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Biodiversity is determined by a myriad of complex processes acting at different scales. Given the current rates of biodiversity loss and change, it is of paramount importance that we improve our understanding of the underlying structure of ecological communities. In this thesis, I focused on Species Abundance Distributions (SAD), as a synthetic measure of biodiversity and community structure, and on Beta (β) diversity patterns, as a description of the spatial variation of species composition. I systematically assessed the effect of scale on both these patterns, analysing a broad range of community data, including different taxa and habitats, from the terrestrial, marine and freshwater realms. Knowledge of the scaling properties of abundance and compositional patterns must be fully integrated in biodiversity research if we are to understand biodiversity and the processes underpinning it, from local to global scales. SADs depict the relative abundance of the species present in a community. Although typically described by unimodal logseries or lognormal distributions, empirical SADs can also exhibit multiple modes. However, the existence of multiple modes in SADs has largely been overlooked, assumed to be due to sampling errors or a rare pattern. Thus, we do not know how prevalent multimodality is, nor do we have an understanding of the factors leading to this pattern. Here, I provided the first global empirical assessment of the prevalence of multimodality across a wide range of taxa, habitats and spatial extents. I employed an improved method combining two model selection tools, and (conservatively) estimated that ~15% of the communities were multimodal with strong support. Furthermore, I showed that the pattern is more common for communities at broader spatial scales and with greater taxonomic diversity (i.e. more phylogenetically diverse communities, since taxonomic diversity was measured as number of families). This suggests a link between multimodality and ecological heterogeneity, broadly defined to incorporate the spatial, environmental, taxonomic and functional variability of ecological systems. Empirical understanding of how spatial scale affects SAD shape is still lacking. Here, I established a gradient in spatial scale spanning several orders of magnitude by decomposing the total extent of several datasets into smaller subsets. I performed an exploratory analysis of how SAD shape is affected by area sampled, species richness, total abundance and taxonomic diversity. Clear shifts in SAD shape can provide information about relevant ecological and spatial mechanisms affecting community structure. There was a clear effect of area, species richness and taxonomic diversity in determining SAD shape, while total abundance did not exhibit any directional effect. The results supported the findings of the previous analysis, with a higher prevalence of multimodal SADs for larger areas and for more taxonomically diverse communities, while also suggesting that species spatial aggregation patterns can be linked to SAD shape. On the other hand, there was a systematic departure from the predictions of two important macroecological theories for SAD across scales, specifically regarding logseries distributions being selected only for smaller scales and when species richness and number of families were proportionally much smaller than the total extent. β diversity quantifies the variation in species composition between sites. Although a fundamental component of biodiversity, its spatial scaling properties are still poorly understood. Here, I tested if two conceptual types of β diversity showed systematic variation with scale, while also explicitly accounting for the two β diversity components, turnover and nestedness (species replacement vs species richness differences). I provided the first empirical analysis of β diversity scaling patterns for different taxa, revealing remarkably consistent scaling curves. Total β diversity and turnover exhibit a power law decay with log area, while nestedness is largely insensitive to scale changes. For the distance decay of similarity analysis, while area sampled affected the overall dissimilarity values, rates of similarity were consistent across large variations in sampled area. Finally, in both these analyses, turnover was the main contributor to compositional change. These results suggest that species are spatially aggregated across spatial scales (from local to regional scales), while also illustrating that substantial change in community structure might occur, despite species richness remaining relatively stable. This systematic and comprehensive analysis of SAD and community similarity patterns highlighted spatial scale, ecological heterogeneity and species spatial aggregation patterns as critical components underlying the results found. This work expanded the range of scales at which both theories deriving SAD and community similarity studies have been developed and tested (from local plots to continents). The results here showed strong departures from two important macroecological theories for SAD at different scales. In addition, the overall findings in this thesis clearly indicate that unified theories of biodiversity (or assuming a set of synthetic minimal assumptions) are unable to accommodate the variability in SADs shape across spatial scales reported here, and cannot fully reproduce community similarity patterns across scales. Incorporating more realistic assumptions, or imposing scale dependent assumptions, may prove to be a fruitful avenue for ecological research regarding the scaling properties of SAD and community similarity patterns. This will allow deriving new predictions and improving the ability of theoretical models to incorporate the variability in abundance and similarity patterns across scales.

