Intercontinental trends in functional and phylogenetic structure of stream fish assemblages

Understanding of community assembly has been improved by phylogenetic and trait‐based approaches, yet there is little consensus regarding the relative importance of alternative mechanisms and few studies have been done at large geographic and phylogenetic scales. Here, we use phylogenetic and trait dispersion approaches to determine the relative contribution of limiting similarity and environmental filtering to community assembly of stream fishes at an intercontinental scale. We sampled stream fishes from five zoogeographic regions. Analysis of traits associated with habitat use, feeding, or both resulted in more occurrences of trait underdispersion than overdispersion regardless of spatial scale or species pool. Our results suggest that environmental filtering and, to a lesser extent, species interactions were important mechanisms of community assembly for fishes inhabiting small, low‐gradient streams in all five regions. However, a large proportion of the trait dispersion values were no different from random. This suggests that stochastic factors or opposing assembly mechanisms also influenced stream fish assemblages and their trait dispersion patterns. Local assemblages tended to have lower functional diversity in microhabitats with high water velocity, shallow water depth, and homogeneous substrates lacking structural complexity, lending support for the stress‐dominance hypothesis. A high prevalence of functional underdispersion coupled with phylogenetic underdispersion could reflect phylogenetic niche conservatism and/or stabilizing selection. These findings imply that environmental filtering of stream fish assemblages is not only deterministic, but also influences assemblage structure in a fairly consistent manner worldwide.     

Past is prologue: host community assembly and the risk of infectious disease over time

Infectious disease risk is often influenced by host diversity, but the causes are unresolved. Changes in diversity are associated with changes in community structure, particularly during community assembly; therefore, by incorporating change over time, host community assembly may provide a framework to resolve causation. In turn, community assembly can be driven by many processes, including resource enrichment. To test the hypothesis that community assembly causally links host diversity to future disease, we experimentally manipulated host diversity and resource supply to hosts, then allowed communities to assemble for 2 years (surveyed 2012–2014). Initially, host diversity increased disease. Subsequently, host diversity did not directly alter disease. However, host diversity determined the trajectory of host community assembly, altering colonisation by exotic host species and richness‐independent host phylogenetic diversity, which together reversed the initial increase in disease. Ultimately, incorporating the temporal dimension of community assembly revealed novel mechanisms linking host diversity to future disease.     

Spatial phylogenetics of the native woody plant species in Hainan,China

To better identify biodiversity hotspots for conservation on Hainan Island, a tropical island in southern China, we assessed spatial variation in phylogenetic diversity and species richness using 18,976 georeferenced specimen records and a newly reconstructed molecular phylogeny of 957 native woody plants. Within this framework, we delineated bioregions based on vegetation composition and mapped areas of neoendemism and paleoendemism to identify areas of priority for conservation. Our results reveal that the southwest of Hainan is the most important hot spot for endemism and plant diversity followed by the southeast area. The distribution of endemic species showed a scattered, rather than clustered, pattern on the island. Based on phylogenetic range‐weighted turnover metrics, we delineated three major vegetational zones in Hainan. These largely correspond to natural secondary growth and managed forests (e.g., rubber and timber forests) in central Hainan, old‐growth forests and natural secondary growth forest at the margins of Hainan, and nature reserves on the island (e.g., Jianfeng and Diaoluo National Nature Reserves). Our study helps to elucidate potential botanical conservation priorities for Hainan within an evolutionary, phylogenetic framework.     

功能多样性和谱系多样性对热带云雾林群落构建的影响

近年来, 功能多样性和谱系多样性为探究群落构建机制提供了新方法。为了更准确地了解海南岛高海拔热带云雾林群落构建机制, 该研究以海南岛霸王岭热带云雾林为对象, 测定7个环境因子和13个植物功能性状。利用主成分分析(PCA)筛选环境因子, 以霸王岭、尖峰岭和黎母山热带云雾林分布物种建立区域物种库, 结合模型, 分析Rao二次熵(RaoQ)和平均成对谱系距离(MPD)变化对植物群落构建的影响。结果表明: 林冠开阔度、土壤全磷含量和坡度是影响植物群落构建的关键环境因子。多数功能性状的系统发育信号很低且不显著, 说明热带云雾林群落的系统发育关系与功能性状随历史进程变化不一致。RaoQ和MPD的实际观测值都显著低于期望值, 且其标准效应值与土壤磷含量显著相关, 说明生境过滤是驱动热带云雾林群落构建的关键因子, 土壤磷含量是群落构建的关键环境筛。     

