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

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

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.     

Community assembly along a species pool gradient: implications for multiple-scale patterns of species diversity

Various ecological processes influence patterns of species diversity at multiple spatial scales. One process that is potentially important but rarely considered is community assembly. I assembled model communities using species pools of differing size to examine how the history of community assembly may affect multi-scale diversity patterns. The model contained three scales at which diversity could be measured: local community, metacommunity, and species pool. Local species saturation occurred, as expected from the competition and predation built in the model. However, local communities did not become resistant to invasions except when the species pool was very small. Depending on dispersal rate and trophic level, the larger the species pool, the harder it was to predict which species invades which local community at a given time. Consequently, local-community dissimilarity maintained by assembly history increased linearly with pool size, even though local diversity was decoupled from pool size. These results have two implications for multi-scale diversity patterns. First, assembly history may provide an explanation for scale-dependent relationships between local and regional diversity: assembly causes the relationship to be curvilinear at one scale (local community), while linear at another (metacommunity). Second, assembly history influences how -diversity is partitioned into - and -diversity: assembly causes the relative contribution of to increase with pool size. Overall, this study suggests that community assembly history interacts with species pool size to regulate multi-scale patterns of species diversity.     

Life-history trait and functional diversity patterns of ground beetles and spiders along a coastal heathland successional gradient

We investigated patterns in species richness and diversity, life-history traits and functional diversity of carabid beetles and spiders along a coastal heathland successional gradient. We sampled in five successional stages on the island of Hiddensee, Germany. Species richness of carabids and spiders and Simpson diversity for spiders did not differ among stages. Functional dispersion (FDis), a measure of functional richness, was lowest in the youngest stage for both carabids and spiders. Older successional stages represent more heterogeneous habitats and offer a broader range of niches, which might enhance functional dispersion. Functional evenness (FEve) differed among successional stages for spiders, indicating an uneven distribution of species abundances in the functional trait space. Functional divergence (FDiv) did not differ among stages for either taxon. Shifts in life-history traits were noted in both arthropod groups, but not always in the same direction: Body size of carabid beetles increased towards older successional stages, but decreased for spiders. The number of flightless carabid species increased towards older habitats. Each successional stage seemingly favors species with distinct life-history traits (species sorting). We conclude that a trait-based approach including the use of functional diversity measurements (FDis) could contribute valuable information for biodiversity conservation.     

青藏高原高寒草甸不同海拔梯度下土壤微生物群落碳代谢多样性

土壤微生物群落功能多样性对维持生态系统功能和稳定性具有非常重要的意义。为探究青藏高原高寒草甸不同海拔梯度下土壤微生物碳源利用差异以及影响机制,运用Biolog微平板技术,研究了西藏当雄县草原站4300—5100 m的6个不同海拔梯度下土壤微生物群落碳源代谢多样性。研究结果表明:(1)不同海拔下高寒草甸土壤微生物碳源的利用程度均随培养时间的延长而升高;微生物代谢活性和群落多样性指数均随海拔升高呈现先上升后下降的单峰变化趋势,整体表现4800 m4950 m4400 m4650 m5100 m4300 m;(2)主成分分析表明不同海拔显著影响了土壤微生物群落碳源代谢多样性,其中碳水化合物类、氨基酸类和胺类碳源是各海拔土壤微生物的偏好碳源;碳水化合物类、羧酸类、氨基酸类和胺类碳源的利用强度受海拔影响较大;(3)分类变异分析表明,土壤、植物和气候因素是影响不同海拔碳源利用变异的主要影响因子,可解释不同海拔的碳源利用差异的79.0%;排除环境因子之间的多重及交互作用,偏曼特尔检验表明土壤含水量、植被丰富度和年均降水量是影响不同海拔微生物碳源利用多样性的最重要的环境因子。综上,研究表明青藏高寒草甸不同海拔土壤微生物碳源代谢多样性呈现显著的海拔差异趋势,其海拔差异主要受到土壤含水量、植被丰富度和年均降水量的影响。     

Using functional trait diversity to infer community assembly mechanisms: an exclosure experiment as an example

AimsThe Qinghai-Tibetan Plateau has a mean altitude exceeding 4000 m and covers about 2.5 million km2. More than 60% of this area is alpine grassland. Exclosures have been widely used in this region to study the sustainable use of grassland resources. We used patterns of functional trait diversity to infer the effects of exclosures on community assembly in alpine meadows.     

