Incorporating intraspecific variation in tests of trait-based community assembly

Environmental filtering and niche differentiation are processes proposed to drive community assembly, generating nonrandom patterns in community trait distributions. Despite the substantial intraspecific trait variation present in plant communities, most previous studies of trait-based community assembly have used species mean trait values and therefore not accounted for intraspecific variation. Using a null model approach, I tested for environmental filtering and niche differentiation acting on three key functional traits??vegetative height, specific leaf area (SLA), and leaf dry matter content (LDMC)??in old-field plant communities. I also examined how accounting for intraspecific variation at the among-plot and individual levels affected the detection of nonrandom assembly patterns. Tests using fixed species mean trait values provided evidence of environmental filtering acting on height and SLA and niche differentiation acting on SLA. Including plot-level intraspecific variation increased the strength of these patterns, indicating an important role of intraspecific variation in community assembly. Tests using individual trait data indicated strong environmental filtering acting on all traits, but provided no evidence of niche differentiation, although these signals may have been obscured by the effects of dispersal limitation and spatial aggregation of conspecific individuals. There was also strong evidence of nonrandom assembly of individuals within single species, with the strength of environmental filtering varying among species. This study demonstrates that, while analyses using fixed species mean trait values can provide insights into community assembly processes, accounting for intraspecific variation provides a more complete view of communities and the processes driving their assembly.     

Trait‐based approach confirms the importance of propagule limitation and assembly rules in old‐field restoration

Community assembly theory is suggested as a guiding principle for ecological restoration to help understand the mechanisms that structure biological communities and identify where restoration interventions are needed. We studied three hypotheses related to propagule limitation, stress‐dominance, and limiting similarity concepts in community assembly in a restoration field experiment with a trait‐based null model approach. The experiment aimed to assist the recovery of sand grassland on former arable land in the Kiskunság, Pannonian biogeographic region, Europe. Treatments included initial seeding of five grassland species, carbon amendment, low‐intensity mowing, and combinations in 1 m by 1 m plots in three old fields from 2003 to 2008. The distribution of 10 individual plant traits was compared to the null model and the effect of time and treatments were tested with linear mixed effect models. Initial seeding had the most visible impact on species and trait composition confirming propagule limitation in grassland recovery. Reducing nutrient availability through carbon amendment strengthened trait convergence for length of flowering as expected based on the stress‐dominance hypothesis. Mowing changed trait divergence to convergence for plant height with a strengthening impact with time, supporting our hypothesis of increasing dominance of limiting similarity with time. Our results support the idea that community assembly is simultaneously influenced by propagule limitation and multiple trait‐based processes that act through different traits. The limited impact of manipulating environmental filtering and limiting similarity compared to seeding, however, supports the view that only targeting the dispersal and environmental filters in parallel would improve restoration outcome.     

Trait but not species convergence during plant community assembly in restored semi‐natural grasslands

Community assembly or succession was traditionally thought of as being deterministic and directional, leading to a clearly defined climax state. The alternative view, however, keeps gaining attention. This view states that community assembly is influenced by historical processes, where differences in the sequence and timing of species arrival result in distinct communities. Here we tested the hypothesis that both views are valid, but at a different level, with increasing dissimilarity in species composition among sites with increasing age (divergence), caused by historical processes (priority effects), and with increasing similarity in mean trait composition (convergence) among sites, indicating a directional development at the niche level. We surveyed a chronosequence of restored semi‐natural grassland patches on former pine plantations over seven restoration age classes, covering 22 grasslands. Pairwise multivariate distances were calculated between the different grassland patches based on species abundance on the one hand, and on mean community trait values for 28 plant life history traits on the other. Trait composition showed a clear decrease in multivariate distance with increasing restoration age, indicating trait convergence through time. At the species level, we found no evidence of convergence through time, with even a trend towards divergence. Furthermore, spatial variation and environmental heterogeneity were found to remain constant through time. These results confirm our hypothesis. At the trait level, limited niches occur, only filled by species having the appropriate traits, resulting in a clear deterministic model of assembly. Species identity, on the contrary, has no role in this niche filling. The first appropriate species to reach a restoration site will be most likely the ones that get established, resulting in divergence of the species composition among restored grasslands.     

