addressed at the scale of the entire Western Ghats using a combination of field surveys, secondary data collection and species distribution modeling. The specific approaches to address these questions and the findings are outlined below. Chapter 2: Bird species in the Western Ghats – Patterns in composition and richness Fine-scale data on species presence and abundance are essential for exploring patterns in species distribution and richness. Despite the fact that birds have been extensively studied in the Western Ghats, systematic data collection and compilation of information over the entire mountain range has not been carried out, especially for the purpose of testing macroecological questions. This chapter describes patterns in bird species presence, abundance, composition and richness within the Western Ghats. The study area, site selection protocol and the sampling technique have also been described in detail. This dataset establishes a baseline of information about birds in the Western Ghats and subsets of this larger dataset will be used to address various questions in the following chapters. Chapter 3: Predicting bird species distribution in the Western Ghats Detailed knowledge of species’ ecological and geographical distributions is fundamental for conservation, as well as for understanding ecological and evolutionary determinants of spatial patterns of biodiversity. However, occurrence data for a vast majority of species are sparse, resulting in information about species distributions that is inadequate for many purposes. Species distribution models attempt to provide detailed predictions of distributions by relating presence or abundance of species to environmental predictors. In this chapter, we describe the usage of Maxent, a species distribution modelling technique based on presence-only data, to predict the distributions of bird species within the Western Ghats. For this purpose, we put together primary locations of bird species presence along with a published dataset. Using a number of important environmental layers, predicted species distribution maps were derived for 98 bird species, including 13 endemics, in the Western Ghats. Additionally, we calculated predicted range sizes for each of these species and obtained percentage contributions of important environmental predictors to each species’ distribution. This is the first study to develop species distribution models for bird species within the Western Ghats. Chapter 4: Patterns of range size among bird species Understanding large-scale patterns of variation in species geographic range size is fundamental to questions in macroecology and conservation biology. In general, range is believed to be influenced by a combination of environmental factors, evolutionary history and biotic interactions, mediated by species specific traits. These patterns need to be examined even for well-studied taxa like birds, especially within biodiversity hotspots faced by persistent degradation due to anthropogenic activities such as the Western Ghats. In this chapter, we use a dataset of 98 bird species within the Western Ghats to examine trends in range sizes, measured as latitudinal extent of occurrence and predicted range size from species distribution models. We show a significant relationship between latitude and range size for these bird species, supporting Rapoport’s rule. As far as we know, this relationship has never been tested at such low latitudes for birds. We also find that species traits such as body size, mean abundance and diet do not seem to show any discernable effect on patterns of range size. Additionally, we found that widely-used bird species range maps (in this case, from BirdLife International) are inaccurate representations of species ranges in comparison to the predicted species distribution maps that were derived in the previous chapter. We quantitatively demonstrated that these expert-drawn maps need to re-evaluated, especially since they are used to make conservation decisions. This is the first study to quantify species range sizes of birds within the Western Ghats and assess such range maps that are used to determine conservation status of species. Chapter 5: Environmental predictors of bird species distribution One of the major goals in ecology is to understand patterns and processes that determine species diversity. The drivers of global species richness gradients have been studied, especially in the case of birds, in terms of contemporary and historical factors. Such broad scale processes may not always reflect the processes affecting richness and distribution at smaller scales. Therefore, understanding the factors that influence individual species distributions is the first step towards this larger goal. In this chapter, we examined the environmental predictors that contributed to the predicted distribution of bird species observed in the Western Ghats, using the variable importance contribution values derived in Chapter 3. We found that a large proportion of the 98 bird species studied were influenced by normalized differential vegetation index, annual precipitation and elevation. The predictors did not differ among birds of different diet guilds and body size classes. Using Prinicipal components analysis, we observed that all 98 bird species are spread out across the environmental ordination space depicted by the PC axes 1 and 2. These axes are governed by measures of habitat heterogeneity and water-energy related variables, consistent with other tropical studies. The insectivorous guild seemed to occupy a variety of environmental niches across this space and other guilds seemed to be nested within the insectivorous guild. Similarly, larger sized birds were spread across the entire environmental ordination space, with species of smaller sizes nested within. This is the first step in trying to understand environmental predictors acting on birds in the Western Ghats. Further detailed studies need to be carried out to come to definite conclusions. Chapter 6: Relative roles of floristics and vegetation structure on bird species composition On the basis of the hierarchical model of habitat selection, it is known that birds select suitable habitats based on vegetation structure (physiognomy) at coarse biogeographic scales, and plant species composition (floristics) at more local scales. This chapter examines the relative influence of tree species composition and vegetation structure on bird species composition in the Western Ghats. These relationships were specifically assessed across the entire Western Ghats, within regions of the Western Ghats as well as within specific forest types. We found that floristics had a strong association with bird species composition across the Western Ghats and within evergreen and mixed deciduous habitat types. This association seems to be independent of the structural variation in the region. There was a decrease in association strength from the southern to the northern Western Ghats, in terms of both floristics and structure. We did not find an association between vegetation structure and insectivore composition, whereas phytophage composition did show a stronger association with floristics than structure. This is the first study at the scale of the entire Western Ghats to test the relative roles of floristics and vegetation structure. Taken as a whole, this dissertation examines large-scale macroecological questions regarding species distribution, range size and patterns of composition using primary data at the scale of the Western Ghats. The findings of this study have established a foundation that will help further our understanding of species distribution and richness in the Western Ghats, and aid in the decision making for conservation strategies in the future.

