Field sampling is biased against small-ranged species of high conservation value: a case study on the sphingid moths of East Africa

The range size of species co-occurring in local assemblages is a pivotal variable in assessments of a site’s conservation value. Assemblages featuring many small-ranged species are given more priority than assemblages consisting mainly of wide-ranging species. However, the assembly of relevant information can be challenging and local range size distributions of tropical invertebrates are rarely available for conservation planning. We present such data for sphingid moths in East Africa, a highly diverse region of high conservation value. We compare geographic range size distributions based on field samples with predictions from modelled range map data. Using this system as a case study, we provide evidence for a systematic sampling bias when inferring average local range sizes from field data. Unseen species (i.e., species present but missed in local sampling) are often those with small ranges (hence, of high conservation value). Using an elevational gradient, we illustrate how this bias can lead to false, counterintuitive assessments of environmental effects on local range size distributions. Furthermore, with particular reference to sphingid moths in the study region, we show that current protected areas appear unrelated to the spatial distribution of species richness or average geographic range sizes at a local scale. We discuss the need to treat field sampled data with caution and in concert with other data sources such as probabilistic models.     

Organization of Squamata (Reptilia) assemblages in Mediterranean archipelagos

Mediterranean islands have complex reptile assemblages, but little is known about the factors that determine their organization. In this study, the structure of assemblages of Squamata was evaluated based on their species richness and two measures of phylogenetic diversity (variability and clustering). I evaluated the composition of the assemblages comparing distinct biogeographic subregions within the Mediterranean: Adriatic, Aegean, Balearic, Corsica–Sardinia, Crete, Gulf of Gabés, Ionian Sea, Ligurian Sea, Malta, Sicily, and Tyrrhenian Sea. The effect of island environments and geographical isolation on the diversity metrics was assessed using generalized linear models. The analyses indicated that species richness was mostly influenced by island area and geographical isolation. Assemblages on smaller islands were poorer in species and phylogenetically dispersed, possibly as an effect of interspecific competition. The species composition of the assemblages was determined by similar environmental drivers within the biogeographic subregions, including island area, island elevation, geographical isolation, and aridity. In several subregions, significant patterns of phylogenetic attraction were found in species co‐occurrences, caused by the limits imposed by the island size on large predatory species.     

Influence of tree shape and evolutionary time‐scale on phylogenetic diversity metrics

During the last decades, describing, analysing and understanding the phylogenetic structure of species assemblages has been a central theme in both community ecology and macro‐ecology. Among the wide variety of phylogenetic structure metrics, three have been predominant in the literature: Faith's phylogenetic diversity (PD_Faith), which represents the sum of the branch lengths of the phylogenetic tree linking all species of a particular assemblage, the mean pairwise distance between all species in an assemblage (MPD) and the pairwise distance between the closest relatives in an assemblage (MNTD). Comparisons between studies using one or several of these metrics are difficult because there has been no comprehensive evaluation of the phylogenetic properties each metric captures. In particular it is unknown how PD_Faith relates to MDP and MNTD. Consequently, it is possible that apparently opposing patterns in different studies might simply reflect differences in metric properties. Here, we aim to fill this gap by comparing these metrics using simulations and empirical data. We first used simulation experiments to test the influence of community structure and size on the mismatch between metrics whilst varying the shape and size of the phylogenetic tree of the species pool. Second we investigated the mismatch between metrics for two empirical datasets (gut microbes and global carnivoran assemblages). We show that MNTD and PD_Faith provide similar information on phylogenetic structure, and respond similarly to variation in species richness and assemblage structure. However, MPD demonstrate a very different behaviour, and is highly sensitive to deep branching structure. We suggest that by combining complementary metrics that are sensitive to processes operating at different phylogenetic depths (i.e. MPD and MNTD or PD_Faith) we can obtain a better understanding of assemblage structure.     