The partitioning of diversity: showing Theseus a way out of the labyrinth

A methodology for partitioning of biodiversity into α, β and γ components has long been debated, resulting in different mathematical frameworks. Recently, use of the Rao quadratic entropy index has been advocated since it allows comparison of various facets of diversity (e.g. taxonomic, phylogenetic and functional) within the same mathematical framework. However, if not well implemented, the Rao index can easily yield biologically meaningless results and lead into a mathematical labyrinth. As a practical guideline for ecologists, we present a critical synthesis of diverging implementations of the index in the recent literature and a new extension of the index for measuring β‐diversity. First, we detail correct computation of the index that needs to be applied in order not to obtain negative β‐diversity values, which are ecologically unacceptable, and elucidate the main approaches to calculate the Rao quadratic entropy at different spatial scales. Then, we emphasize that, similar to other entropy measures, the Rao index often produces lower‐than‐expected β‐diversity values. To solve this, we extend a correction based on equivalent numbers, as proposed by Jost (2007), to the Rao index. We further show that this correction can be applied to additive partitioning of diversity and not only its multiplicative form. These developments around the Rao index open up an exciting avenue to develop an estimator of turnover diversity across different environmental and temporal scales, allowing meaningful comparisons of partitioning across species, phylogenetic and functional diversities within the same mathematical framework. We also propose a set of R functions, based on existing developments, which perform different key computations to apply this framework in biodiversity science.     

Functional diversity: a tool for answering challenging ecological questions

Functional trait diversity provides a powerful means of addressing ecology's persistent questions, through its dual role as an indicator of mechanisms driving differences in species composition between communities and as a predictor of ecosystem‐level processes. Functional traits provide a means of testing mechanisms behind species turnover between communities because environmental heterogeneity, competition and disturbance influence species fitness via their traits. Functional traits also provide a link between species and multiple ecosystem‐level processes, such as primary productivity, nutrient fluxes and resilience, since species influence these processes via their traits. This special issue demonstrates that functional diversity offers a practical means of investigating ecology's persistent questions.     

Maximal ecological diversity exceeds evolutionary diversity in model ecosystems

Understanding community saturation is fundamental to ecological theory. While investigations of the diversity of evolutionary stable states (ESSs) are widespread, the diversity of communities that have yet to reach an evolutionary endpoint is poorly understood. We use Lotka–Volterra dynamics and trait-based competition to compare the diversity of randomly assembled communities to the diversity of the ESS. We show that, with a large enough founding diversity (whether assembled at once or through sequential invasions), the number of long-time surviving species exceeds that of the ESS. However, the excessive founding diversity required to assemble a saturated community increases rapidly with the dimension of phenotype space. Additionally, traits present in communities resulting from random assembly are more clustered in phenotype space compared to random, although still markedly less ordered than the ESS. By combining theories of random assembly and ESSs we bring a new viewpoint to both the saturation and random assembly literature.     

华北区域环境梯度上阔叶林构建模式及分布成因

森林群落的构建即多样性维持机制是当今生态学研究的热点问题。然而, 当前群落构建和群落多样性的研究多在间接梯度上进行, 而在水、热等影响物种在区域内定植的关键且直接的环境梯度上研究群落构建和多样性模式则鲜有报道。结合环境因子, 基于物种组成和谱系方法探讨不同群落的分布成因, 有助于解释群落构建过程中的关键问题。该研究基于华北森林群落调查数据和环境数据, 涉及7个省市区的29个以壳斗科、桦木科为优势种的群落, 探讨了直接环境梯度上的群落构建和多样性模式, 同时用典范对应分析研究了不同群落分布的环境解释。结果发现, 相似的群落具有相似的生境偏好, 相似的生境条件会形成物种组成相同或相似的群落。环境热量主导了本区域的谱系关系, 在年平均气温较低的地区, 群落构建主要表现为生境过滤的模式。此外, 随着年降水量的增加, 生境过滤作用逐渐增加。在温度梯度上, 谱系多样性表现为钟形模式, 而降水量的增加能导致谱系多样性的增加。     

Contrasting effects of space and environment on functional and phylogenetic dissimilarity in a tropical forest

Aims The evolutionary history and functional traits of species can illuminate ecological processes supporting coexistence in diverse forest communities. However, little has been done in decoupling the relative importance of these mechanisms on the turnover of phylogenetic and functional characteristics across life stages and spatial scales. Therefore, this study aims to estimate the contribution of environment and dispersal on the turnover of phylogenetic and functional diversity across life stages and spatial scales, in order to build a coherent picture of the processes responsible for species coexistence.     

Introductions do not compensate for functional and phylogenetic losses following extinctions in insular bird assemblages

The ratio of species extinctions to introductions has been comparable for many insular assemblages, suggesting that introductions could have ‘compensated’ for extinctions. However, the capacity for introduced species to replace ecological roles and evolutionary history lost following extinction is unclear. We investigated changes in bird functional and phylogenetic diversity in the wake of extinctions and introductions across a sample of 32 islands worldwide. We found that extinct and introduced species have comparable functional and phylogenetic alpha diversity. However, this was distributed at different positions in functional space and in the phylogeny, indicating a ‘false compensation’. Introduced and extinct species did not have equivalent functional roles nor belong to similar lineages. This makes it unlikely that novel island biotas composed of introduced taxa will be able to maintain ecological roles and represent the evolutionary histories of pre‐disturbance assemblages and highlights the importance of evaluating changes in alpha and beta diversity concurrently.     