Beta‐diversity of central European forests decreases along an elevational gradient due to the variation in local community assembly processes

Beta‐diversity has been repeatedly shown to decline with increasing elevation, but the causes of this pattern remain unclear, partly because they are confounded by coincident variation in alpha‐ and gamma‐diversity. We used 8795 forest vegetation‐plot records from the Czech National Phytosociological Database to compare the observed patterns of beta diversity to null‐model expectations (beta‐deviation) controlling for the effects of alpha‐ and gamma‐diversity. We tested whether β‐diversity patterns along a 1200 m elevation gradient exclusively depend on the effect of varying species pool size, or also on the variation of the magnitude of community assembly mechanisms determining the distribution of species across communities (e.g. environmental filtering, dispersal limitation). The null model we used is a novel extension of an existing null‐model designed for presence/absence data and was specifically designed to disrupt the effect of community assembly mechanisms, while retaining some key features of observed communities such as average species richness and species abundance distribution. Analyses were replicated in ten subregions with comparable elevation ranges. Beta‐diversity declined along the elevation gradient due to a decrease in gamma‐diversity, which was steeper than the decrease in alpha‐diversity. This pattern persisted after controlling for alpha‐ and gamma‐diversity variation, and the results were robust when different resampling schemes and diversity metrics were used. We conclude that in temperate forests the pattern of decreasing beta‐diversity with elevation does not exclusively depend on variation in species pool size, as has been hypothesized, but also on variation in community assembly mechanisms. The results were consistent across resampling schemes and diversity measures, thus supporting the use of vegetation‐plot databases for understanding patterns of beta‐diversity at the regional scale.     

The microbial gene diversity along an elevation gradient of the Tibetan grassland

Tibet is one of the most threatened regions by climate warming, thus understanding how its microbial communities function may be of high importance for predicting microbial responses to climate changes. Here, we report a study to profile soil microbial structural genes, which infers functional roles of microbial communities, along four sites/elevations of a Tibetan mountainous grassland, aiming to explore the potential microbial responses to climate changes via a strategy of space-for-time substitution. Using a microarray-based metagenomics tool named GeoChip 4.0, we showed that microbial communities were distinct for most but not all of the sites. Substantial variations were apparent in stress, N and C-cycling genes, but they were in line with the functional roles of these genes. Cold shock genes were more abundant at higher elevations. Also, gdh converting ammonium into urea was more abundant at higher elevations, whereas ureC converting urea into ammonium was less abundant, which was consistent with soil ammonium contents. Significant correlations were observed between N-cycling genes (ureC, gdh and amoA) and nitrous oxide flux, suggesting that they contributed to community metabolism. Lastly, we found by Canonical correspondence analysis, Mantel tests and the similarity tests that soil pH, temperature, NH₄⁺–N and vegetation diversity accounted for the majority (81.4%) of microbial community variations, suggesting that these four attributes were major factors affecting soil microbial communities. On the basis of these observations, we predict that climate changes in the Tibetan grasslands are very likely to change soil microbial community functional structure, with particular impacts on microbial N-cycling genes and consequently microbe-mediated soil N dynamics.     

Deterministic tropical tree community turnover: evidence from patterns of functional beta diversity along an elevational gradient

Explaining the mechanisms that produce the enormous diversity within and between tropical tree communities is a pressing challenge for plant community ecologists. Mechanistic hypotheses range from niche-based deterministic to dispersal-based stochastic models. Strong tests of these hypotheses require detailed information regarding the functional strategies of species. A few tropical studies to date have examined trait dispersion within individual forest plots using species trait means in order to ask whether coexisting species tend to be more or less functionally similar than expected given a null model. The present work takes an alternative approach by: (i) explicitly incorporating population-level trait variability; and (ii) quantifying the functional beta diversity in a series of 15 tropical forest plots arrayed along an elevational gradient. The results show a strong pattern of decay in community functional similarity with elevation. These observed patterns of functional beta diversity are shown to be highly non-random and support a deterministic model of tropical tree community assembly and turnover.     