Phylogenetic structure and phylogenetic diversity of angiosperm assemblages in forests along an elevational gradient in Changbaishan,China

Aims Understanding what drives the variation in species composition and diversity among local communities can provide insights into the mechanisms of community assembly. Because ecological traits are often thought to be phylogenetically conserved, there should be patterns in phylogenetic structure and phylogenetic diversity in local communities along ecological gradients. We investigate potential patterns in angiosperm assemblages along an elevational gradient with a steep ecological gradient in Changbaishan, China.Methods We used 13 angiosperm assemblages in forest plots (32×32 m) distributed along an elevational gradient from 720 to 1900 m above sea level. We used Faith's phylogenetic diversity metric to quantify the phylogenetic alpha diversity of each forest plot, used the net relatedness index to quantify the degree of phylogenetic relatedness among angiosperm species within each forest plot and used a phylogenetic dissimilarity index to quantify phylogenetic beta diversity among forest plots. We related the measures of phylogenetic structure and phylogenetic diversity to environmental (climatic and edaphic) factors.Important findings Our study showed that angiosperm assemblages tended to be more phylogenetically clustered at higher elevations in Changbaishan. This finding is consistent with the prediction of the phylogenetic niche conservatism hypothesis, which highlights the role of niche constraints in governing the phylogenetic structure of assemblages. Our study also showed that woody assemblages differ from herbaceous assemblages in several major aspects. First, phylogenetic clustering dominated in woody assemblages, whereas phylogenetic overdispersion dominated in herbaceous assemblages; second, patterns in phylogenetic relatedness along the elevational and temperature gradients of Changbaishan were stronger for woody assemblages than for herbaceous assemblages; third, environmental variables explained much more variations in phylogenetic relatedness, phylogenetic alpha diversity and phylogenetic beta diversity for woody assemblages than for herbaceous assemblages.     

Phylocom: software for the analysis of phylogenetic community structure and trait evolution

MOTIVATION: The increasing availability of phylogenetic and trait data for communities of co-occurring species has created a need for software that integrates ecological and evolutionary analyses. Capabilities: Phylocom calculates numerous metrics of phylogenetic community structure and trait similarity within communities. Hypothesis testing is implemented using several null models. Within the same framework, it measures phylogenetic signal and correlated evolution for species traits. A range of utility functions allow community and phylogenetic data manipulation, tree and trait generation, and integration into scientific workflows. Availability: Open source at: http://phylodiversity.net/phylocom/.     

Joint effect of phylogenetic relatedness and trait selection on the elevational distribution of Rhododendron species

Congeneric species may coexist at fine spatial scales through niche differentiation, however, the magnitude to which the effects of functional traits and phylogenetic relatedness contribute to their distribution along elevational gradients remains understudied. To test the hypothesis that trait and elevational range overlap can affect local speciesʼ coexistence, we first compared phylogenetic relatedness and trait (including morphological traits and leaf elements) divergence among closely related species of Rhododendron L. on Yulong Mountain, China. We then assessed relationships between the overlap of multiple functional traits and the degree of elevational range overlap among species pairs in a phylogenetic context. We found that phylogeny was a good predictor for most functional traits, where closely related species showed higher trait similarity and occupied different elevational niches at our study site. Species pairs of R. subgen. Hymenanthes (Blume) K. Koch showed low elevational range overlap and some species pairs of R. subgen. Rhododendron showed obvious niche differentiation. Trait divergence is greater for species in R. subgen. Rhododendron, and it plays an important role between species pairs with low elevational range overlap. Trait convergent selection takes place between co-occurring closely related species that have high elevational range overlap, which share more functional trait space due to environmental filtering or ecological adaptation in more extreme habitats. Our results highlight the importance of evolutionary history and trait selection for species coexistence at fine ecological scales along environmental gradients.     

Contrasting patterns of phylogenetic assemblage structure along the elevational gradient for major hummingbird clades

Aim We evaluated the hypothesis that, given niche conservatism, relatedness of co‐occurring hummingbird species of a given clade will increase at greater distances from the elevation where it originated. We also used prior knowledge of flight biomechanics and feeding specialization of hummingbird species (family Trochilidae) to evaluate which environmental variables were important predictors of changes in phylogenetic structure for each hummingbird clade. Location Ecuador. Methods We compiled species lists for 189 local hummingbird assemblages across major environmental gradients in Ecuador from a variety of published and non‐published sources. For the entire family and each of the major hummingbird clades (hermits, emeralds, mangoes, coquettes and brilliants) we quantified the phylogenetic structure of each assemblage using the net relatedness index (NRI). This index calculates the standardized mean of all possible pairwise phylogenetic distances between co‐occurring species. We related NRI for each clade to elevation, precipitation and vegetation‐related variables using generalized additive models. Results Our findings support the prediction of an increase in the co‐occurrence of close relatives away from the elevation of origin at the family level and for assemblages of mangoes and brilliants. The opposite pattern was found for assemblages of coquettes and emeralds. For the hermits, variation in phylogenetic structure was not explained by elevation. Clades with high levels of feeding specialization (hermits and brilliants) always included a vegetation‐related variable as an important predictor of change in phylogenetic structure. Main conclusions We found no overall support for the conservatism and zone of origin hypotheses. Knowledge of each clade’s natural history proved useful for predicting which environmental variables correlated with phylogenetic structure of local assemblages. Clades with the same elevation of origin appear to have radiated along the elevational gradient in association with different environmental factors.     

