The role of evolutionary modes for trait-based cascades in mutualistic networks

The erosion of functional diversity may foster the collapse of ecological systems. Functional diversity is ultimately determined by the distribution of species traits. As species traits are a legacy of species evolutionary history, one might expect that the mode of trait evolution influences community resistance under the loss of functional diversity. In this paper, we investigate the role of trait evolutionary dynamics on the robustness of mutualistic networks undergoing the following scenarios of species loss: (i) random extinctions, (ii) loss of functional distinctiveness and (iii) extinctions biased towards larger sizes. We simulated networks defined by models of single trait complementary and evolutionary modes where traits can arise in recent diversification events with weak phylogenetic signal, in early diversification events with strong phylogenetic signal, or as a random walk through evolutionary time. Our simulations show that mutualistic networks are especially vulnerable to extinctions based on trait distinctiveness and more robust to random extinction dynamics. The networks show an intermediate level of robustness against size-based extinctions. Despite the small range of variation in network robustness, our results show that the mode of trait evolution matters for network robustness in all three scenarios. Networks with low phylogenetic signal are more robust than networks with high phylogenetic signal across all scenarios. As a consequence, our results predict that mutualistic networks based upon current adaptations are more likely to cope with extinction dynamics than those networks that are based upon conserved traits

Is evolution faster at ecotones? A test using rates and tempo of diet transitions in Neotropical Sigmodontinae (Rodentia, Cricetidae)

We evaluated whether evolution is faster at ecotones as niche shifts may be needed to persist under unstable environment. We mapped diet evolution along the evolutionary history of 350 sigmodontine species. Mapping was used in three new tipbased metrics of trait evolution – Transition Rates, Stasis Time, and Last Transition Time – which were spatialized at the assemblage level (aTR, aST, aTL). Assemblages were obtained by superimposing range maps on points located at core and ecotone of the 93 South American ecoregions. Using Linear Mixed Models, we tested whether ecotones have species with more changes from the ancestral diet (higher aTR), have maintained the current diet for a shorter time (lower aST), and have more recent transitions to the current diet (lower aLT) than cores. We found lower aTR, and higher aST and aLT at ecotones than at cores. Although ecotones are more heterogeneous, both environmentally and in relation to selection pressures they exert on organisms, ecotone species change little from the ancestral diet as generalist habits are necessary toward feeding in ephemeral environments. The need to incorporate phylogenetic uncertainty in tip-based metrics was evident from large uncertainty detected. Our study integrates ecology and evolution by analyzing how fast trait evolution is across space

Elevational shifts in tree community composition in the Brazilian Atlantic Forest related to climate change

Question: Climate change induces shifts in species distributions, ultimately changing community composition. Mountains are especially sensitive to climate change, and tree species are predicted to move towards higher elevations, but observed changes are not always unidirectional. The diversity and complexity of tropical and subtropical systems limits our understanding of climate‐induced responses of subtropical mountain forests. Here, we investigated migration trends in tree community composition along an elevational gradient, and between the transition from lowland to montane forests in subtropical forests. Location: Brazilian Atlantic Forest. Methods: We used thermal affiliations of 627 tree species to calculate community temperature scores (CTS) for different life‐history stages of trees in 96 permanent plots. We compared CTS of different life‐history stages across space and time. Results: Most tree communities (58%) did not show a significant difference of CTS between life‐history stages, indicating a non‐migration trend. On the other hand, 27% of tree communities showed upward migration and 15% downward migration. Upward migration was more common in montane forests, and downward migration in lowland forests. Our temporal analysis shows significant changes in CTS values for juvenile communities with 0.36°C decrease in lowland forests and 0.34°C increase in montane forests. Conclusions: Contrasting results between lowland and montane forest communities indicate that the transition zone influences migration patterns and may reflect differences in species’ thermal limitations, as well as by non‐thermal factors such as biotic interactions. Our findings provide the first evidence of climate change‐induced community shifts in the Brazilian Atlantic Forest. We demonstrated that upward migration trends were predominantly observed in montane–upper mountain forests, while downward migrations were noted in lowland–submontane forests. The thermophilization of montane forests may suggest an increased risk of reduction for cold‐demanding species under climate change scenarios

How do distinct facets of tree diversity and community assembly respond to environmental variables in the subtropical Atlantic Forest?