Em diversos estudos ao redor do globo, a estrutura e heterogeneidade da vegetação têm se mostrado um fator determinante na diversidade de espécies de aves e também de outros grupos de animais. O Cerrado é o segundo mais extenso e mais ameaçado bioma de ocorrência no Brasil. Este bioma também é caracterizado por um evidente gradiente ambiental de estrutura e heterogeneidade de vegetação. Na presente tese analisamos a influência da estrutura e heterogeneidade da vegetação sobre a diversidade em comunidades de aves do Cerrado. Nossa expectativa era corroborar a “Hipótese de Heterogeneidade de Habitats”, que propõe que quanto maior a estrutura e heterogeneidade da vegetação, maior será a diversidade de espécies. No primeiro capítulo, realizamos uma compilação sistemática de estudos publicados sobre a diversidade de aves em áreas ocupadas por algumas fisionomias típicas de Cerrado lato sensu, com o intuito de analisar o conhecimento obtido até então acerca da relação entre diversidade de aves e a estrutura da vegetação no Cerrado. Foram selecionadas 72 amostras de 22 estudos, sendo que estas amostras variaram quanto ao tipo fisionomia amostrada e o método amostral empregado, além de também estarem disponíveis em diferentes artigos e serem realizadas em diferentes regiões geográficas. Para análises destes dados, utilizamos a análise de modelos lineares generalizados de efeitos mistos (modelo com distribuição de erros poisson), que permite analisar os efeitos de variáveis fixas e aleatórias sobre a variável explicativa (riqueza de espécies). As variáveis fixas foram o tipo de vegetação amostrada (vegetação campestre, savânica e florestal) e o método amostral empregado (ponto fixo, transecto e redes de neblina). Já as variáveis de efeito aleatório utilizadas foram o estudo onde os dados foram publicados, o autor de cada estudo e a localidade geográfica. O efeito destas variáveis aleatórias poderiam afetar somente os interceptos das relações entre as variáveis fixas e a variável explicativa ou poderiam alterar a relação entre as variáveis fixas e explicativa. Construímos diversos modelos a partir da combinação de variáveis de efeito fixo e aleatório e a seleção do modelo mais parcimonioso foi feito por meio do critério AICc (critério de informação de Akaike corrigido para pequenas amostras). O modelo que apresentou menor valor de AICc (mais parcimonioso) foi aquele que incluiu os efeitos de ambas variáveis de efeito fixo (fisionomia e método amostral) e também um efeito da interação entre estas duas variáveis. Neste modelo também foram incluídos os efeitos das variáveis aleatórias estudo e localidade geográfica sobre os interceptos das relações entre as variáveis de efeito fixo e a variável explicativa. Estes resultados mostraram que a riqueza de espécies de aves em nosso estudo variou não só em função da fisionomia e do método amostral empregado, mas dependendo do método amostral utilizado a relação entre riqueza e fisionomia também foi alterada. Portanto, esta interação não permitiu que fosse estimada a relação entre fisionomia e riqueza sem considerar o efeito dos métodos. Já os efeitos das variáveis aleatórias mostraram que a variação estimada nos interceptos entre estudos foi duas vezes maior do que a variação estimada entre localidades geográficas. O efeito da interação entre as variáveis fisionomia e método amostral apontou para a existência de heterogeneidade de detecção entre locais com diferentes fisionomias, além também de um efeito das fisionomias na efetividade dos diferentes métodos amostrais. A influência dos métodos amostrais no número de espécies observadas em cada fisonomia pode ser esperada devido às diferenças intrínsecas dos métodos, já que ponto fixo e transecto são baseados em contatos visuais e auditivos com as espécies, enquanto que o método de rede de neblina consiste na captura passiva das espécies que voam na altura das redes. Assim, redes de neblina podem ser mais efetivas em habitats menos estruturados (por ex. campos limpos e sujos), onde a rede alcança quase todo os estratos de vegetação. No entanto, o método de transecto pode ser mais efetivo que o método de ponto fixo em áreas de florestas, pois nestes hábitats as espécies tendem a ter territórios menores e o deslocamento do observador proporciona ao observador cobrir um maior número de terrítórios. Por outro lado, o ponto fixo pode ser mais vantajoso por não produzir ruído e afugentar as espécies, o que pode ser uma desvantagem do método de transecto. Outros fatores, como a experiência e número de observadores, número de pontos amostrais, número de redes utilizadas e comprimento de transectos, podem explicar a grande variação estimada entre os estudos. Uma das maneiras de se contornar estes efeitos metodológicos é utilizar métodos desenvolvidos especialmente para lidar com diferentes probabilidades de detecção entre