Phylogenetic structure of angiosperm trees in local forest communities along latitudinal and elevational gradients in eastern North America

Latitudinal and elevational gradients both represent thermal gradients. Assessing the consistency of the relationships between phylogenetic structure and climate between latitudinal and elevational gradients can provide insight into the mechanisms driving assembly of species from regional pools into local assemblages. The aim of this study is to compare patterns of phylogenetic structure measures for angiosperm tree species between latitudinal and elevational gradients, using a dataset of angiosperm tree species in 14 092 forest plots in eastern North America. We assessed whether these two gradients produce similar relationships between climate and phylogenetic structure, hypothesizing that they should differ in magnitude but not direction. We used correlation and regression analyses to assess the relation of measures of phylogenetic structure to elevation, latitude and climatic variables, which included minimum temperature, temperature seasonality, annual precipitation and precipitation seasonality. We found that 1) phylogenetic relatedness of angiosperm trees increases with decreasing temperature along both latitudinal and elevational gradients but the relationship between phylogenetic relatedness and temperature is steeper for elevational gradients than for latitudinal gradients; 2) the tip-weighted metric of phylogenetic relatedness (nearest taxon index) is more strongly correlated with climatic variables than the basal-weighted metric of phylogenetic relatedness (net relatedness index); 3) winter cold temperature exerts a stronger effect on community assembly of angiosperm trees than does temperature seasonality. These results suggest that winter cold temperature, rather than temperature seasonality, drives phylogenetic structure of plants in local forest communities, and that species distributions along elevational gradients are more in equilibrium with temperature, compared with those along latitudinal gradients.     

Functional structure and specialization in three tropical plant–hummingbird interaction networks across an elevational gradient in Costa Rica

Understanding causes of variation in multispecies assemblages along spatial environmental gradients is a long‐standing research topic in ecology and biogeography. Ecological networks comprising interacting species of plants and pollinators are particularly suitable for testing effects of environmental gradients on the functional structure and specialization in multispecies assemblages. In this study, we investigated patterns in functional assemblage structure and specialization of hummingbirds at the individual and species level along a tropical elevational gradient. We mist‐netted hummingbirds at three elevations in Costa Rica in seven temporally distinct sampling periods and used the pollen carried by hummingbird individuals to construct plant–hummingbird networks at each elevation. We measured four functional traits of hummingbird species and quantified different metrics of functional community structure. We tested the effect of elevation on functional metrics of hummingbird assemblages and specialization within the networks, employing the variability across sampling periods and hummingbird species to compare the respective metrics among elevations. Hummingbird species and individuals were more specialized at low and mid elevations than at the highest elevation. This pattern corresponded to a more even and over‐dispersed assemblage structure at the lower elevations throughout the year and suggests a high level of floral resource partitioning in functionally diversified communities. In contrast, an uneven and clustered functional structure of the highland assemblage across all sampling periods suggests that this assemblage was structured by environmental filtering and by niche expansion of hummingbird individuals and species at this elevation. We conclude that high degrees of specialization on specific floral resources might be crucial for the coexistence of hummingbird species in diversified lowland communities. Spatial variation in animal resource use may be an important crucial driver of spatial patterns in the functional structure of diversified species assemblages also in other types of ecological networks.     

Phylogenetic diversity of macromycetes and woody plants along an elevational gradient in Eastern Mexico

Phylogenetic information provides insight into the ecological and evolutionary processes that organize species assemblages. We compared patterns of phylogenetic diversity among macromycete and woody plant communities along a steep elevational gradient in eastern Mexico to better understand the evolutionary processes that structure their communities. Macrofungi and trees were counted and identified in eight sites from 100 to 3500 m asl, and sequence data retrieved from GenBank for the same or closely related species were used to reconstruct their phylogenies. Patterns of species richness and phylogenetic diversity were similar for both macrofungi and trees, but macromycete richness and diversity peaked at mid‐elevations, whereas woody plant richness and diversity did not show significant trends with elevation. Phylogenetic similarity among sites was low for both groups and decreased as elevational distance between sites increased. Macromycete communities displayed phylogenetic overdispersion at low elevations and phylogenetic clustering at high elevations; the latter is consistent with environmental filtering at high elevation sites. Woody plants generally exhibited phylogenetic clustering, consistent with the potential importance of environmental filtering throughout the elevational gradient.     

