Stochastic and deterministic assembly processes in subsurface microbial communities

A major goal of microbial community ecology is to understand the forces that structure community composition. Deterministic selection by specific environmental factors is sometimes important, but in other cases stochastic or ecologically neutral processes dominate. Lacking is a unified conceptual framework aiming to understand why deterministic processes dominate in some contexts but not others. Here we work toward such a framework. By testing predictions derived from general ecological theory we aim to uncover factors that govern the relative influences of deterministic and stochastic processes. We couple spatiotemporal data on subsurface microbial communities and environmental parameters with metrics and null models of within and between community phylogenetic composition. Testing for phylogenetic signal in organismal niches showed that more closely related taxa have more similar habitat associations. Community phylogenetic analyses further showed that ecologically similar taxa coexist to a greater degree than expected by chance. Environmental filtering thus deterministically governs subsurface microbial community composition. More importantly, the influence of deterministic environmental filtering relative to stochastic factors was maximized at both ends of an environmental variation gradient. A stronger role of stochastic factors was, however, supported through analyses of phylogenetic temporal turnover. Although phylogenetic turnover was on average faster than expected, most pairwise comparisons were not themselves significantly non-random. The relative influence of deterministic environmental filtering over community dynamics was elevated, however, in the most temporally and spatially variable environments. Our results point to general rules governing the relative influences of stochastic and deterministic processes across micro- and macro-organisms.     

The role of recruitment and dispersal limitation in tree community assembly in Amazonian forests

Background: Species composition of plant communities is shaped by the interplay between dispersal limitation, environmental filters and stochastic events.

Aims: The aim of this work was to investigate the effects of dispersal limitation and environmental filtering on tree recruitment. To accomplish this, we employed the unified neutral theory of biodiversity and biogeography to examine migration within the metacommunity, defined as a set of interacting local communities linked by the dispersal of multiple potentially interacting species.

Methods: We sampled 12,975 individuals with dbh ≥ 1 cm in 26 1-ha permanent plots, including habitats of terra firme, transitional forests, várzea and campinarana, on the upper Madeira River, Brazilian Amazon.

Results: Campinarana drew individuals from outside the metacommunity species pool at a mean probability of recruitment of 0.06, a much lower probability than terra firme (0.31), transitional (0.21) and várzea forests (0.22). Environmental variables, such as water table depth, soil texture and fertility, were related to differences in community assembly.

Conclusions: Species abundance distribution and diversity patterns of plant assemblages in a large river landscape in the Amazon highlight the importance of environmental heterogeneity that conditions beta-diversity. The high variation in recruitment probabilities from the metacommunity species pool to local communities suggests high habitat variability in the process of maintaining patterns of local diversity.     

黄土高原腹地人工林下草本层群落构建机制

群落构建机制是生态学研究的核心论题。生态位理论和中性理论是阐明群落构建的两种主要理论, 但这两种理论分别强调的环境过滤和扩散限制对群落构建的影响尚存争议。该研究以黄土高原人工林下草本层群落为研究对象, 将群落物种组成数据与地理空间、气候、地形以及生物因子相结合, 运用随机森林模型(RF)和基于距离矩阵的多元回归方法(MRM), 探究了各个因子的重要性, 并通过邻体主坐标矩阵(PCNM)和基于距离的冗余分析(db-RDA)分别筛选显著的空间结构和环境因子, 最后结合筛选出的PCNM特征值和环境变量进行变差分解。研究结果显示: 1)林下草本层群落组成相似性随地理距离和生境差异的增加而减小。2)地理距离与生境差异共同解释了群落组成相似性变化的47.8%。其中, 空间因子和环境因子的独立解释率分别为14.1%和9.8%, 两者的联合解释率为23.9%。3)地理距离和年降水量是引起研究区内物种组成变化的关键因子, 且地理距离的重要性大于年降水量。在黄土高原腹地, 扩散限制与环境过滤共同主导了人工林下草本层群落的构建过程。     