This study assessed the impact of altitude, precipitation, and soil conditions on species richness (SR), phylogenetic diversity (PD), and functional diversity (FD) standardized effect sizes in subtropical Brazilian Atlantic Forest tree communities. We considered specific trait information (FDs) for FD, reflecting recent adaptive evolution, contrasting with deeper phylogenetic constraints in FD. Three functional traits (leaf area‐LA, wood density‐WD, and seed mass‐SM) were examined for their response to these gradients. Generalized least squares models with environmental variables as predictors and diversity metrics as response variables were used, and a fourth‐corner correlation test explored trait‐environmental relationships. SR decreased with altitude, while PD increased, indicating niche convergence at higher altitudes. Leaf area and seed mass diversity also decreased with altitude. For LA, both FD and FDs were significant, reflecting filtering processes influenced by phylogenetic inheritance and recent trait evolution. For SM, only the specific trait structure responded to altitude. LA and SM showed significant trait‐environmental relationships, with smaller‐leaved and lighter‐seeded species dominant at higher altitudes. Soil gradients affect diversity. Fertile soils have a wider range of LA, indicating coexistence of species with different nutrient acquisition strategies. WD variation is lower for FDs. SM diversity has different relationships with soil fertility for FDs and FD, suggesting phylogeny influences trait variation. Soil pH influences WD and LA under acidic soils, with deeper phylogenetic constraints (FD). Environmental factors impact tree communities, with evidence of trait variation constraints driven by conditions and resources. Subtropical Atlantic forests' tree assemblies are mainly influenced by altitude, pH, and soil fertility, selecting fewer species and narrower trait spectra under specific conditions (e.g., higher altitudes, pH). Functional diversity patterns reflect both phylogenetic and recent evolution constraints, with varying strength across traits and conditions. These findings highlight the intricate processes shaping long‐lived species assembly across diverse environments in the Southern Brazilian Atlantic Forest

Species niches, not traits, determine abundance and occupancy patterns: A multi‐site synthesis

International audienceAim: Locally abundant species are usually widespread, and this pattern has been related to properties of the niches and traits of species. However, such explanations fail to account for the potential of traits to determine species niches and often overlook statistical artefacts. Here, we examine how trait distinctiveness determines the abilities of species to exploit either common habitats (niche position) or a range of habitats (niche breadth) and how niche position and breadth, in turn, affect abundance and occupancy. We also examine how statistical artefacts moderate these relationships. Location: Sixteen sites in the Neotropics. Time period 1993–2014. Major taxa studied Aquatic invertebrates from tank bromeliads. Methods: We measured the environmental niche position and breadth of each species and calculated its trait distinctiveness as the average trait difference from all other species at each site. Then, we used a combination of structural equation models and a meta-analytical approach to test trait–niche relationships and a null model to control for statistical artefacts. Results: The trait distinctiveness of each species was unrelated to its niche properties, abundance and occupancy. In contrast, niche position was the main predictor of abundance and occupancy; species that used the most common environmental conditions found across bromeliads were locally abundant and widespread. Contributions of niche breadth to such patterns were attributable to statistical artefacts, indicating that effects of niche breadth might have been overestimated in previous studies. Main conclusions: Our study reveals the generality of niche position in explaining one of the most common ecological patterns. The robustness of this result is underscored by the geographical extent of our study and our control of statistical artefacts. We call for a similar examination across other systems, which is an essential task to understand the drivers of commonness across the tree of life

Estrutura filogenética de comunidades de plantas lenhosas em ecótonos vegetacionais