espécies, entre sítios e até métodos amostrais, o que poderia render dados mais confiáveis para o estudo da ecologia das espécies e para a elaboração de planos de manejo e/ou conservação. No segundo capítulo, a relação entre diversidade de aves e estrutura da vegetação foi analisada a partir de dados coletados em campo e utilizando um protocolo de amostragem específico para se estimar e considerar os efeitos da vegetação sobre a detecção das espécies. As amostragens foram realizadas em um dos maiores e mais preservados remanescentes de Cerrado (Parque Nacional Grande Sertão Veredas-PARNA GSV) e consistiram do registro das espécies de aves em 32 áreas dispostas em um gradiente de vegetação de Cerrado, que variaram desde campos limpos e sujos, campos cerrado a cerrados sensu stricto. O cálculo da riqueza de espécies de aves em cada sítio foi realizado através de modelos de ocupação-detecção, adaptados para estimar a riqueza de espécies em comunidades. A vegetação, por sua vez, foi medida a partir de estimativas de presença da vegetação entre 0 e 4 m de altura (16 intervalos de 22,5 cm cada um) e duas variáveis de estrutura foram obtidas a partir de uma análise de componentes principais, que foi aplicada para resumir a variação da presença de vegetação nos 16 intervalos de altura. Estas variáveis de vegetação foram relacionadas tanto com a ocupação quanto com a detecção das espécies, já que a estrutura da vegetação poderia influenciar não só a ocorrência mas também a detecção das espécies. O dia da amostragem e também a temperatura no momento da amostragem também foram incluídas como covariáveis que poderiam afetar a detecção. Após a estimativa da riqueza de espécies pelo modelo de ocupação-detecção para comunidades, esta riqueza estimada foi relacionada por uma função quadrática com a estrutura da vegetação usando um modelo bayesiano de metanálise, que permitiu incluir a incerteza nas estimativas de riqueza na análise. A título de comparação, também foi ajustado um modelo quadrático GLM (distribuição de erros normal) aos dados de riqueza observada. Os resultados mostraram que a riqueza estimada a partir dos dados das 38 espécies mais detectadas durante as amostragens teve uma fraca relação com as duas covariáveis de estrutura de vegetação, sendo que houve uma maior riqueza de espécies em sítios com vegetação intermediária em altura e uma maior riqueza de espécies de aves em sítios onde houve maior presença de vegetação abaixo de 2 m de altura. No entanto, as relações entre riqueza estimada e estas covariáveis foi menos intensa mas qualitativamente similar às relações entre a riqueza observada e as covariáveis de vegetação. A menor intensidade nas relações da riqueza estimada foi evidenciada principalmente em ambos os extremos do gradiente de estrutura vertical da vegetação e também nas áreas com menor presença de vegetação abaixo de 2 m. Estes resultados mostraram que o efeito da detecção pode alterar o efeito da relação entre riqueza de espécies e estrutura de vegetação. Além disso, ao menos para as 38 espécies mais comumente encontradas na área de estudo, os resultados apontam para a importância de todo o gradiente de estrutura da vegetação para a manutenção da riqueza de espécies de aves no Cerrado. Futuros estudos que visem aprimorar o uso destes modelos de ocupação e detecção para comunidades são fundamentais para permitir o uso dos dados de todas as espécies da comunidade. Além disto, outros estudos que se proponham a analisar a dinâmica e composição das comunidades de aves nestes gradientes de estrutura de vegetação são fundamentais para um maior conhecimento sobre a ecologia e conservação das aves no Cerrado / In several studies around the globe, the structure and diversity of vegetation have been shown to be a determining factor in the diversity of species of birds and also other groups of animals. The Cerrado is the second most extensive and most threatened biome occurrence in Brazil. This biome is also characterized by an obvious environmental gradient of vegetation structure and heterogeneity. In this thesis we analysed the influence of the structure and diversity of the vegetation on the diversity in the Cerrado bird communities. Our expectation was to support the “Habitat Heterogeneity Hypothesis” which suggests that the higher the structure and diversity of vegetation, the greater the diversity of species. In the first chapter, we conducted a systematic compilation of published studies on the diversity of birds in areas occupied by some typical physiognomy of Cerrado textit lato sensu, in order to analyze the knowledge obtained so far about the relationship between diversity of birds and the structure of the vegetation in the Cerrado. We selected 72 samples from 22 studies, and these samples varied as the sampled vegetation physiognomy, the sampling method used, and they also are