Community assembly during vegetation succession after metal mining is driven by multiple processes with temporal variation

The mechanisms governing community assembly is fundamental to ecological restoration and clarification of the assembly processes associated with severe disturbances (characterized by no biological legacy and serious environmental problems) is essential. However, a systematic understanding of community assembly in the context of severe anthropogenic disturbance remains lacking. Here, we explored community assembly processes after metal mining, which is considered to be a highly destructive activity to provide insight into the assembly rules associated with severe anthropogenic disturbance. Using a chronosequence approach, we selected vegetation patches representing different successional stages and collected data on eight plant functional traits from each stage. The traits were classified as establishment and regenerative traits. Based on these traits, null models were constructed to identify the processes driving assembly at various successional stages. Comparison of our observations with the null models indicated that establishment and regenerative traits converged in the primary stage of succession. As succession progressed, establishment traits shifted to neutral assembly, whereas regeneration traits alternately converged and diverged. The observed establishment traits were equal to expected values, whereas regenerative traits diverged significantly after more than 20 years of succession. Furthermore, the available Cr content was linked strongly to species'' ecological strategies. In the initial stages of vegetation succession in an abandoned metal mine, the plant community was mainly affected by the available metal content and dispersal limitation. It was probably further affected by strong interspecific interaction after the environmental conditions had improved, and stochastic processes became dominant during the stage with a successional age of more than 20 years.     

Advances, challenges and a developing synthesis of ecological community assembly theory

Ecological approaches to community assembly have emphasized the interplay between neutral processes, niche-based environmental filtering and niche-based species sorting in an interactive milieu. Recently, progress has been made in terms of aligning our vocabulary with conceptual advances, assessing how trait-based community functional parameters differ from neutral expectation and assessing how traits vary along environmental gradients. Experiments have confirmed the influence of these processes on assembly and have addressed the role of dispersal in shaping local assemblages. Community phylogenetics has forged common ground between ecologists and biogeographers, but it is not a proxy for trait-based approaches. Community assembly theory is in need of a comparative synthesis that addresses how the relative importance of niche and neutral processes varies among taxa, along environmental gradients, and across scales. Towards that goal, we suggest a set of traits that probably confer increasing community neutrality and regionality and review the influences of stress, disturbance and scale on the importance of niche assembly. We advocate increasing the complexity of experiments in order to assess the relative importance of multiple processes. As an example, we provide evidence that dispersal, niche processes and trait interdependencies have about equal influence on trait-based assembly in an experimental grassland.     

Global to community scale differences in the prevalence of convergent over divergent leaf trait distributions in plant assemblages

Aim The drivers of species assembly, by limiting the possible range of functional trait values, can lead to either convergent or divergent distributions of traits in realized assemblages. Here, to evaluate the strengths of these species assembly drivers, we partition trait variance across global, regional and community scales. We then test the hypothesis that, from global to community scales, the outcome of co‐occurring trait convergence and divergence is highly variable across biomes and communities. Location Global: nine biomes ranging from subarctic highland to tropical rain forest. Methods We analysed functional trait diversity at progressively finer spatial scales using a global, balanced, hierarchically structured dataset from 9 biomes, 58 communities and 652 species. Analyses were based on two key leaf traits (foliar nitrogen content and specific leaf area) that are known to drive biogeochemical cycling. Results While 35% of the global variance in these traits was between biomes, only 15% was between communities within biomes and as much as 50% occurred within communities. Despite this relatively high within‐community variance in trait values, we found that trait convergence dominated over divergence at both global and regional scales through comparisons of functional trait diversity in regional and community assemblages against random (null) models of species assembly. Main conclusions We demonstrate that the convergence of traits occurring from global to regional assemblages can be twice as strong as that from regional to community assemblages, and argue that large differences in the nature and strength of abiotic and biotic drivers of dominant species assembly can, at least partly, explain the variable outcome of simultaneous trait convergence and divergence across sites. Ultimately, these findings stress the urgent need to extend species assembly research to address those scales where trait variance is the highest, i.e. between biomes and within communities.     