A busca de padrões consistentes na natureza tem sido a principal meta dos ecólogos. Essa dissertação teve como objetivo usar abordagens filogenéticas na tentativa de compreender melhor o processo ecológico da expansão florestal sobre áreas abertas. O uso da informação filogenética em análises ecológicas considera as espécies não independentes umas das outras, pois estas compartilham grande parte da história evolutiva. Essa hierarquia de organização das espécies é muito importante para determinar as regras que governam os processos de montagem das comunidades locais. Nesta dissertação foram avaliados padrões filogenéticos de estruturação da vegetação lenhosa florestal ocorrente em ecótonos de áreas abertas com vegetação florestal distribuídos em diferentes regiões do extremo sul do Brasil. Estes ecótonos são formados por diversas formações florestais, as quais tendem a expandir sobre as áreas abertas. Dados sobre composição de espécies provieram de estudos já realizados e de amostragens em alguns sítios. Duas métricas filogenéticas complementares foram usadas para avaliar a estrutura filogenética em cada categoria de habitat nos ecótonos: índice de parentesco líquido (NRI) e coordenadas principais da estrutura filogenética (PCPS). As análises dos valores de NRI não mostraram um padrão nítido de estruturação filogenética das comunidades. Já a análise dos PCPS mostrou padrões consistentes nas três escalas espaciais abordadas e independente da composição de espécies. Clados basais associaram-se às áreas florestais, enquanto clados de diversificação recente associaram-se às áreas abertas. Estes resultados indicam que áreas abertas atuam como um filtro filogenético de habitat para as espécies lenhosas florestais em todos os locais analisados, independentemente da escala e da composição de espécies de cada local. Os resultados sugerem que os clados de Rosanae e Asteranae estão na linha de frente do processo de expansão florestal sobre as áreas abertas, e o clado de Magnolianae restrito às áreas florestais. A busca por padrões gerais de organização das comunidades ecológicas a partir de sua estrutura filogenética parece consistir numa ferramenta útil para a exploração e entendimento sobre o funcionamento de sistemas ecológicos. Estas abordagens poderiam beneficiar estratégias de gerenciamento e conservação destes sistemas, por simplificarem sistemas ecológicos complexos e por mostrarem padrões gerais independentes da escala espacial analisada.The search for consistent patterns in nature has been a major goal of ecologists. This study aimed to employ phylogenetic analyses to improve the understanding of an ecological process, the expansion of forest expansion over open areas. The use of evolutionary information considers species as not independent units in relation to each other, as they share their evolutionary history. Such hierarchical organization of species is very important to determine the rules governing assembly processes in local communities. Phylogenetic approaches were used to evaluate phylogenetic patterns in forest woody vegetation occurring in ecotones comprising open areas and forest vegetation, and distributed across different regions in the southernmost Brazilian region. Those ecotones are composed by different forest vegetation types, which tend to expand over open areas. Data on species composition were compiled from previous studies, and vegetation sampling was carried out in sites without available information on species composition in ecotones. Two complementary phylogenetic metrics were used to evaluate the phylogenetic structure in each habitat type occurring in ecotones: net relatedness index (NRI) and principal coordinates of phylogenetic structure (PCPS). Analyses of NRI values did not show any clear pattern of phylogenetic structuring of the communities. Nonetheless, PCPS analysis showed consistent patterns across the threes spatial scales evaluated, which were independent of the species composition of the sites. Basal clades were associated with forest areas, while late-divergence clades were associated with open areas. These findings indicate that open areas act as a phylogenetic habitat filtering to forest woody species throughout the region, independently of the spatial scale and of the species composition in each site. The results suggest that the clades Rosanae and Asteranae represent the vanguard in theforest expansion process over open areas, while the distribution of the basal clade Magnolianae is restricted to forest sites. The search for general organization patterns in ecological communities based on their phylogenetic structure seems to be a useful tool for exploring and understanding the functioning of ecological systems. Such approach might benefit ecosystem managing and conservation strategies, as it simplifies complex ecological systems, and shows general patterns independently of the scale analyzed

Sinal filogenético e conservação filogenética de nicho : integrando métodos aos conceitos ecológicos