Using phylogeny and functional traits for assessing community assembly along environmental gradients: A deterministic process driven by elevation

Community assembly processes is the primary focus of community ecology. Using phylogenetic‐based and functional trait‐based methods jointly to explore these processes along environmental gradients are useful ways to explain the change of assembly mechanisms under changing world. Our study combined these methods to test assembly processes in wide range gradients of elevation and other habitat environmental factors. We collected our data at 40 plots in Taibai Mountain, China, with more than 2,300 m altitude difference in study area and then measured traits and environmental factors. Variance partitioning was used to distinguish the main environment factors leading to phylogeny and traits change among 40 plots. Principal component analysis (PCA) was applied to colligate other environment factors. Community assembly patterns along environmental gradients based on phylogenetic and functional methods were studied for exploring assembly mechanisms. Phylogenetic signal was calculated for each community along environmental gradients in order to detect the variation of trait performance on phylogeny. Elevation showed a better explanatory power than other environment factors for phylogenetic and most traits’ variance. Phylogenetic and several functional structure clustered at high elevation while some conserved traits overdispersed. Convergent tendency which might be caused by filtering or competition along elevation was detected based on functional traits. Leaf dry matter content (LDMC) and leaf nitrogen content along PCA 1 axis showed conflicting patterns comparing to patterns showed on elevation. LDMC exhibited the strongest phylogenetic signal. Only the phylogenetic signal of maximum plant height showed explicable change along environmental gradients. Synthesis. Elevation is the best environment factors for predicting phylogeny and traits change. Plant's phylogenetic and some functional structures show environmental filtering in alpine region while it shows different assembly processes in middle‐ and low‐altitude region by different trait/phylogeny. The results highlight deterministic processes dominate community assembly in large‐scale environmental gradients. Performance of phylogeny and traits along gradients may be independent with each other. The novel method for calculating functional structure which we used in this study and the focus of phylogenetic signal change along gradients may provide more useful ways to detect community assembly mechanisms.     

Patterns of macromycete community assemblage along an elevation gradient: options for fungal gradient and metacommunity analyse

Gradient analysis is rarely used in studies of fungal communities. Data on macromycetes from eight sites along an elevation gradient in central Veracruz, Mexico, were used to demonstrate methods for gradient analysis that can be applied to studies of communities of fungi. Selected sites from 100 to 3,500?m altitude represent tropical dry forest, tropical montane cloud forest, conifer forest, and their ecotones. From May to October 2010, macromycetes were collected monthly within ten 10?×?10?m permanent plots per site. In total, 672 individuals of 213 species of macromycetes were recorded. Models for richness and diversity for all macromycete and ectomycorrhizal communities displayed peaks in the mid-part of the gradient, and a tendency to increase with elevation, whereas xylophagous fungi displayed a peak in the mid-lower part but tended to decrease with elevation. Cluster and Maximum Likelihood analyses distinguished four communities for both macromycetes and trees, but plant and fungal communities were only partly concordant. Canonical correspondence analysis indicated that macromycete distribution along the gradient is related to slope, relative humidity, soil temperature, soil water content, canopy openness, and litter depth. Spearman’s correlation and regression trees suggested that air and soil temperature, relative humidity, soil water content, canopy openness, vegetation structure and tree species richness were most strongly related to macrofungal functional groups, but these environmental variables were often correlated to the forest type and may not be causal. Variation in the environment along the elevation gradient differentially affected macromycete functional groups. Results from the different methods used in this work were concordant and showed significant patterns.     