Coordenação de Aperfeiçoamento de Pessoal de Nível Superior / Right after the arising of the Panama isthmus, the family Canidae colonized South America, around 2.6-2.4 million years ago. Although canids radiation in South America is recent, this region holds the largest canid diversity in the world, with more than 10 extant species. This great diversity is also notable when dealing with the ecomorphological variation of these animals. The maned-wolf (Chrysocyon brachyurus), a large omnivore, and the bush-dog (Speothos venaticus), a small hypercarnivore, are the extreme exemples of this variation. Such shape diversity can be potentially explained by both abiotic, such as climate, and biotic factors, like competition. These factors may have contributed for stabilizing the south-american canids community. Thus, this study aimed to investigate what drives this ecomorphological amplitude, as well as to understand how can similar species coexist when overlapping their distributional ranges, such as the crab-eating fox (Cerdocyon thous), which has sympatric areas with Lycalopex vetulus and L. gymnocercus, two foxes ecologically similar to Cerdocyon thous. To answer these questions, 431 especies were photographed in nine South-American museums. Through geometric morphometric procedures, it was possible to quantify the phenotypic variation of eight canid species (Atelocynus microtis, C. thous, C. brachyurus, L. culpaeus, L. griseus, L. gymnocercus, L. vetulus e S. venaticus) throughout their geographical range and then test the contribution of biotic and abiotic factors driving this variation. The evidences presented in this study suggest C. thous alters its shape and body size when in sympatry with two Lycalopex species, pattern described by character displacement , which is when similar species shift their phenotype in order to minimize competition. Besides, C. thous also follows the Bergmann s rule, which predicts that body size increases at larger latitudes. When considering the canid community as a whole, climate was identified as the main factor contributing to phenotypic variation in these animals. Competition has a weaker impact in south-american canids skull morphology, although it may have played a larger role in the past, when the ecomorphological diversity in the subcontinent was even larger. / Logo após a emergência completa do istmo do Panamá, a família Canidae colonizou a América do Sul, há, aproximadamente, 2.6-2.4 milhões de anos atrás. Embora a radiação dos canídeos seja recente na América do Sul, esta região compõe a maior diversidade atual de espécies desta família no mundo, com mais de 10 espécies viventes. Esta grande biodiversidade também é notável na variação ecomorfológica destes animais. Exemplo disso são os dois extremos desta variação: o lobo-guará (Chrysocyon thous), um animal onívoro de grande tamanho corporal; e o cachorro-vinagre (Speothos venaticus), espécie hipercarnívora de pequeno porte. Tamanha diversidade de formas pode potencialmente ser explicada tanto por fatores abióticos, como o clima, e bióticos, como a competição. Ambos os fatores podem ter contribuído para a estabilização da comunidade de canídeos sul-americana. Por isso, este estudo se propôs a investigar o que impulsionou esta amplitude ecomorfológica, bem como entender como espécies mais similares coexistem quando em sobreposição distribucional, como o graxaim-do-mato (Cerdocyon thous), que possui área simpátrica a Lycalopex vetulus e L. gymnocercus, duas raposas ecologicamente similares a Cerdocyon thous. Para tanto, 431 espécimens foram fotografados em nove museus da América do Sul. Através de procedimentos de morfometria geométrica, foi possível quantificar a variação fenotípica de oito espécies de canídeos (Atelocynus microtis, C. thous, C. brachyurus, L. culpaeus, L. griseus, L. gymnocercus, L. vetulus e S. venaticus), ao longo de sua distribuição geográfica, e então testar a contribuição dos fatores bióticos e abióticos nesta variação. As evidências deste estudo sugerem que C. thous altera sua forma e tamanho corporal quando em simpatria com as duas espécies de Lycalopex, padrão descrito pelo deslocamento de caráter , quando espécies semelhantes alteram seu fenótipo em simpatria, a fim de minimizar a competição. Além disso, C. thous também segue a regra de Bergmann, que prevê um aumento de tamanho corporal com o aumento da latitude. Ao considerar a comunidade de canídeos como um todo, o clima foi identificado como o fator que mais contribuiu para a variação fenotípica destes animais. A competição, por sua vez, tem um impacto mais fraco na morfologia do crânio dos canídeos sul-americanos, embora possa ter tido uma maior contribuição no passado, quando a diversidade ecomorfológica no subcontinente era ainda maior.