Compreender os fatores que afetam a distribuição das espécies tem sido um dos principais objetivos dos ecólogos. Atualmente, sabe-se que os processos ecológicos e evolutivos moldam a dinâmica de especiação e extinção de espécies, e determinam a distribuição e abundância das mesmas. Ao longo dos últimos anos, tem havido um aumento no número de estudos que utilizam informação filogenética para explicar as dinâmicas populacionais e as distribuições de espécies, e que buscam identificar os mecanismos responsáveis pela montagem das comunidades. Interações das espécies, sejam elas intraespecífica, interespecífica ou com o ambiente, ocorrem baseadas nas diferenças e semelhanças fenotípicas. Essas variações fenotípicas tem origem na evolução das espécies, e com isso espera-se que as espécies proximamente relacionadas tendam a ser ecologicamente mais semelhantes entre si do que as espécies distantemente relacionadas. Esta concepção tem dado origem a um conceito importante, com implicações para estudos tanto ecológicos quanto evolutivos: o conceito de conservação filogenética de nicho, isto é, quando as espécies relacionadas mantêm seus nichos ancestrais ao longo do tempo evolutivo. Esse padrão tem importância para diversas áreas de ecologia, permitindo a ligação das espécies aos processos ecológicos e auxiliando na maior compreensão da ecologia evolutiva das diferentes linhagens. Devido à sua importância, é fundamental o desenvolvimento de métodos estatísticos adequados para quantificar esses padrões e inferir os processos que o subjazem. Atualmente, os métodos utilizados para inferir conservação filogenética de nicho são, em sua maioria, incompatíveis com determinados conceitos ecológicos e não abrangem todos os tipos de dados e esse fato explica uma visão incompleta dos processos presentes nas comunidades e conflitante com o objetivo de muitos estudos ecológicos e conservacionistas que buscam vincular as espécies aos processos ecológicos e evolutivos. Desta forma, o principal objetivo desta tese é propor novos métodos para quantificar o sinal filogenético que integrem diferentes aspectos do conceito de nicho ecológico. Apresentamos aqui os novos métodos em detalhes e avaliamos suas propriedades estatísticas (erro tipo I e poder estatístico) por meio de dados simulados. No capítulo 1, nós propomos um método para medir sinal filogenético utilizando o teste de Mantel, incorporando modelos evolutivos para testar hipóteses específicas da evolução dos atributos. No capítulo 2, descrevemos um conjunto de funções e um novo pacote estatístico para explorar os padrões filogenéticos no nível de metacomunidade. Este pacote permite explorar a distribuição de linhagens filogenéticas através de gradientes ecológicos, a análise de sinal filogenético no nível da metacomunidade e explorar a associação entre clados e gradientes ecológicos. No capítulo 3, investigamos a relação entre sinal filogenético dos atributos com os padrões de coocorrência das espécies nos níveis da comunidade. Esta abordagem permite testar se espécies filogeneticamente relacionadas que coocorrem expressam as suas dimensões de nicho com maior semelhança do que seria esperado por modelos neutros de evolução. Por fim, testamos as propriedades estatísticas destes métodos em relação dois modelos nulos, que incorporam diferentes aspectos da estrutura da comunidade e evolução dos atributos das espécies. Os três capítulos representam diferentes trabalhos que se interconectam no sentido de elucidar o conceito de sinal filogenético e conservação filogenética de nicho.Understanding the factors that can affect species distributions has been a main goal of ecologists. Currently, it is known that evolutionary and ecological processes shape the speciation dynamics, species extinction and determine the distribution and abundance of species. Over the last years, there has been an increase in the number of studies using phylogenetic information to explain the dynamics of population, species distribution and identifying the mechanisms of community assembly. Species interactions – intraspecific, interspecific or with the environment – occur based on their phenotypic differences and similarities. As phenotypic variation has a basis in evolutionary history, it is expected that closely related species tend to be more ecologically similar to each other than distantly related ones. This notion has given rise to an important concept, with implications for both evolutionary and ecological studies: the concept of phylogenetic niche conservatism, that is, when related species maintain their ancestral niches over evolutionary time. This pattern is important for several areas of ecology, and allows to link species to ecological processes and to understand the evolutionary ecology of different lineages. Despite its importance, it is crucial the development of appropriate statistical method to measure this pattern and to infer the processes behind it. The methods currently available to infer phylogenetic niche conservatism are sometimes incompatible with some ecological concepts and do not cover all kind of data, this fact leads to an incomplete view of the process acting in the currents communities and conflict with the goal of many ecological and conservation studies that need to link species to ecological and evolutionary processes. The main goal of this dissertation is to propose novel methods to measure phylogenetic signal incorporating different aspects of ecological niche. We introduce novel methods in detail and evaluate its statistical properties (type I error and statistical power) by means of simulated data with known structure. In chapter 1 we propose a method to measure phylogenetic signal using the Mantel test, incorporating evolutionary model to test specific hypothesis of trait evolution. In chapter 2, we describe a set of function and a new statistical package for exploring the phylogenetic patterns at the metacommunity level. This package allows the exploration of distribution of phylogenetic lineages across ecological gradients, the analysis of phylogenetic signal at metacommunity level and to explore the association between clades and ecological gradients. In the chapter 3, we access the relationship between phylogenetic signal in traits and species co-occurrence patterns in the community levels. This approach allows one to test whether phylogenetic close related species cooccurring in metacommunities express their niche dimensions more similarly than would be expected by neutral expectation. We tested the statistical properties of these methods in relation to two null models, which incorporate these different aspects of the community structure and evolution of species traits. The three chapters represent different works that are interconnected in order to elucidate the concept of phylogenetic signal and phylogenetic niche conservatism

Estrutura filogenética de comunidades de plantas lenhosas em ecótonos vegetacionais