Behavior influences range limits and patterns of coexistence across an elevational gradient in tropical birds

Does competition influence patterns of coexistence between closely related taxa? Here we address this question by analyzing patterns of range overlap between related species of birds (‘sister pairs’) co‐occurring on a tropical elevational gradient. We explicitly contrast the behavioral dimension of interspecific competition (interference competition) with similarity in resource acquisition traits (exploitative competition). Specifically, we ask whether elevational range overlap in 118 sister pairs that live along the Manu Transect in southeastern Peru is predicted by proxies for competition (intraspecific territorial behavior) or niche divergence (beak divergence and divergence times, an estimate of evolutionary age). We find that close relatives that defend year‐round territories tend to live in non‐overlapping elevational distributions, while close relatives that do not defend territories tend to broadly overlap in elevational distribution. In contrast, neither beak divergence nor evolutionary age was associated with patterns of range limitation. We interpret these findings as evidence that behavioral interactions – particularly direct territorial aggression – can be important in setting elevational range limits and preventing coexistence of closely related species, though this depends upon the extent to which intraspecific territorial behavior can be extended to territorial interactions between species. Our results suggest that interference competition can be an important driver of species range limits in diverse assemblages, and thus highlight the importance of considering behavioral dimensions of the niche in macroecological studies.     

Phylogenetic diversity,trait diversity and niches: species assembly of ferns along a tropical elevational gradient

Aim To analyse the structure of pteridophyte assemblages, based on phylogenetic relatedness and trait properties, along an elevational gradient. Ecological theory predicts that co‐occurring species may be: randomly selected from a regional pool; ecologically sorted so that they are functionally different hence resulting in reduced competition (overdispersion); or functionally similar as an adaptation to specific ecological conditions (clustering). Location Braulio Carrillo National Park and Cerro de la Muerte, Costa Rica, Central America. Methods We used an empirical dataset of the quantitative pattern of species occurrences and individual numbers of ferns within 156 plots along a tropical elevational gradient to test whether directed ecological sorting might cause deviations in patterns of trait and phylogenetic diversity. Mean pairwise distances of species based on phylogenetic and trait properties were compared with two different sets of null assemblages, one maintaining species frequency distributions (constrained) and one not (unconstrained). Results Applying different null models resulted in varying degrees of overdispersion and clustering, but overall patterns of deviation from random expectations remained the same. Contrary to theoretical predictions, phylogenetic and trait diversity were relatively independent from one another. Phylogenetic diversity showed no patterns along the elevational gradient, whereas trait diversity showed significant trends for epiphytes. Main conclusions Under stressful environmental conditions (drought at low elevations and frost at high elevations), epiphytic fern assemblages tended to be clustered with respect to trait characteristics, which suggests environmental filtering. Conversely, under less extreme environmental conditions (middle of the transect), the sorting was biased towards high differentiation (overdispersion), presumably because of interspecific competition and trait shifts among closely related species (character displacement).     

Elevational filtering and the evolution of planthoppers (Hemiptera,Fulgoromorpha) in Papua New Guinea

Along elevational gradients, phylogenetic relatedness patterns constitute a considerable source of information and may shed light on ecological processes that structure communities. This study focuses on community phylogenetic structure of planthoppers, specifically the species-rich and abundant Fulgoromorpha families (Hemiptera, Auchenorrhyncha), Cixiidae and Derbidae + Achilidae, along an elevational gradient on Mount Wilhelm (Papua New Guinea). In order to assess the factors driving planthoppers community composition, we recorded abundance data for planthoppers species at each elevation and we generated a molecular phylogeny of the local species, using Bayesian inference. We analyzed 168 individuals representing 59 local morphospecies. Using a fully resolved and well-supported phylogeny, we then investigated the phylogenetic structure of the communities by performing a Spatial Analysis of Community Diversity. We show that Cixiidae are phylogenetically clustered along the elevational gradient, whereas Derbidae + Achilidae harbor a random structure, suggesting that local adaptation to elevation shapes community structure of Cixiidae, but not that of Derbidae + Achilidae. Our findings highlight the importance of phylogenies in the study of tropical elevational gradients.     