Disentangling niche and neutral influences on community assembly: assessing the performance of community phylogenetic structure tests

Patterns of phylogenetic relatedness within communities have been widely used to infer the importance of different ecological and evolutionary processes during community assembly, but little is known about the relative ability of community phylogenetics methods and null models to detect the signature of processes such as dispersal, competition and filtering under different models of trait evolution. Using a metacommunity simulation incorporating quantitative models of trait evolution and community assembly, I assessed the performance of different tests that have been used to measure community phylogenetic structure. All tests were sensitive to the relative phylogenetic signal in species metacommunity abundances and traits; methods that were most sensitive to the effects of niche-based processes on community structure were also more likely to find non-random patterns of community phylogenetic structure under dispersal assembly. When used with a null model that maintained species occurrence frequency in random communities, several metrics could detect niche-based assembly when there was strong phylogenetic signal in species traits, when multiple traits were involved in community assembly, and in the presence of environmental heterogeneity. Interpretations of the causes of community phylogenetic structure should be modified to account for the influence of dispersal.     

Using functional trait diversity to evaluate the contribution of multiple ecological processes to community assembly during succession

Successional chronosequences provide a unique opportunity to study the effects of multiple ecological processes on plant community assembly. Using a series of 0.5 × 0.5 m² plots (n = 30) from five successional sub‐alpine meadow plant communities (ages 3, 5, 9, 12, and undisturbed) in the Qinghai‐Tibetan Plateau, we investigated whether community assembly is stochastic or deterministic for species and functional traits. We tested directional change in species composition, functional trait composition, and then functional trait diversity measured by Rao's quadratic entropy for four traits – plant height, leaf dry matter content, specific leaf area, and seed mass – along two comparable successional chronosequences. We then evaluated the importance of species interactions, habitat filtering and stochasticity by comparing with random communities and partitioning the environmental and spatial components of Rao's quadratic entropy. We found no directional change in species composition, but clear directionality in functional trait composition. None of the abiotic environmental variables (except P) showed linear change with successional age, but soil moisture and nitrogen were positively related to functional diversity within meadows. Functional trait diversity increased significantly with the increase in successional age. Comparison with random communities showed a significant shift from trait divergence in early stages of succession (3‐ and 5‐yr) to convergence in the later stages of succession 9‐, 12‐yr and undisturbed). The relative importance of abiotic variables and spatial structure for functional trait diversity changed in a predictable manner with successional age. Stochasticity at the species level may indicate dispersal limitation, but deterministic effects on functional trait distributions show the role of both habitat effects and biotic interactions.     

Mechanisms of ecological succession: insights from plant functional strategies

Successions are a central issue of ecological theory. They are governed by changes in community assembly processes that can be tracked by species’ traits. While single‐trait‐based approaches have been mostly promoted to address community assembly, ecological strategies actually encompass tradeoffs between multiple traits that are relevant to succession theory. We analyzed plant ecological strategies along a 140‐year‐long succession primary succession of 52 vertical outcrop communities after roadwork. We performed a RLQ analysis to relate six functional traits, associated with resource acquisition, competition, colonization ability and phenology, to the age of the outcrops. We found the prominence of two main axes of specialization, one related to resource acquisition and the other to reproduction and regeneration. We further examined the community‐level variation in ecological strategies to assess the abiotic and biotic drivers of community assembly. Using trait‐based statistics of functional richness, regularity and divergence, we found that different processes drove the variation in ecological strategies along the axes of specialization. In late succession, functional convergence was detected for the traits related to resource acquisition as a signature of habitat filtering, while the coexistence of contrasted strategies was found for the traits related to reproduction and regeneration as a result of spatial micro‐heterogeneity. We observed a lack of niche differentiation along the succession, revealing a weak importance of biotic interactions for the regulation of community assembly in the outcrops. Overall, we highlight a prominent role of habitat filtering and spatial micro‐heterogeneity in driving the primary succession governed by water and nutrient limitation.     