A busca de padrões consistentes na natureza tem sido a principal meta dos ecólogos. Essa dissertação teve como objetivo usar abordagens filogenéticas na tentativa de compreender melhor o processo ecológico da expansão florestal sobre áreas abertas. O uso da informação filogenética em análises ecológicas considera as espécies não independentes umas das outras, pois estas compartilham grande parte da história evolutiva. Essa hierarquia de organização das espécies é muito importante para determinar as regras que governam os processos de montagem das comunidades locais. Nesta dissertação foram avaliados padrões filogenéticos de estruturação da vegetação lenhosa florestal ocorrente em ecótonos de áreas abertas com vegetação florestal distribuídos em diferentes regiões do extremo sul do Brasil. Estes ecótonos são formados por diversas formações florestais, as quais tendem a expandir sobre as áreas abertas. Dados sobre composição de espécies provieram de estudos já realizados e de amostragens em alguns sítios. Duas métricas filogenéticas complementares foram usadas para avaliar a estrutura filogenética em cada categoria de habitat nos ecótonos: índice de parentesco líquido (NRI) e coordenadas principais da estrutura filogenética (PCPS). As análises dos valores de NRI não mostraram um padrão nítido de estruturação filogenética das comunidades. Já a análise dos PCPS mostrou padrões consistentes nas três escalas espaciais abordadas e independente da composição de espécies. Clados basais associaram-se às áreas florestais, enquanto clados de diversificação recente associaram-se às áreas abertas. Estes resultados indicam que áreas abertas atuam como um filtro filogenético de habitat para as espécies lenhosas florestais em todos os locais analisados, independentemente da escala e da composição de espécies de cada local. Os resultados sugerem que os clados de Rosanae e Asteranae estão na linha de frente do processo de expansão florestal sobre as áreas abertas, e o clado de Magnolianae restrito às áreas florestais. A busca por padrões gerais de organização das comunidades ecológicas a partir de sua estrutura filogenética parece consistir numa ferramenta útil para a exploração e entendimento sobre o funcionamento de sistemas ecológicos. Estas abordagens poderiam beneficiar estratégias de gerenciamento e conservação destes sistemas, por simplificarem sistemas ecológicos complexos e por mostrarem padrões gerais independentes da escala espacial analisada.The search for consistent patterns in nature has been a major goal of ecologists. This study aimed to employ phylogenetic analyses to improve the understanding of an ecological process, the expansion of forest expansion over open areas. The use of evolutionary information considers species as not independent units in relation to each other, as they share their evolutionary history. Such hierarchical organization of species is very important to determine the rules governing assembly processes in local communities. Phylogenetic approaches were used to evaluate phylogenetic patterns in forest woody vegetation occurring in ecotones comprising open areas and forest vegetation, and distributed across different regions in the southernmost Brazilian region. Those ecotones are composed by different forest vegetation types, which tend to expand over open areas. Data on species composition were compiled from previous studies, and vegetation sampling was carried out in sites without available information on species composition in ecotones. Two complementary phylogenetic metrics were used to evaluate the phylogenetic structure in each habitat type occurring in ecotones: net relatedness index (NRI) and principal coordinates of phylogenetic structure (PCPS). Analyses of NRI values did not show any clear pattern of phylogenetic structuring of the communities. Nonetheless, PCPS analysis showed consistent patterns across the threes spatial scales evaluated, which were independent of the species composition of the sites. Basal clades were associated with forest areas, while late-divergence clades were associated with open areas. These findings indicate that open areas act as a phylogenetic habitat filtering to forest woody species throughout the region, independently of the spatial scale and of the species composition in each site. The results suggest that the clades Rosanae and Asteranae represent the vanguard in theforest expansion process over open areas, while the distribution of the basal clade Magnolianae is restricted to forest sites. The search for general organization patterns in ecological communities based on their phylogenetic structure seems to be a useful tool for exploring and understanding the functioning of ecological systems. Such approach might benefit ecosystem managing and conservation strategies, as it simplifies complex ecological systems, and shows general patterns independently of the scale analyzed

PCPS – an R-package for exploring phylogenetic eigenvectors across metacommunities

PCPS is an R package for exploring phylogenetic eigenvectors across metacommunities. It offers a set of functions for analyzing principal coordinates of phylogenetic structure (PCPS), allowing analysis of phylogenetic signal in ecological traits of species at the metacommunity level, and the association between each PCPS and environmental, spatial and historical factors. The package is a flexible solution for exploring the distribution of major phylogenetic lineages across ecological or biogeographic gradients. The package is freely available on the CRAN official web server for R.&nbsp