Exploring the phylogenetic history of mammal species richness

Aim At broad geographical scales, species richness is a product of three basic processes: speciation, extinction and migration. However, determining which of these processes predominates is a major challenge. Whilst palaeontological studies can provide information on speciation and extinction rates, data are frequently lacking. Here we use a recent dated phylogenetic tree of mammals to explore the relative importance of these three processes in structuring present‐day richness gradients. Location The global terrestrial biosphere. Methods We combine macroecological data with phylogenetic methods more typically used in community ecology to describe the phylogenetic history of regional faunas. Using simulations, we explore two simple phylogenetic metrics, the mean and variance in the pairwise distances between taxa, and describe their relationship to phylogenetic tree topology. We then use these two metrics to characterize the evolutionary relationships among mammal species assemblages across the terrestrial biome. Results We show that the mean and variance in the pairwise distances describe phylogenetic tree topology well, but are less sensitive to phylogenetic uncertainty than more direct measures of tree shape. We find the phylogeny for South American mammals is imbalanced and ‘stemmy’ (long branches towards the root), consistent with recent diversification within evolutionarily disparate lineages. In contrast, the phylogeny for African mammals is balanced and ‘tippy’ (long branches towards the tips), more consistent with the slow accumulation of diversity over long times, reflecting the Old World origin of many mammal clades. Main conclusions We show that phylogeny can accurately capture biogeographical processes operating at broad spatial scales and over long time periods. Our results support inferences from the fossil record – that the New World tropics are a diversity cradle whereas the Old World tropics are a museum of old diversity.     

The influence of variability in species trait data on community‐level ecological prediction and inference

Species trait data have been used to predict and infer ecological processes and the responses of biological communities to environmental changes. It has also been suggested that, in lieu of trait, data niche differences can be inferred from phylogenetic distance. It remains unclear how variation in trait data may influence the strength and character of ecological inference. Using species‐level trait data in community ecology assumes intraspecific variation is small in comparison with interspecific variation. Intraspecific variation across species ranges or within populations may lead to variability in trait data derived from different scales (i.e., local or regional) and methods (i.e., mean or maximum values). Variation in trait data across species can affect community‐level relationships. I examined variability in body size, a key trait often measured across taxa. I collected 12 metrics of fish species length (including common and maximum values) for 40 species from literature, online databases, museum collections, and field data. I then tested whether different metrics of fish length could consistently predict observed species range boundary shifts and the impacts of an introduced predator on inland lake fish communities across Ontario, Canada. I also investigated whether phylogenetic signal, an indicator of niche‐conservativism, changed among measures. I found strong correlations between length metrics and limited variation across metrics. Accordingly, length was a consistently significant predictor of the response of fish communities to environmental change. Additionally, I found significant evidence of phylogenetic signal in fish length across metrics. Limited variation in length across metrics (within species), in comparison with variation within metrics (across species), made fish species length a reliable predictor at a community‐level. When considering species‐level trait data from different sources, researchers should examine the potential influence of intraspecific trait variation on data derived by different metrics and at different scales.     

The spatial structure of phylogenetic and functional diversity in the United States and Canada: An example using the sedge family (Cyperaceae)

Systematically quantifying diversity across landscapes is necessary to understand how clade history and ecological heterogeneity contribute to the origin, distribution, and maintenance of biodiversity. Here, we chart the spatial structure of diversity among all species in the sedge family (Cyperaceae) throughout the USA and Canada. We first identify areas of remarkable species richness, phylogenetic diversity, and functional trait diversity, and highlight regions of conservation priority. We then test predictions about the spatial structure of this diversity based on the historical biogeography of the family. Incorporating a phylogeny, over 400 000 herbarium records, and a database of functional traits mined from online floras, we find that species richness and functional trait diversity peak in the Northeastern USA, while phylogenetic diversity peaks along the Gulf of Mexico. Floristic turnover among assemblages increases significantly with distance, but phylogenetic turnover is twice as rapid along latitudinal gradients as along longitudinal gradients. These patterns reflect the expected distribution of Cyperaceae, which originated in the tropics but radiated in temperate regions. We identify assemblages with an abundance of rare, range‐restricted lineages, and assemblages composed of species generally lacking from diverse regions. We argue that both of these metrics are useful for developing targeted conservation strategies. We use the data generated here to establish future research priorities, including the testing of a series of hypotheses regarding the distribution of chromosome numbers, photosynthetic pathways, and resource partitioning in sedges.     