The quest for trait convergence and divergence in community assembly: are null‐models the magic wand?

The relevance of neutral versus niche‐based community assembly rules (i.e. the processes sorting species present in a larger geographical region into local communities) remains to be demonstrated in ecology and biogeography. To attempt to do this, a number of complex null models are increasingly being used that compare observed community functional diversity (FD, i.e. the extent of trait dissimilarity between coexisting species) with randomly simulated FD. However, little is known about the performance of these null models in detecting non‐neutral community assembly rules such as trait convergence and divergence of communities (supposedly revealing habitat selection and limiting similarity, respectively). Here, using both simulated and field communities, I show that assembly rule detection varies systematically with the magnitude of the observed FD, so that these null models do not really succeed in breaking down the observed functional relationships between species. This is a particular concern, making detection of community assembly dependent on: (1) the pool of samples considered, and (2) the capacity of observed FD to correctly discriminate these rules. Null models should be more thoroughly described and validated before being considered as a magic wand to reveal assembly patterns.     

Species divergence and trait convergence in experimental plant community assembly

Despite decades of research, it remains controversial whether ecological communities converge towards a common structure determined by environmental conditions irrespective of assembly history. Here, we show experimentally that the answer depends on the level of community organization considered. In a 9‐year grassland experiment, we manipulated initial plant composition on abandoned arable land and subsequently allowed natural colonization. Initial compositional variation caused plant communities to remain divergent in species identities, even though these same communities converged strongly in species traits. This contrast between species divergence and trait convergence could not be explained by dispersal limitation or community neutrality alone. Our results show that the simultaneous operation of trait‐based assembly rules and species‐level priority effects drives community assembly, making it both deterministic and historically contingent, but at different levels of community organization.     

片段化森林群落构建的生态过程及其检验方法

鉴于全球森林均呈现片段化(破碎化)的分布状态, 理解片段化森林群落构建的过程很有必要。该文通过综述群落构建的主要生态过程如生态漂变、扩散、选择和物种形成等在片段化森林群落构建中的相对作用, 发现因片段化森林形成方式的不同, 重构群落(片段化生境中通过次生演替重新形成的森林群落)和解构群落(原有森林被片段化后形成的森林群落)在不同演替阶段所受到的主要生态过程的相对作用有所不同。虽然利用基于群落内物种分布格局推测构建过程(如物种多度分布、零模型结合β多样性的方法、功能特征的收敛和发散等)、人工控制实验、群落结构动态分析等方法对片段化森林中群落构建的过程进行了有效的检验, 但是针对片段化森林群落构建过程的实验性研究仍然不足。未来有待在理论模型、群落构建过程的检验以及理论与物种保护相结合等方面继续开展深入的研究。     

Underdispersion and overdispersion of traits in terrestrial snail communities on islands