available in different articles and be carried out in different geographical regions. We performed the analysis of generalized linear mixed effects models (model poisson distribution errors), which allows us to analyse the effects of fixed and random variables on the explanatory variable (species richness). Fixed variables were the type of sampled vegetation (grassland, savanna and forest) and the sample method employed (fixed point, transect and mist nets). The random variables used were the study where the data were published, the author of each study and geographic location. These random variables could only affect the intercepts of the relationship between fixed and variable explanatory variable or could alter the relationship between fixed and explanatory variables. We built several models from the combination of fixed and random effects variables and selection the most parsimonious model was made by the AIC criterion (Akaike information criterion corrected for small samples). The model that showed lower value of AIC (more parsimonious) was the one that included the effects of both fixed effect variables (physiognomy and sampling method) and also an effect of the interaction between these two variables. In this model were also included the effects of random variables study and geographic location of the intercepts of the relationship between the fixed effect variables and the explanatory variable. These results showed that the bird species richness in our study varied not only in terms of physiognomy and sample method, but depending on the sampling method used the relationship between richness and physiognomy has also changed. Therefore, this interaction does not allowed us to estimate the relationship between physiognomy and richness without considering the effect of the methods. Since the effects of random variables showed that the variation in the estimated intercept between studies was twice larger than the estimated variation between geographic locations. The effect of interaction between the vegetation physiognomy and sampling method variables pointed to the existence of heterogeneity detection between locations with different physiognomies, in addition also of an effect of the physiognomies in the effectiveness of different sampling methods. The influence of the sampling method in the number of species observed in each physiognomy may be expected due to intrinsic differences in the methods, since fixed point counts and transect are based on visual and aural contacts with the species, while the mist net method consists in passive capture of species flying at the time of the networks. Thus, mist nets may be more effective in less structured environments (eg. Clean and dirty fields) where the net reaches virtually all vegetation layers. However, transect method can be more effective than the fixed point method in areas of forests since in these habitats species tend to have smaller territory areas, and the observer movement provides the observer cover greater areas. On the other hand, the point counts can be more advantageous not to produce noise and chase species, which may be a disadvantage of transect method. Other factors, such as experience and number of observers, the number of sampling points, the number of nets used and length of transects, may explain the wide variation between studies estimated. One of the ways to overcome these methodological effects is to use methods developed especially to deal with different probabilities of detection of species, between sites and sampling methods, which could yield more reliable data for the ecological study of the species and the development of management plans and/or conservation. In the second chapter, the relationship between diversity of birds and vegetation structure was analysed from data collected in the field and using a specific sampling protocol to estimate and consider the effects of vegetation on the detection of species. The samples were taken in one of the largest and well preserved remnants of Cerrado (Grande Sertão Veredas National Park-PARNA GSV) and consisted of the record of bird speciesin 32 areas arranged in a Cerrado vegetation gradient, ranging from grasslands, open and dense savannas. The calculation of the bird species richness at each site was conducted using occupancy-detection models adapted to estimate the number of species in communities. The vegetation, in turn, was measured from estimates of the presence of vegetation in height intervals between 0 and 4 m (16 intervals of 22.5 cm each) and two structure variables were obtained from a principal component analysis applied to summarize the variation of the vegetation presence in height intervals. These vegetation variables were related to both the occupation and detection of species, since the vegetation structure could influence not only the occurrence but also