Temperature, but not productivity or geometry, predicts elevational diversity gradients in ants across spatial grains

Aim  This research aims to understand the factors that shape elevational diversity gradients and how those factors vary with spatial grain. Specifically, we test the predictions of the species–productivity hypothesis, species–temperature hypothesis, the metabolic theory of ecology and the mid-domain effects null model. We also examine how the effects of productivity and temperature on richness depend on spatial grain.
Location  Deciduous forests along an elevational gradient in Great Smoky Mountains National Park, USA.
Methods  We sampled 22 leaf litter ant assemblages at three spatial grains, from 1-m² quadrats to 50 × 50 m plots using Winkler samplers.
Results  Across spatial grains, warmer sites had more species than did cooler sites, and primary productivity did not predict ant species richness. We found some support for the predictions of the metabolic theory of ecology, but no support for the mid-domain effects null model. Thus, our data are best explained by some version of a species–temperature hypothesis.
Main conclusions  Our results suggest that temperature indirectly affects ant species diversity across spatial grains, perhaps by limiting access to resources. Warmer sites support more species because they support more individuals, thereby reducing the probability of local extinction. Many of our results from this elevational gradient agree with studies at more global scales, suggesting that some mechanisms shaping ant diversity gradients are common across scales.     

Body size, rarity, and phylogenetic community structure: insights from diving beetle assemblages of Alberta

Although it is implicit that interactions between species depend on their traits, studies on the probability of finding related species in a community are in their infancy. Community composition and species richness of predaceous diving beetles (Dytiscidae: Coleoptera) have been used as indicators of freshwater ecosystem function yet no incorporation of phylogenetic relationships of coexisting dytiscids has been attempted to date. Improved knowledge of phylogenetic relationships and phylogenetic community structure analysis methods may provide additional insight into the relationships between community composition and species richness, thus impacting our interpretation of aquatic indicator species metrics. Here, we use museum records of dytiscid beetles in 53 lakes of Alberta, Canada to: (1) compile a supertree of dytiscid beetles that live in the province, (2) examine whether coexisting dytiscids tend to be more or less related than expected by chance, and (3) examine whether phylogenetic structuring depends on species richness or mean size of coexisting species. We find that, although the majority of dytiscid assemblages exhibited phylogenetic clustering, the extent to which this occurred depended on the mean size of dytiscids. We discuss the potential mechanisms and implications of the observed patterns in phylogenetic clustering, along with data that would further improve our understanding of community dynamics in dytiscid beetles.     

Functional and phylogenetic structure of forest and savanna bird assemblages across spatial scales

Ecological and evolutionary mechanisms that drive community assembly vary in space and time. However, little is known about how such mechanisms act in contrasting habitats. Here, we estimated the functional and phylogenetic structure of forest and savanna bird assemblages across different spatial scales to understand: 1) the mechanisms that govern the structure of assemblages in these habitats; 2) the relationship between phylogenetic and functional structure; and 3) the influence of species richness on the functional and phylogenetic structure of assemblages. We used a null model where forest and savanna bird species were allowed to occur in the same null assemblages and other where species were separated based on their habitats. According to the first null model, forest bird assemblages were functionally and phylogenetically clustered at all spatial scales, whereas savanna bird assemblages generally showed random functional and phylogenetic structure. These results can be explained by the low dispersal rate of forest species across of the patchy habitats and the widespread distribution of savanna species. However, in the second null model, both forest and savanna bird assemblages showed random functional and phylogenetic structure at regional and local scales. This suggests that trait‐based assembly might not play an important role in both habitats and across different spatial scales. In addition, the phylogenetic and functional structure of assemblages were not correlated, evidencing that caution is necessary when using phylogenetic relationships as a surrogate to functional distances among species. Finally, the relationships between species richness and functional and phylogenetic structure indicated that an increase in the number of species can promote both clustering and overdispersion, depending on the studied habitat and scale. Our study shows that integrating different types of habitat, spatial scales and biodiversity components in a single framework can shed light on the mechanisms that determine the community assembly.