Understanding and disentangling different processes underlying the assembly and diversity of communities remains a key challenge in ecology. Species can assemble into communities either randomly or due to deterministic processes. Deterministic assembly leads to species being more similar (underdispersed) or more different (overdispersed) in certain traits than would be expected by chance. However, the relative importance of those processes is not well understood for many organisms, including terrestrial invertebrates. Based on knowledge of a broad range of species traits, we tested for the presence of trait underdispersion (indicating dispersal or environmental filtering) and trait overdispersion (indicating niche partitioning) and their relative importance in explaining land snail community composition on lake islands. The analysis of community assembly was performed using a functional diversity index (Rao's quadratic entropy) in combination with a null model approach. Regression analysis with the effect sizes of the assembly tests and environmental variables gave information on the strength of under‐ and overdispersion along environmental gradients. Additionally, we examined the link between community weighted mean trait values and environmental variables using a CWM‐RDA. We found both trait underdispersion and trait overdispersion, but underdispersion (eight traits) was more frequently detected than overdispersion (two traits). Underdispersion was related to four environmental variables (tree cover, habitat diversity, productivity of ground vegetation, and location on an esker ridge). Our results show clear evidence for underdispersion in traits driven by environmental filtering, but no clear evidence for dispersal filtering. We did not find evidence for overdispersion of traits due to diet or body size, but overdispersion in shell shape may indicate niche differentiation between snail species driven by small‐scale habitat heterogeneity. The use of species traits enabled us to identify key traits involved in snail community assembly and to detect the simultaneous occurrence of trait underdispersion and overdispersion.     

植物群落构建机制研究进展

群落构建研究对于解释物种共存和物种多样性的维持是至关重要的,因此一直是生态学研究的中心论题。尽管近年来关于生态位和中性理论的验证研究已经取得了显著的成果,但对于局域群落构建机制的认识仍存在很大争议。随着统计和理论上的进步使得用功能性状和群落谱系结构解释群落构建机制变为可能,主要是通过验证共存物种的性状和谱系距离分布模式来实现。然而,谱系和功能性状不能相互替代,多种生物和非生物因子同时控制着群落构建,基于中性理论的扩散限制、基于生态位的环境过滤和竞争排斥等多个过程可能同时影响着群落的构建。所以,综合考虑多种方法和影响因素探讨植物群落的构建机制,对于预测和解释植被对干扰的响应,理解生物多样性维持机制有重要意义。试图在简要回顾群落构建理论及研究方法发展的基础上,梳理其最新研究进展,并探讨整合功能性状及群落谱系结构的研究方法,解释群落构建和物种多样性维持机制的可能途径。在结合功能性状和谱系结构研究群落构建时,除了考虑空间尺度、环境因子、植被类型外,还应该关注时间尺度、选择性状的种类和数量、性状的种内变异、以及人为干扰等因素对群落构建的影响。     

Trends in single trait dispersion between early-mid successional stages: the importance of species pool extension and habitat scale

Aims Are there trends of increasing/decreasing dispersion of single, categorical traits related to early/late-successional species between stages of community development? If yes, are these trends dependent on species pool extension and habitat scale? Is there a consistent reduction in single trait convergence or divergence in any seral stage when scaling down from ecological to local species pool?Methods Presence of all vascular species rooted within plots of 5 × 5 m was recorded in assemblages of exposed mining spoils (EMS) and heathlands (HTL), which form a chronosequence on two abandoned ore tailing heaps located close to each other in the south-eastern Carpathians (Romania). Fifteen nominal, trait attributes of plant species co-occurring in the two seral assemblages were collected from available databases and subsequently classified as either successionally 'pioneer' or 'mature'. The strength of single trait convergence or divergence was estimated by comparison with null plant assemblages at patch type (meta-community) level by reference to the ecological or local species pool, and at community level.Important findings At patch type level, all pioneer and mature trait attributes (apart from short life span), with significant variation between the two seral stages, increased and, respectively, decreased in dispersion irrespective of species pool extension. However, these trends were more conspicuous when using the ecological species pool, very likely due to relaxation in abiotic filtering and dispersal limitation. At community level, no consistent trends were observed between EMS and HTL assemblages, probably because most trait attributes were sorted by microenvironmental filters displaying high variation, like topography or habitat patch geometry. In both seral stages, there was a general weakening of trait convergence or divergence at patch type level when scaling down from the ecological to the local species pool, which was due to niche space contraction. At community level, there was a trend of rise in dispersion of pioneer attributes along the observed chronosequence, presumably imputable to increasing competition for light and underground water, but an opposite trend of dispersion drop in mature attributes was not so evident. Based on these findings, we proposed two rules of thumb concerning the expected changes in dispersion of trait attributes at patch level along successions and between levels of species pool extension. In conclusion, trends in the successional dynamics of pioneer and mature trait dispersion are clearly detectable at meta-community level, especially by reference to the ecological species pool. Habitat scale and species pool extension are key factors to consider and report when estimating the magnitude of single trait dispersion.     