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.

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No. of bitstreams: 2 Tese - Lucas Lacerda Caldas Zanini Jardim - 2018.pdf: 5066072 bytes, checksum: 4280b5b19a9111a59fea8065049fd5b3 (MD5) license_rdf: 0 bytes, checksum: d41d8cd98f00b204e9800998ecf8427e (MD5) Previous issue date: 2018-04-27 / Coordenação de Aperfeiçoamento de Pessoal de Nível Superior - CAPES / Macroecology studies ecological pattern at large geographical and temporal scales. At these scales, information about hundreds or even thousands of studied species. This lack of information may potentially bias studies’ conclusions related with macroecological processes and patterns. In this thesis, we evaluated phylogenetic imputation methods, their uses and effects in macroecological studies. The first chapter evaluated different methods used to deal with missing data, taking into account different scenarios of species trait evolution, as well as percentage and pattern of missing data. We found that dealing with missing data relies on the specific goals and data of the study. Therefore, we suggested caution while using imputed database. In the second chapter, we tested the island rule effect in body mass and brain volume of primates. To do so, we fitted evolutionary models to those traits and then imputed the body mass and brain volume for Homo floresiensis. We concluded that primates do not follow the island rule and even though our models overestimated, on average, brain and body size of Homo floresiensis, its evolution did not deviate from primates’ evolutionary expectation. Lastly, in the third chapter, we tested existence of Bergmann’s rule in mammals using multiple imputation methods, in addition to considering the consequences of ignoring missing data while testing the rule. We found that ignoring missing data can invert (eg. changing from positive to negative effect) the effect of temperature on body mass, but this bias did not turn the effect statistically significant. Therefore, we concluded that mammals do not follow Bergmann’s rule, when evaluated at the class taxonomic level. Finally, this thesis discussed pros, cons and future research avenues in order to make phylogenetic imputation a more robust tool to deal with missing data in macroecology. / A macroecologia estuda padrões ecológicos em grandes escalas geográficas e temporais, em busca de quais processos moldam esses padrões. Nessas escalas de estudo, há raramente informações completas sobre as centenas ou até milhares de espécies estudadas. Essa ausência de informações tem o potencial de enviesar as conclusões dos estudos sobre padrões e processos macroecológicos. Nessa tese, nós avaliamos métodos de imputação filogenética, a sua aplicação e consequências em estudos macroecológicos. Para avaliar potenciais vieses do uso de banco de dados imputados, no primeiro capítulo, nós aplicamos diferentes métodos utilizados para tratar dados faltantes, sob diferentes cenários de evolução dos atributos das espécies, porcentagem e padrão dos dados faltantes. Nós encontramos que a forma de tratar o dado faltante pode ser dependente dos objetivos e dos dados de cada estudo e, portanto, nós sugerimos cautela ao utilizarmos bancos de dados imputados. No segundo capítulo, nós testamos o efeito da regra de ilha na evolução da massa corpórea e do volume cerebral de primatas. A partir dos melhores modelos evolutivos ajustados a esses atributos, nós imputamos a massa corpórea e volume cerebral de Homo floresiensis. Nós concluímos que primatas não seguem regra de ilha e que apesar de nossos modelos superestimarem, em média, o tamanho do corpo e cérebro de Homo floresiensis, a sua evolução não se desvia do esperado pela evolução de primatas. Por fim, no terceiro capítulo testamos a regra de Bergmann em mamíferos, utilizando métodos de imputação múltipla e avaliamos as consequências de desconsiderar os dados faltantes na detecção da regra. Nós encontramos que testar a regra sem considerar os dados faltantes pode inverter o efeito da temperatura na massa do corpo, mas esse viés não tornou o efeito estatisticamente significante. Portanto, concluímos que mamíferos não seguem a regra de Bergmann, quando toda a classe é avaliada. Por fim, essa tese discutiu vantagens, desvantagens e futuras linhas de pesquisa para tornar a imputação filogenética uma ferramenta mais robusta para tratarmos dados faltantes em macroecologia.