Loss of forest cover and host functional diversity increases prevalence of avian malaria parasites in the Atlantic Forest

Host phylogenetic relatedness and ecological similarity are thought to contribute to parasite community assembly and infection rates. However, recent landscape level anthropogenic changes may disrupt host-parasite systems by impacting functional and phylogenetic diversity of host communities. We examined whether changes in host functional and phylogenetic diversity, forest cover, and minimum temperature influence the prevalence, diversity, and distributions of avian haemosporidian parasites (genera Haemoproteus and Plasmodium) across 18 avian communities in the Atlantic Forest. To explore spatial patterns in avian haemosporidian prevalence and taxonomic and phylogenetic diversity, we surveyed 2241 individuals belonging to 233 avian species across a deforestation gradient. Mean prevalence and parasite diversity varied considerably across avian communities and parasites responded differently to host attributes and anthropogenic changes. Avian malaria prevalence (termed herein as an infection caused by Plasmodium parasites) was higher in deforested sites, and both Plasmodium prevalence and taxonomic diversity were negatively related to host functional diversity. Increased diversity of avian hosts increased local taxonomic diversity of Plasmodium lineages but decreased phylogenetic diversity of this parasite genus. Temperature and host phylogenetic diversity did not influence prevalence and diversity of haemosporidian parasites. Variation in the diversity of avian host traits that promote parasite encounter and vector exposure (host functional diversity) partially explained the variation in avian malaria prevalence and diversity. Recent anthropogenic landscape transformation (reduced proportion of native forest cover) had a major influence on avian malaria occurrence across the Atlantic Forest. This suggests that, for Plasmodium, host phylogenetic diversity was not a biotic filter to parasite transmission as prevalence was largely explained by host ecological attributes and recent anthropogenic factors. Our results demonstrate that, similar to human malaria and other vector-transmitted pathogens, prevalence of avian malaria parasites will likely increase with deforestation.     

Community assembly after long-term fragmentation: a case study of tropical rainforest in Xishuangbanna,south-west China

Background: Tropical rainforests represent the most species-rich and at the same time the most fragmented terrestrial biome on Earth. Fragmentation of tropical rainforests is having wide-ranging consequences for the maintenance of local species diversity and community assembly patterns.

Aims: To examine floristic changes and changes in community phylogenetic structure in the forest fragment over the past five decades.

Methods: A new taxonomic diversity algorithm (within-family diversity) was developed to assess floristic changes in the forest fragment. Community phylogenetic structure was then compared before and after fragmentation.

Results: Taxonomic diversity changed greatly among families, with changes occurring randomly across the phylogeny. The forest fragment had higher phylogenetic diversity, higher mean pair-wise phylogenetic distance, but lower mean nearest-neighbour distance. The community phylogenetic structure has changed significantly from clustering to dispersion.

Conclusions: High species turnover occurred in the forest fragment. While shade-tolerant species have been lost, and ruderal and alien species have been added, overall phylogenetic diversity has increased with species being more phylogenetically distant. Competitive exclusion, which was related to the relatively drier conditions in the forest after fragmentation, led the plant community phylogenetic structure to be more dispersed.     

Plant functional assemblages as indicators of the resilience of grassland ecosystem service provision

Ecosystems provide a variety of ecosystem services (ES), which act as key linkages between social and ecological systems. ES respond spatially and temporally to abiotic and biotic variation, and to management. Thus, resistant and resilient ES provision is expected to remain within a stable range when facing disturbances. In this study, generic indicators to evaluate resistance, potential resilience and capacity for transformation of ES provision are developed and their relevance demonstrated for a mountain grassland system. Indicators are based on plant trait composition (i.e. functional composition) and abiotic parameters determining ES provision at community, meta-community and landscape scales. First the resistance of an ES is indicated by its normal operating range characterized by observed values under current conditions. Second its resilience is assessed by its potential operating range − under hypotheses of reassembly from the community’s species pool. Third its transformation potential is assessed for reassembly at meta-community and landscape scales. Using a state-and-transition model, possible management-related transitions between mountain grassland states were identified, and indicators calculated for two provisioning and two regulating ES. Overall, resilience properties varied across individual ES, supporting a focus on resilience of specific ES. The resilience potential of the two provisioning services was greater than for the two regulating services, both being linked to functional complementarity within communities. We also found high transformation potential reflecting functional redundancy among communities within each meta-community, and across meta-communities in the landscape. Presented indicators are promising for the projection of future ES provision and the identification of management options under environmental change.     

