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Species richness of virtually all high-level taxonomic groups is strongly statistically related to climatic variables such as temperature and precipitation, and consistently so across space and time. These observations are consistent with a causal link between the number of species that occur in a given region and its climate. Although dozens of hypotheses have been proposed, the main mechanisms underlying this pattern remain largely unresolved. And, few ecological studies have attempted to identify regularities in the individual species distributions that make up the richness–climate relationship. Despite the complexities of species’ biologies, I found that, to a first approximation, species’ probability of occupancy at continental scales were generally well statistically explained by a Gaussian function of temperature and precipitation. This simple model appeared general among species, taxa and regions. However, although individual species’ ranges are strongly statistically related to climate, spatial variations in richness cannot be explained by systematic variations in species’ climatic niches. And, individual species track changes in climatic variables through time much more weakly than species richness tracks these changes, suggesting that richness is at least partly constrained by mechanisms independent of species identities. Moreover, at macro-scales, species richness was also not strongly predictable from the temperature at which clades have originated, from historical variability in climatic variables nor from local short-term extirpation rates. In sum, I rejected several prominent hypotheses aiming to explain richness–climate relationship and found several lines of evidence inconsistent with the common idea that climatic constraints on individual species, by themselves, can explain richness–climate relationship. I propose a mechanism to explain, as a first approximation, the continental biogeography of species distributions that relies on neutral processes of dispersal and local extinctions within species’ broad deterministic thermal tolerances.

The decline in biodiversity over the last decade has motivated researchers to investigate the relationship between species richness (biodiversity) and ecosystem function. Empirical approaches are becoming more realistic as more factors have been included. Spatial heterogeneity is an example. Heterogeneity is an inherent part of the environment and apparent in all habitat types creating a patchy, mosaic of natural landscape. Researchers have reported the extent of heterogeneity in the landscape, but surprisingly not yet included heterogeneity into biodiversity and ecosystem function (BEF) studies. In recent years, empirical studies of marine systems have enhanced the BEF debate. Depauperate estuarine systems are ideal candidates for establishing model systems. In this study, estuarine microphytobenthos (MPB) were used as a response variable since the relationship between MPB and primary productivity is well-known. This relationship was exploited to employ MPB biomass as a proxy for primary productivity. Benthic chambers were used to assess the effect of macrofauna in single species and multi-species treatments on both ecosystem function and net macrofaunal movement. Heterogeneity was created through enriching sediment ‘patches’ with Enteromorpha intestinalis, providing areas of high and low nutrient. Heterogeneity, macrofaunal biomass, species richness, species diversity and flow were all varied in order to assess combined effects on the functioning of the system. Heterogeneity was found to have a significant influence on ecosystem functioning and on macrofaunal movement, however, patch arrangement did not. MPB biomass was highest in patches containing organic enrichment suggesting that nutrients were obtained locally from the sediment/water interface rather than the water column. There was variation in MPB biomass with macrofaunal species, probably resulting from differences in behavioural traits. It was also evident that flow altered species behaviour, as there was a significant difference between static and flow treatments. This work shows the importance of heterogeneity for BEF relationships.

Animal phyla vary dramatically in species richness (from one species to >1.2 million), but the causes of this variation remain largely unknown. Animals have also evolved striking variation in morphology and ecology, including sessile marine taxa lacking heads, eyes, limbs, and complex organs (e.g., sponges), parasitic worms (e.g., nematodes, platyhelminths), and taxa with eyes, skeletons, limbs, and complex organs that dominate terrestrial ecosystems (arthropods, chordates). Relating this remarkable variation in traits to the diversification and richness of animal phyla is a fundamental yet unresolved problem in biology. Here, we test the impacts of 18 traits (including morphology, ecology, reproduction, and development) on diversification and richness of extant animal phyla. Using phylogenetic multiple regression, the best-fitting model includes five traits that explain approximate to 74% of the variation in diversification rates (dioecy, parasitism, eyes/photoreceptors, a skeleton, nonmarine habitat). However, a model including just three (skeleton, parasitism, habitat) explains nearly as much variation (approximate to 67%). Diversification rates then largely explain richness patterns. Our results also identify many striking traits that have surprisingly little impact on diversification (e.g., head, limbs, and complex circulatory and digestive systems). Overall, our results reveal the key factors that shape large-scale patterns of diversification and richness across >80% of all extant, described species.