Early subtropical forest growth is driven by community mean trait values and functional diversity rather than the abiotic environment

While functional diversity (FD) has been shown to be positively related to a number of ecosystem functions including biomass production, it may have a much less pronounced effect than that of environmental factors or species‐specific properties. Leaf and wood traits can be considered particularly relevant to tree growth, as they reflect a trade‐off between resources invested into growth and persistence. Our study focussed on the degree to which early forest growth was driven by FD, the environment (11 variables characterizing abiotic habitat conditions), and community‐weighted mean (CWM) values of species traits in the context of a large‐scale tree diversity experiment (BEF‐China). Growth rates of trees with respect to crown diameter were aggregated across 231 plots (hosting between one and 23 tree species) and related to environmental variables, FD, and CWM, the latter two of which were based on 41 plant functional traits. The effects of each of the three predictor groups were analyzed separately by mixed model optimization and jointly by variance partitioning. Numerous single traits predicted plot‐level tree growth, both in the models based on CWMs and FD, but none of the environmental variables was able to predict tree growth. In the best models, environment and FD explained only 4 and 31% of variation in crown growth rates, respectively, while CWM trait values explained 42%. In total, the best models accounted for 51% of crown growth. The marginal role of the selected environmental variables was unexpected, given the high topographic heterogeneity and large size of the experiment, as was the significant impact of FD, demonstrating that positive diversity effects already occur during the early stages in tree plantations.     

From diversity indices to community assembly processes: a test with simulated data

Ecological theory suggests that spatial distribution of biodiversity is strongly driven by community assembly processes. Thus the study of diversity patterns combined with null model testing has become increasingly common to infer assembly processes from observed distributions of diversity indices. However, results in both empirical and simulation studies are inconsistent. The aim of our study is to determine with simulated data which facets of biodiversity, if any, may unravel the processes driving its spatial patterns, and to provide practical considerations about the combination of diversity indices that would produce significant and congruent signals when using null models. The study is based on simulated species’ assemblages that emerge under various landscape structures in a spatially explicit individual‐based model with contrasting, predefined assembly processes. We focus on four assembly processes (species‐sorting, mass effect, neutral dynamics and competition colonization trade‐off) and investigate the emerging species’ distributions with varied diversity indices (alpha, beta and gamma) measured at different spatial scales and for different diversity facets (taxonomic, functional and phylogenetic). We find that 1) the four assembly processes result in distinct spatial distributions of species under any landscape structure, 2) a broad range of diversity indices allows distinguishing between communities driven by different assembly processes, 3) null models provide congruent results only for a small fraction of diversity indices and 4) only a combination of these diversity indices allows identifying the correct assembly processes. Our study supports the inference of assembly processes from patterns of diversity only when different types of indices are combined. It highlights the need to combine phylogenetic, functional and taxonomic diversity indices at multiple spatial scales to effectively infer underlying assembly processes from diversity patterns by illustrating how combination of different indices might help disentangling the complex question of coexistence.     

Soil fungal diversity and assembly along a xeric stress gradient in the central Namib Desert

The Namib Desert of south-western Africa is one of the oldest deserts in the world and possesses unique geographical, biological and climatic features. While research through the last decade has generated a comprehensive survey of the prokaryotic communities in Namib Desert soils, little is yet known about the diversity and function of edaphic fungal communities, and even less of their responses to aridity. In this study, we have characterized soil fungal community diversity across the longitudinal xeric gradient across the Namib desert (for convenience, divided into the western fog zone, the central low-rainfall zone and the eastern high-rainfall zone), using internal transcribed sequence (ITS) metabarcoding. Ascomycota, Basidiomycota and Chytridiomycota consistently dominated the Namib Desert edaphic fungal communities and a core mycobiome composed of only 15 taxa, dominated by members of the class Dothideomycetes (Ascomycota), was identified. However, fungal community structures were significantly different in the fog, low-rainfall and high-rainfall zones. Furthermore, Namib Desert gravel plain fungal community assembly was driven by both deterministic and stochastic processes; the latter dominating in the all three xeric zones. We also present data that suggest that the inland limit of fog penetration represents an ecological barrier to fungal dispersal across the Namib Desert.