Vole disturbances and plant diversity in a grassland metacommunity

We studied the disturbance associated with prairie vole burrows and its effects on grassland plant diversity at the patch (1 m²) and metacommunity (>5 ha) scales. We expected vole burrows to increase patch-scale plant species diversity by locally reducing competition for resources or creating niche opportunities that increase the presence of fugitive species. At the metacommunity scale, we expected burrows to increase resource heterogeneity and have a community composition distinct from the matrix. We measured resource variables and plant community composition in 30 paired plots representing disturbed burrows and undisturbed matrix patches in a cool-season grassland. Vole disturbance affected the mean values of nine resource variables measured and contributed more to resource heterogeneity in the metacommunity than matrix plots. Disturbance increased local plant species richness, metacommunity evenness, and the presence and abundance of fugitive species. To learn more about the contribution of burrow and matrix habitats to metacommunity diversity, we compared community similarity among burrow and matrix plots. Using Sorenson’s similarity index, which considers only presence–absence data, we found no difference in community similarity among burrows and matrix plots. Using a proportional similarity index, which considers both presence–absence and relative abundance data, we found low community similarity among burrows. Burrows appeared to shift the identity of dominant species away from the species dominant in the matrix. They also allowed subordinate species to persist in higher abundances. The patterns we observed are consistent with several diversity-maintaining mechanisms, including a successional mosaic and alternative successional trajectories. We also found evidence that prairie voles may be ecosystem engineers.     

Species turnover (β‐diversity) in ectomycorrhizal fungi linked to uptake capacity

Ectomycorrhizal (EcM) fungal communities may be shaped by both deterministic and stochastic processes, potentially influencing ecosystem development and function. We evaluated community assembly processes for EcM fungi of Pseudotsuga menziesii among 12 sites up to 400 km apart in southwest British Columbia (Canada) by investigating species turnover (β‐diversity) in relation to soil nitrogen (N) availability and physical distance. We then examined functional traits for an N‐related niche by quantifying net fluxes of , and protons on excised root tips from three contrasting sites using a microelectrode ion flux measurement system. EcM fungal communities were well aligned with soil N availability and pH, with no effect of site proximity (distance–decay curve) on species assemblages. Species turnover was significant (β_1/2 = 1.48) along soil N gradients, with many more Tomentella species on high N than low N soils, in contrast to Cortinarius species. Ammonium uptake was greatest in the spring on the medium and rich sites and averaged over 190 nmol/m²/s for Tomentella species. The lowest uptake rates of were by nonmycorrhizal roots of axenically grown seedlings (10 nmol/m²/s), followed by Cortinarius species (60 nmol/m²/s). EcM roots from all sites displayed only marginal uptake of nitrate (8.3 nmol/m²/s). These results suggest uptake capacity is an important functional trait influencing the assembly of EcM fungal communities. The diversity of EcM fungal species across the region arguably provides critical belowground adaptations to organic and inorganic N supply that are integral to temperate rainforest ecology.     

广西地区喀斯特与非喀斯特山地森林树木物种多样性及系统发育结构比较

以广西地区植被调查集资料总共365个群落样方(样方面积在300—1000 m2,50%左右为400 m2)为基础,将样方分为海拔≤1000 m喀斯特和非喀斯特森林以及海拔1000 m的非喀斯特森林三类生境,探讨这3种生境森林物种丰富度和群落系统发育结构差异及其与年均温的关系。结果表明,海拔≤1000 m非喀斯特森林含有4种以上的科比海拔1000 m非喀斯特森林多24科,比喀斯特森林多18科,其中樟科、壳斗科、山茶科、杜鹃花科等都是两种非喀斯特森林生境的重要组成科;大戟科为喀斯特森林含有种数最多的科,其中出现在非喀斯特森林的山茶科和杜鹃花科等没有在喀斯特森林出现。森林物种多样性及系统发育指数与年均温不相关或相关不强;海拔≤1000 m的喀斯特与非喀斯特森林的Gleason物种多样性指数没有显著差异,而海拔1000 m的非喀斯特森林的Gleason指数比另外两种生境显著高。海拔≤1000 m非喀斯特森林系统发育多样性PD指数比中山地带的非喀斯特生境的森林高。喀斯特森林和海拔1000 m非喀斯特的绝大部分样地以及海拔≤1000 m的非喀斯特大部分样地都表现出系统发育结构聚集,推测生境过滤在广西地区喀斯特和非喀斯特森林群落构建中发挥着主要作用。但是喀斯特和海拔1000 m的非喀斯特两种生境森林群落发生系统发育发散的比例都比海拔≤1000 m的非喀斯特森林低,揭示生境过滤在这两个生境更为明显。