Aquesta tesi examina els mecanismes ecològics i evolutius que mantenen els gradients de riquesa específica en ocells. S'examinen en primer lloc el patrons de diversitat de l'avifauna de Nord Amèrica. (capitol 1). Tot seguit es testen mecanismes a escala regional, tot estudiant el cas de l'avifauna de Catalunya (capitols 2 i 3). Finalment, es testen diferents mecanismes que regulen la diversitat a escala local en una comunitat d'ocells frugívors al Parc nacional de Doñana. El capitol 6 integra tots els anteriors i els relaciona tot fent una revisió bibliogràfica en profunditat.Paraules clau: biodiversitat, ocells, adaptació, filogènia, switching behaviour, xarxes d'interacció, Nord Amèrica, Catalunya, Doñana, gradients altitudinals, productivitat. / Species richness gradients have been analyzed during many decades and they have progressively emerged as a central topic in community ecology (Darwin 1859, Wallace 1878, Willis 1922, Dobzhansky 1950, Fisher 1960, Hillebrand 2004, Riclkefs 2004, Mittelbach et al. 2007, among others). Historically, species richness gradients have been analyzed from two main points of view: the ecological and the evolutionary perspective (Ricklefs 2004, 2006b). The ecological perspective assumes that populations are evolutionarily fixed and studies species richness gradients as the result of regional colonization and extinction processes, and ecological interactions (MacArthur and Wilson 1967b, Scheiner and Willig 2005). Contrarily, the evolutionary approach states that species richness gradients are the result of geographic differences in the long-term evolutionary processes of speciation and lineage extinction (Rohde 1978, 1992, Mittelbach et al. 2007). For instance, according to the evolutionary view, tropical regions achieve higher species numbers due to increased diversification rates in low latitude areas (i.e. increased speciation and/or reduced extinction rates). Interestingly enough, recent empirical evidence derived from phylogenetic studies suggests that bird diversification rates are effectively higher in the tropics thus providing empirical support for the evolutionary view (Cardillo 1999, Cardillo et al. 2005, Golberg et al. 2005, Ricklefs 2006a, Jablonski et al. 2006, Weir and Schluter 2007). In contrast with the evolutionary approach, the ecological approach assumes that populations are evolutionary stable units and focus on the effect of regional colonization-extinction dynamics, local conditions and local interactions in determining species richness gradients (Hutchinson 1959, MacArthur and Wilson 1963, 1967ab, MacArthur and Levins 1967, May 1975, Riclkefs 2006b). The ecological approach highlights that species richness gradients can be generated by ecological mechanisms independently of the evolutionary processes of speciation and lineage extinction that conform the regional pool of available species. For instance, an environmental gradient can cause a species richness gradient by limiting the number of successful colonizations in environmentally severe localities without the participation of any speciation process in the regional pool (Wright 1983, Hanski 1997, Boulinier et al. 1998).These two contrasting and complementary views, the evolutionary and ecological approaches, are progressively being merged in a unified framework (Ricklefs 2006b, Johnson and Stinchcombe 2007). For instance, Johnson and Stinchcombe have recently proposed that two general hypotheses (H1, H2) are underlying a new synthesis between community ecology and evolutionary biology. On one hand (H1), evolutionary processes explain present-day community patterns and the ecological dynamics of species interactions. For instance, extinction and colonization processes that generate species richness gradients might be shaped by species' adaptations, history and phylogenetic relationships (Wiens and Donoghue 2004, Wiens and Graham 2005, Kraft et al. 2007). On the other hand (H2), Johnson and Stinchcombe highlighted that both species interactions and community context strongly influence the direction, rate and outcome of present-day evolutionary processes (Benkman 1999, Thompson 2005). Overall, the emerging synthesis of community ecology and evolutionary ecology highlights that ecological and evolutionary views are profoundly interlinked. Here we applied an evolutionary and ecological synthetic perspective to the study of the generation and maintenance of species richness in bird communities (Johnson and Stinchcombe 2007). As a first preliminary step, we examined geographical patterns of bird species richness in North America in several functional groups and the associated environmental correlates at the continental scale (Chapter 1). Our results showed that global large-scale patterns of avian diversity in temperate regions were best viewed as the overlayed response of distinct species groups to diverse ecological factors. These results strongly suggested the convenience of choosing specific functional groups in order to examine specific macroecological evolutionary and ecological hypotheses. Consequently, in Chapter 2, we next chose forest birds as a study group for the analysis of macroecological patterns, because they are the largest group in Nearctic and Palaeoarctic regions among terrestrial birds, and hold strong species-energy relationships (Mönkkönnen et al. 2006). Regional data with colonization and extinction estimates for forest birds were available in Catalonia along an altitudinal gradient (Estrada et al. 2004) providing an exceptional dataset to study the ecological and evolutionary processes behind species richness gradients (Chapters 2 and 3). We examined the altitudinal species richness gradient in Catalonia from an evolutionary and ecological synthetic perspective. First, the role of evolutionary processes in the gradient was assessed studying changes in community phylogenetic structure along the altitudinal gradient. Secondly, the role of ecological processes was examined by studying colonization and extinction dynamics at an ecological time-scale (20 years). These analyses allowed us to examine the role of phylogeny, colonizations, extinctions, community size, productivity, habitat availability, and dispersal limitation in generating bird species richness gradients in Catalonia. However, due to the coarse structure of macroecological data used, the role of local species interactions in such processes remained yet obscure and elusive.Understanding the role of species interactions in the processes of coexistence and diversification of bird communities thus remained as a big challenge. Since long ago, ecological interactions have been hypothesized to play a role in the maintenance of diversity (Hutchinson 1959, MacArthur and Levins 1967, McPeek 1997, Mittelbach et al. 2007). Therefore, the quantitative study of species interactions was expected to provide insights on the processes that ultimately generate species richness in bird communities (Cattin et al. 2006, Rezende et al. 2007). However, data on species interactions is usually available only for specific local communities. Therefore, a macroecological approach is precluded because the bulk of the studies of community interaction networks are carried out at local scales. Accordingly, we examined the role of species interactions in the generation of species richness in a local Mediterranian bird community for which interaction good-quality data was available (Chapters 4 and 5) (Jordano 1984, 1987). We addressed the two general hypotheses delineated by Johnson and Stinchcombe (2007). First, in chapter 4, we analyzed if evolutionary processes causally affect present-day community interaction patterns and the dynamics of species interactions (H1). Secondly, in chapter five, we assessed if species interactions and community context might influence the direction, rate and outcome of present-day diversification processes that ultimately generate species richness (H2). Overall, our results provide strong support for the two hypotheses examined, thus highlighting the idea that evolutionary and ecological processes are effectively profoundly interlinked. On one hand, we show that long-term evolutionary processes effectively modulate present-day community interaction patterns and dynamics (Webb et al. 2002, Cavender-Bares and Wilczek 2003, Johnson and Stinchcombe 2007, Chapter 4). Likewise, we demonstrate that community context can potentially shape bird morphological diversification processes and drive speciation (Thompson 2005, Abrams 2006, Johnson and Stinchcombe 2007, Chapter 5). Overall, Chapters 4 and 5 provide some new empirical and theoretical insights dealing with the processes that generate and maintain local bird diversity from an integrated evolutionary and ecological perspective.

