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BACKGROUND AND AIMS: The Atlantic Forest biodiversity hotspot is a complex mosaic of habitat types. However, the diversity of the rain forest at the core of this complex has received far more attention than that of its marginal habitats, such as cloud forest, semi-deciduous forest or restinga. Here, we investigate broad-scale angiosperm tree diversity patterns along elevation gradients in the south-east Atlantic Forest and test if the diversity of marginal habitats is shaped from the neighbouring rain forest, as commonly thought. METHODS: We calculated phylogenetic indices that capture basal [mean pairwise phylogenetic distance (MPD)] and terminal [mean nearest taxon distance (MNTD)] phylogenetic variation, phylogenetic endemism (PE) and taxonomic and phylogenetic beta diversity (BD and PBD) for 2074 angiosperm tree species distributed in 108 circular sites of 10 km diameter across four habitat types i.e. rain forest, cloud forest, semi-deciduous forest and coastal vegetation known as restinga. We then related these metrics to elevation and environmental variables. KEY RESULTS: Communities in wetter and colder forests show basal phylogenetic overdispersion and short phylogenetic distances towards the tips, respectively. In contrast, communities associated with water deficit and salinity show basal phylogenetic clustering and no phylogenetic structure toward the tips. Unexpectedly, rain forest shows low PE given its species richness, whereas cloud and semi-deciduous forests show unusually high PE. The BD and PBD between most habitat types are driven by the turnover of species and lineages, except for restinga. CONCLUSIONS: Our results contradict the idea that all marginal habitat types of the Atlantic Forest are sub-sets of the rain forest. We show that marginal habitat types have different evolutionary histories and may act as 'equilibrium zones for biodiversity' in the Atlantic Forest, generating new species or conserving others. Overall, our results add evolutionary insights that reinforce the urgency of encompassing all habitat types in the Atlantic Forest concept.

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Factors driving the spatial configuration of centres of endemism have long been a topic of broad interest and debate. Due to different eco-evolutionary processes, these highly biodiverse areas may harbour different amounts of ancient and recently diverged organisms (paleo- and neo-endemism, respectively). Patterns of endemism still need to be measured at distinct phylogenetic levels for most clades and, consequently, little is known about the distribution, the age and the causes of such patterns. Here we tested for the presence of centres with high phylogenetic endemism (PE) in the highly diverse Neotropical snakes, testing the age of these patterns (paleo- or neo-endemism), and the presence of PE centres with distinct phylogenetic composition. We then tested whether PE is predicted by topography, by climate (seasonality, stability, buffering and relictualness), or biome size. We found that most areas of high PE for Neotropical snakes present a combination of both ancient and recently diverged diversity, which is distributed mostly in the Caribbean region, Central America, the Andes, the Atlantic Forest and on scattered highlands in central Brazil. Turnover of lineages is higher across Central America, resulting in more phylogenetically distinct PE centres compared to South America, which presents a more phylogenetically uniform snake fauna. Finally, we found that elevational range (topographic roughness) is the main predictor of PE, especially for paleo-endemism, whereas low paleo-endemism levels coincide with areas of high climatic seasonality. Our study highlights the importance of mountain systems to both ancient and recent narrowly distributed diversity. Mountains are both museums and cradles of snake diversity in the Neotropics, which has important implications for conservation in this region.

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Environmental and geographical variables are known drivers of community assembly, however their influence on phylogenetic structure and phylogenetic beta diversity of lineages within different bioregions is not well-understood. Using Neotropical palms as a model, we investigate how environmental and geographical variables affect the assembly of lineages into bioregions across an evolutionary time scale. We also determine lineage shifts between tropical (TRF) and non-tropical (non-TRF) forests. Our results identify that distance and area explain phylogenetic dissimilarity among bioregions. Lineages in smaller bioregions are a subset of larger bioregions and contribute significantly to the nestedness component of phylogenetic dissimilarity, here interpreted as evidence for a bioregional shift. We found a significant tendency of habitat shifts occurring preferentially between TRF and non-TRF bioregions (31 shifts) than from non-TRF to TRF (24) or from TRF to TRF (11) and non-TRF to non-TRF (9). Our results also present cases where low dissimilarity is found between TRF and non-TRF bioregions. Most bioregions showed phylogenetic clustering and larger bioregions tended to be more clustered than smaller ones, with a higher species turnover component of phylogenetic dissimilarity. However, phylogenetic structure did not differ between TRF and non-TRF bioregions and diversification rates were higher in only two lineages, Attaleinae and Bactridinae, which are widespread and overabundant in both TRF and non-TRF bioregions. Area and distance significantly affected Neotropical palm community assembly and contributed more than environmental variables. Despite palms being emblematic humid forest elements, we found multiple shifts from humid to dry bioregions, showing that palms are also important components of these environments.