Compositional divergence and convergence in local communities and spatially structured landscapes

Community structure depends on both deterministic and stochastic processes. However, patterns of community dissimilarity (e.g. difference in species composition) are difficult to interpret in terms of the relative roles of these processes. Local communities can be more dissimilar (divergence) than, less dissimilar (convergence) than, or as dissimilar as a hypothetical control based on either null or neutral models. However, several mechanisms may result in the same pattern, or act concurrently to generate a pattern, and much research has recently been focusing on unravelling these mechanisms and their relative contributions. Using a simulation approach, we addressed the effect of a complex but realistic spatial structure in the distribution of the niche axis and we analysed patterns of species co-occurrence and beta diversity as measured by dissimilarity indices (e.g. Jaccard index) using either expectations under a null model or neutral dynamics (i.e., based on switching off the niche effect). The strength of niche processes, dispersal, and environmental noise strongly interacted so that niche-driven dynamics may result in local communities that either diverge or converge depending on the combination of these factors. Thus, a fundamental result is that, in real systems, interacting processes of community assembly can be disentangled only by measuring traits such as niche breadth and dispersal. The ability to detect the signal of the niche was also dependent on the spatial resolution of the sampling strategy, which must account for the multiple scale spatial patterns in the niche axis. Notably, some of the patterns we observed correspond to patterns of community dissimilarities previously observed in the field and suggest mechanistic explanations for them or the data required to solve them. Our framework offers a synthesis of the patterns of community dissimilarity produced by the interaction of deterministic and stochastic determinants of community assembly in a spatially explicit and complex context.     

A conceptual framework for the spatial analysis of functional trait diversity

Functional trait diversity is a popular tool in modern ecology, mainly used to infer assembly processes and ecosystem functioning. Patterns of functional trait diversity are shaped by ecological processes such as environmental filtering, species interactions and dispersal that are inherently spatial, and different processes may operate at different spatial scales. Adding a spatial dimension to the analysis of functional trait diversity may thus increase our ability to infer community assembly processes and to predict change in assembly processes following disturbance or land‐use change. Richness, evenness and divergence of functional traits are commonly used indices of functional trait diversity that are known to respond differently to large‐scale filters related to environmental heterogeneity and dispersal and fine‐scale filters related to species interactions (competition). Recent developments in spatial statistics make it possible to separately quantify large‐scale patterns (variation in local means) and fine‐scale patterns (variation around local means) by decomposing overall spatial autocorrelation quantified by Moran's coefficient into its positive and negative components using Moran eigenvector maps (MEM). We thus propose to identify the spatial signature of multiple ecological processes that are potentially acting at different spatial scales by contrasting positive and negative components of spatial autocorrelation for each of the three indices of functional trait diversity. We illustrate this approach with a case study from riparian plant communities, where we test the effects of disturbance on spatial patterns of functional trait diversity. The fine‐scale pattern of all three indices was increased in the disturbed versus control habitat, suggesting an increase in local scale competition and an overall increase in unexplained variance in the post‐disturbance versus control community. Further research using simulation modeling should focus on establishing the proposed link between community assembly rules and spatial patterns of functional trait diversity to maximize our ability to infer multiple processes from spatial community structure.     