the detection of species. The day of sampling and also the temperature at the time of sampling were also included as covariates that may a_ect the detection. After the estimation of species richness by model occupancy detection for communities, this estimated richness was related by a quadratic function with the vegetation structure using a Bayesian meta-analysis model, which allowed us include uncertainty in richness estimates. By way of comparison, we also fit a quadratic model GLM (normal distribution errors) to the observed richness data. The results showed that the richness estimated from the data of the 38 most detected species during sampling had a weak relationship with both covariates vegetation structure, and there was a greater number of species at sites with intermediate vegetation height and greater bird species richness in places where there was a greater presence of vegetation below 2 m in height. However, relations between estimated richness and these covariates was less intense but qualitatively similar to the relationship between observed richness and vegetation covariates. The lowest intensity in the estimated richness relationship was observed mainly at both ends of the vertical gradient of vegetation and also in areas with less presence of vegetation below 2 m. These results showed that the effect of detection can change the effect of the relationship between species richness and vegetation structure. Moreover, at least for the 38 species most commonly found in the study area, the results point to the importance of the entire vegetation structure gradient to maintain the bird species richness in Cerrado. Future studies aiming to improve the use of these models of occupation and detection for communities are essential to allow the use of data of all species in the community. In addition, other studies that propose to analyse the dynamics and composition of bird communities in these vegetation structure gradients are fundamental for a better understanding of the ecology and conservation of Cerrado birds

Les nombreuses menaces qui pèsent sur le milieu marin et les preuves d’une perte de biodiversité globale au cours des dernières décennies ont rendu indispensable la mise en place de mesures de conservation dans les années à venir. Au sein des Terres Australes et Antarctiques Françaises (TAAFs) en particulier, qui offrent une biodiversité exceptionnelle et sont le cadre d’activités anthropiques intensives à travers les grandes pêcheries industrielles, la délimitation d’Aires Marines Protégées (AMP) serait extrêmement bénéfique pour la conservation du milieu. Pour définir les zones d’importance pour la biodiversité, qui représenteraient de bonnes candidates pour un statut d’AMP, l’utilisation des données de distribution de prédateurs supérieurs apparait idéale. En effet, leur position dans le réseau trophique en fait de bons intégrateurs des niveaux inférieurs, et de plus, ils sont facilement observables lors des campagnes d’observations et offrent des facilités pour l’équipement de dispositifs télémétriques puisqu’ils reviennent régulièrement à terre pendant la période de reproduction. Grâce à l’analyse spatiale des données de distribution obtenues pour un grand nombre de prédateurs supérieurs au sein d’écosystèmes subantarctiques aussi bien qu’en milieu tropical et grâce au développement de modèles d’habitat, nous avons ici déterminé des ‘hotspots’ de biodiversité pour les prédateurs supérieurs des TAAFs. Ces travaux pourraient ainsi servir de base aux propositions de limites pour de potentielles futures AMPs. / The numerous threats that marine environment face, coupled with the evidence for a global biodiversity loss during last decades have lead to an increasing need for setting up conservation measures. Particularly, delimiting Marine Protected Areas (MPAs) within the French Southern Territories, which are home to an exceptional biodiversity and extensive human activities through industrial fisheries, would be extremely beneficial for species conservation. In order to identify areas of ecological significance for biodiversity, which could be good MPA candidates, the use of distribution data from top marine predators seems to be ideal. Thanks to their high position in trophic network, they are likely to integrate lowers trophic level species distribution. Moreover, they are easy to observe trough at-sea observation campaigns, and easy to equip with telemetric devices thank to their central place foraging that lead them to return regularly to their colony during breeding season. Here, we analyse distribution data and develop habitat models from several top predators species in both subantarctic and tropical regions to highlight biodiversity hotspots within the French Southern Territories. This work could thus be use as a basis to define potential boundaries for a future MPA.