La thématique centrale de cette thèse, située à l’interface entre macroécologie et macroévolution, consiste en l’analyse des dynamiques spatio-temporelles de distribution des ammonites du Pliensbachien inf. de la Téthys de l’Ouest. Deux problématiques sont abordées : comment la similarité compositionnelle des assemblages décroît-elle avec la distance géographique? Quel est le lien entre la taille des aires de répartition des espèces et leur phylogénie, leur durée de vie, et leur position latitudinale?Grâce à une analyse des relations de SDD, nous montrons que plus l’environnement est homogène, plus les ammonites se dispersent librement ; et que leur dispersion sur de grandes distances n’est pas liée à leur morphologie, ce qui suggère qu’elle se faisait essentiellement via une dérive planctonique aux cours des stades juvéniles.Notre étude de l’étendue de la répartition des ammonites révèle que cette caractéristique n’est ni stable, ni propre à un clade. Son héritabilité phylogénétique dépend de la stabilité spatio-temporelle de l’environnement et varie selon l’estimateur de répartition et l’intervalle de temps considérés, et entre des espèces d’un même groupe contemporaines les unes des autres. De plus, l’étendue de répartition n’a pas d’effet sur la durée de vie des espèces. Cette absence de corrélation semble prévaloir chez les ammonoïdes, ce groupe étant connu pour présenter de grandes aires de répartition associées à des courtes durées de vie. Enfin, les étendues latitudinales des ammonites étudiées tendent à être maximales aux basses latitudes et augmentent des moyennes vers les hautes latitudes, conformément à la Règle de Rapoport. Ceci suggère qu’elles étaient vraisemblablement température-dépendantes et qu’un gradient de température des eaux existait au Pliensbachien inf. dans la province NW européenne. / The present work, which lies between macroecology and macroevolution, focuses on the spatio-temporal dynamics of early Pliensbachian ammonites of the western Tethys. Two main questions are addressed: how does inter-assemblage compositional similarity decrease with geographical distance? What is the relationship between species range size and phylogeny, species duration and latitudinal position?Ammonite dispersal dynamics are studied through a multi-scale Similarity Distance Decay analysis. We show that their long-distance dispersal is facilitated when the environment is more homogeneous and that it is not related to their morphology. This suggests that their long-distance dispersal occurred essentially through a passive planktonic drift during the first juvenile stages.We also show that ammonite geographical range size is neither stable, nor clade-specific. Indeed, its phylogenetic heritability depends on the spatio-temporal stability of the environment and it varies according to the range variable chosen, the temporal interval considered, and among contemporary species of the same group. Furthermore, the size of their geographical range has no effect on species duration. In fact, this lack of correlation seems to prevail among ammonoids, as this group is well-known for showing both large geographical distributions and short species durations. Finally, species latitudinal ranges tend to be maximal at low latitudes and to increase from middle to high latitudes during the early Pliensbachian, in accordance with the Rapoport’s rule. This suggests that these ammonites were probably temperature-dependent and that a latitudinal gradient of sea-surface temperature occurred in the NW European province during the early Pliensbachian.

Each species has specific adaptations to its environment, and since environmental parameters reveal geographic trends, it is reasonable to expect the existence of geographic trends in species characteristics as well. The aim of this study has been to evaluate the effect of environmental conditions on geographic variability of functional traits of European birds. I have analysed the effect of temperature, precipitation, productivity, altitude and habitat type (forest, open habitats, bush, settlements, wetlands) on clutch size, number of clutches per breeding season, egg size, incubation length, age of maturity, body mass, wing, tail, bill and tarsus length. I have used data from the European breeding bird atlas, so that I have calculated mean values of all the traits for quadrats 50x50 km, and then related them to environmental characteristics using OLS and GLS. Clutch size increases with temperature, whereas the number of clutches decreases with it, indicating possible trade-off between clutch size and the number of clutches, whose result is determined by the length of breeding season. Egg size decreases with temperature, possibly due to higher survival of large eggs (and consequently juveniles) in cold regions. Incubation length increases with both temperature and environmental productivity,...

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