Discriminating trait‐convergence and trait‐divergence assembly patterns in ecological community gradients

Question: Whereas similar ecological requirements lead to trait‐convergence assembly patterns (TCAP) of species in communities, the interactions controlling how species associate produce trait‐divergence assembly patterns (TDAP). Yet, the linking of the latter to community processes has so far only been suggested. We offer a method to elucidate TCAP and TDAP in ecological community gradients that will help fill this gap. Method: We evaluated the correlation between trait‐based described communities and ecological gradients, and using partial correlation, we separated the fractions reflecting TCAP and TDAP. The required input data matrices describe operational taxonomic units (OTUs) by traits, communities by the quantities or presence‐absence of these OTUs, and community sites by ecological variables. We defined plant functional types (PFTs) or species as community components after fuzzy weighting by the traits. The measured correlations for TCAP and TDAP were tested by permutation. The null model for TDAP preserves the trait convergence, the structure intrinsic in the fuzzy types, and community total abundances and autocorrelation. Results: We applied the method to trait‐based data from plant communities in south Brazil, one set in natural grassland experimental plots under different nitrogen and grazing levels, and another in sapling communities colonizing Araucaria forest patches of increasing size in a forest‐grassland mosaic. In these cases, depending on the traits considered, we found strong evidence of either TCAP or TDAP, or both, that was related to the environmental gradients. Conclusions: The method developed is able to reveal TCAP and TDAP that are more likely to be functional for specified ecological gradients, allowing establishment of objective hypotheses on their links to community processes.     

Both traits and phylogenetic history influence community structure in snakes over steep environmental gradients

Assemblages of closely related organisms are generated on axes of deep time diversification, biogeographic processes related to dispersal and habitat filtering, and competition. Using models that account for phylogeny, ecology, and traits, we examine how the interaction among biogeography, habitat filtering, and trait convergence influences community assemblage in Nearctic snakes. With 122 community surveys, environmental niche, trait data including size, diet, parity and habitat preference, and a nearly complete phylogeny of snakes from the United States, we ask 1) do phylogenetic species variability (PSV) and traits change in predictable and correlated ways given ecology and geographic distance, 2) are the measured traits variable within and across communities and how is this related to PSV at local scales, and 3) is there evidence of habitat filtering or trait divergence? Following a general trend of western to eastern North American origin and dispersal of major groups, we similarly show a significant decrease in PSV in this direction but unexpectedly with stable trait variance, showing that traits and phylogenetic variability are disconnected at the community level. We also demonstrate that trait variability and not PSV dominates local communities. Finally, regardless of phylogeny, we show that certain traits, such as reproductive mode (parity) frequency, change within communities in response to steep ecological gradients.     

Change in community phylogenetic structure during tropical forest succession: evidence from New Guinea

Ecologists have recently interpreted patterns of phylogenetic distance among coexisting species as indicative of processes affecting community assembly during forest succession. We investigated plant community phylogenetic structure along a successional gradient in New Guinean lowland rain forest. We surveyed all trees with diameter at breast height ≥ 5 cm in nineteen 0.25 ha plots representing younger secondary, older secondary, and primary forest. We estimated plant community phylogeny from rbcL gene sequences to quantify change in phylogenetic structure during succession. Mean phylogenetic distance among co‐occurring trees increased with total basal area per plot, a proxy for forest age. Significant phylogenetic clustering was detected in secondary forest whereas primary forest was significantly over‐dispersed relative to null expectations. We examined the sensitivity of these patterns to various methods of branch length estimation and phylogenetic uncertainty. Power to detect community phylogenetic patterns when equal branch lengths were assumed was weak in comparison to direct molecular and time‐calibrated measures of divergence. Inferred change during forest succession was also robust to phylogenetic uncertainty so long as temporal information was incorporated in estimates of divergence. The observed patterns are consistent with processes of environmental filtering during tropical forest succession giving way to other processes in primary forests including density‐dependence.