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Although inselbergs from around the world are iconic ecosystems, little is known on the underlying mechanisms of community assembly, especially in their characteristic patchy outcrop vegetation. Environmental constraints are expected to cause phylogenetic clustering when ecological niches are conserved within evolutionary lineages. We tested whether vegetation patches from rock outcrops of the Piedra La Tortuga Natural Monument, in the northern Amazon region, are phylogenetically clustered, indicating that environmental filtering is the dominant driver of community assemblage therein. We classified all patches according to their size as very small (< 1 m2), small (1-4 m2), medium-sized (4-8 m2), and large patches (8-15 m2). From each class, we randomly selected 10 patches, totalizing 40 patches covering 226 m2. All individuals found in the 40 isolated patches were identified to the species level. We also correlated measurements of phylogenetic community structure with patch size. We found that species from patches are restricted to the clades monocots, fabids, malvids, and lamiids. We conclude that vegetation in this rock outcrop is phylogenetically clustered. Furthermore, we found that phylogenetic turnover between pairs of patches increases with patch size, which is consistent with a scenario of higher environmental stress in smaller patches. Further research is necessary to identify nurse species in inselberg vegetation, which is pivotal for conservation and restoration of this particular ecosystem.(AU)

Ainda que os inselbergs ao redor do mundo sejam ecossistemas icônicos, pouco se sabe sobre os mecanismos subjacentes que estruturam suas comunidades vegetais, especialmente nas manchas de vegetação sobre afloramentos rochosos. Espera-se que as restrições ambientais causem agrupamento filogenético quando os nichos ecológicos são conservados dentro das linhagens evolutivas. Nós testamos se as manchas de vegetação dos afloramentos rochosos do Monumento Natural Piedra La Tortuga, no norte da região amazônica, apresentam indicadores filogenéticos de que a filtragem ambiental é o principal direcionador da estruturação da comunidade. Classificamos todas as manchas de acordo com seu tamanho como muito pequenas (<1 m2), pequenas (1-4 m2), médias (4-8 m2) e grandes (8-15 m2). Selecionamos aleatoriamente 10 manchas em cada classe de tamanho, totalizando 40 manchas cobrindo 226 m2. Todos os indivíduos encontrados nas 40 manchas foram identificados ao nível de espécie. Correlacionamos as medidas da estrutura filogenética da comunidade com o tamanho das manchas e encontramos que as espécies das manchas são restritas aos clados das monocotiledôneas, fabídeas, malvídeas e lamiídeas. Concluímos que a vegetação neste afloramento rochoso é agrupada filogeneticamente. Além disso, encontramos que o turnover filogenético entre pares de manchas aumenta com o tamanho da mancha, o que é consistente com um cenário de alto estresse ambiental nas manchas menores. São necessárias mais pesquisas para identificar espécies facilitadoras, que são fundamentais para a conservação e restauração destes ecossistemas.(AU)

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RESUMEN

Although inselbergs from around the world are iconic ecosystems, little is known on the underlying mechanisms of community assembly, especially in their characteristic patchy outcrop vegetation. Environmental constraints are expected to cause phylogenetic clustering when ecological niches are conserved within evolutionary lineages. We tested whether vegetation patches from rock outcrops of the Piedra La Tortuga Natural Monument, in the northern Amazon region, are phylogenetically clustered, indicating that environmental filtering is the dominant driver of community assemblage therein. We classified all patches according to their size as very small (< 1 m2), small (1-4 m2), medium-sized (4-8 m2), and large patches (8-15 m2). From each class, we randomly selected 10 patches, totalizing 40 patches covering 226 m2. All individuals found in the 40 isolated patches were identified to the species level. We also correlated measurements of phylogenetic community structure with patch size. We found that species from patches are restricted to the clades monocots, fabids, malvids, and lamiids. We conclude that vegetation in this rock outcrop is phylogenetically clustered. Furthermore, we found that phylogenetic turnover between pairs of patches increases with patch size, which is consistent with a scenario of higher environmental stress in smaller patches. Further research is necessary to identify nurse species in inselberg vegetation, which is pivotal for conservation and restoration of this particular ecosystem.

Ainda que os inselbergs ao redor do mundo sejam ecossistemas icônicos, pouco se sabe sobre os mecanismos subjacentes que estruturam suas comunidades vegetais, especialmente nas manchas de vegetação sobre afloramentos rochosos. Espera-se que as restrições ambientais causem agrupamento filogenético quando os nichos ecológicos são conservados dentro das linhagens evolutivas. Nós testamos se as manchas de vegetação dos afloramentos rochosos do Monumento Natural Piedra La Tortuga, no norte da região amazônica, apresentam indicadores filogenéticos de que a filtragem ambiental é o principal direcionador da estruturação da comunidade. Classificamos todas as manchas de acordo com seu tamanho como muito pequenas (<1 m2), pequenas (1-4 m2), médias (4-8 m2) e grandes (8-15 m2). Selecionamos aleatoriamente 10 manchas em cada classe de tamanho, totalizando 40 manchas cobrindo 226 m2. Todos os indivíduos encontrados nas 40 manchas foram identificados ao nível de espécie. Correlacionamos as medidas da estrutura filogenética da comunidade com o tamanho das manchas e encontramos que as espécies das manchas são restritas aos clados das monocotiledôneas, fabídeas, malvídeas e lamiídeas. Concluímos que a vegetação neste afloramento rochoso é agrupada filogeneticamente. Além disso, encontramos que o turnover filogenético entre pares de manchas aumenta com o tamanho da mancha, o que é consistente com um cenário de alto estresse ambiental nas manchas menores. São necessárias mais pesquisas para identificar espécies facilitadoras, que são fundamentais para a conservação e restauração destes ecossistemas.

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