Restoration of streams that were formerly channelized for timber-floating has become increasingly common. Generally, this restoration returns boulders from riparian zones to streams, leading to wider, more heterogeneous channels with slower flows. The primary goal is to enhance fish populations, but riparian vegetation is also expected to be favoured. However, increases in floristic diversity have not been observed and reasons for this slow response are still unknown. One possible explanation might be the lack of colonist pools. I therefore investigated how surrounding plant compositions influence riparian recovery. The vascular plant flora was identified in riparian sites and in adjacent upstream riparian and upland sites. Four reach types were included: unchannelized, channelized, restored and demonstration restored. Species richness and floristic similarities among types of sites and reaches were compared. Correlations with upland and upstream channel slopes were made and the importance of variation in seed floating ability was tested. The results show that unchannelized reaches were floristically similar to their adjacent upstream riparian and upland sites, whereas channelized reaches showed more different floras. Restoration created a somewhat more homogeneous flora among the three site types and demonstration restored reaches were most similar to upstream sites. Soil moisture conditions (i.e. wetland vs. forest) in the uplands had stronger impacts on species similarities than upland or upstream channel slopes. I conclude that adjacent sites are important for floristic recovery of riparian reaches and that demonstration restoration is most advantageous for riparian recovery. I recommend protection of upland sites from forestry to facilitate recovery. / Local- and landscape-scale effects on biodiversity after stream restoration

With close to 2100 species, the flowering plant genus Carex (Cyperaceae; sedges) is an example of an evolutionary radiation. Despite its potential for use as a model taxon in evolutionary studies, the diversification of sedges remains largely unexplored. This thesis realizes the potential of Carex as an evolutionary model group by using it to ask questions about species richness patterns. More specifically, it seeks to determine the relationship, if any, between rates of trait evolution and species richness. This tests the hypothesis that organisms with increased abilities to evolve new traits, speciate more rapidly. Morphological and ecological (habitat and climatic niche) traits are modelled on a nearly complete regional (North America north of Mexico) phylogeny and rates of trait evolution are compared among non-nested sister groups. However, before trait evolution is modelled, this work evaluates the sensitivity of climatic niche estimates to underlying distribution datasets. It tests the agreement of niche estimates derived from the commonly used online repository GBIF (the Global Biodiversity Information Facility) and county-level distributions via BONAP (the Biota of North America Program). Results showed that in the context of phylogenetic comparative analyses, it is not vital to obtain highly accurate climatic niche estimates. The second study found significant positive correlations between the rates of climatic niche, habitat and reproductive morphological evolution and species richness. This result supports the role of high trait lability in generating species richness and more generally, the idea that high trait disparity through evolutionary time leads to species success.

I evaluated effects of 5 pine plantation establishment regimes 6 – 8 years postestablishment on loblolly pine (Pinus taeda) growth, vegetation characteristics, nutritional carrying capacity for white-tailed deer, and breeding birds in the Lower Coastal Plain of Mississippi. Treatments combined mechanical site preparation (MSP), chemical site preparation (CSP), and herbaceous weed control (HWC) designed to represent a range of operational intensities. Chemical SP provided greater long-term control of woody competition than MSP, but did not provide significant pine growth advantage. Vegetation richness, diversity, and structure were best maintained with MSP and year 1 banded HWC. Canopy cover appears to be shading out herbaceous understory and altering composition of woody understory toward more shade-tolerant species. Total forage biomass and 3 levels of carrying capacity declined on average 54% each year. Avian metrics decreased as treatment intensity increased. Regionally important species were influenced positively by greater vegetation coverage attained by banded HWC.

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