地理距离及环境差异对云南元江干热河谷植物群落beta多样性的影响

beta多样性反映了群落间物种组成的差异, 是生物多样性研究的热点之一。本研究通过对云南元江干热河谷41个植物群落样方进行调查, 用Jaccard相异系数表征物种beta多样性, 用样方之间的最近谱系距离(mean nearest taxon distance, MNTD)及平均谱系距离(mean pairwise distance, MPD)表征谱系beta多样性, 采用基于距离矩阵的多元回归和方差分解方法, 探讨了该区域干热河谷典型植物群落的物种beta多样性和谱系beta多样性与样方间环境差异(主要是气候)及地理距离之间的关系。结果表明: (1)群落间的地理距离和年平均温度差异对干热河谷植物群落的物种beta多样性和谱系beta多样性有显著影响; (2)地理距离对物种beta多样性和MNTD的影响最大; 地理距离和年平均温度差异对MPD的影响均较大; (3)样方间年平均温度与年平均降水量的差异和地理距离能够解释群落间beta多样性及谱系beta多样性11-13%的变异。以上结果表明, 生态位分化和扩散限制对该地区植物群落的beta多样性均有显著影响, 其中扩散限制的影响可能更大。此外, 人类活动等其他因素也很可能对元江干热河谷的群落组成具有非常重要的影响。     

Biodiversity dynamics in isolated island communities: interaction between natural and human-mediated processes

The flora and fauna of oceanic islands have inspired research since the early scientific explorations. Islands can be considered 'nature's test tubes'- simple systems with multiple replicates. Our research has used the simplicity of island systems to understand ecological community dynamics and to compare the properties of island communities with those in more complex mainland systems. Here, we present three topics: (i) current patterns of biodiversity on isolated islands of the Pacific; (ii) current patterns of disturbance and invasion on islands; and (iii) future trajectories inferred from these patterns. We examine features of islands (in particular, topography and isolation) that have allowed for given levels and distribution of endemicity. The extent to which island communities are impacted by, resist or accommodate disturbance and/or invasions by nonindigenous species appears to be dictated to a large extent by properties of the native communities and how these communities were originally assembled. Accordingly, patterns of disturbance and invasion are very different for high (montane) islands that are extremely isolated compared to those that are nearer to a source of natural migrants. As with all biotas, those on islands are dynamic entities. However, the unique aspect of islands is their isolation, and extreme isolation has largely been lost over the course of the last few centuries due to the development of transportation routes. We argue that such a modified dynamic will affect the future of the biota and the processes that gave rise to the biota. Specifically for isolated habitats, ecological processes will become increasingly more likely to generate biodiversity than evolutionary processes which have been relatively more important in the past. In the short term, island biotas and other similar biotas that occur in montane habitats may fare well as species are often abundant locally in the habitat to which they are indigenous, and may demonstrate considerable resistance and resilience to invasion. However, island biotas - and other biotas that show high local endemism - will likely not fare well in the face of prolonged disturbance. The biotas in these areas generally display a relatively low dispersal capacity; therefore, under conditions of long-term habitat modification, isolated biotas are likely to be swamped by non-natives, which - simply because of random processes and higher propagule pressure - will move more readily into available habitats. Thus, despite the importance of incorporating the evolutionary process into conservation efforts, we must also be careful to evaluate the likely form that the processes will take when the context (specifically, extent of isolation) has been highly modified.     

Cryptic diversity in a fig wasp community—morphologically differentiated species are sympatric but cryptic species are parapatric

A key debate in ecology centres on the relative importance of niche and neutral processes in determining patterns of community assembly with particular focus on whether ecologically similar species with similar functional traits are able to coexist. Meanwhile, molecular studies are increasingly revealing morphologically indistinguishable cryptic species with presumably similar ecological roles. Determining the geographic distribution of such cryptic species provides opportunities to contrast predictions of niche vs. neutral models. Discovery of sympatric cryptic species increases alpha diversity and supports neutral models, while documentation of allopatric/parapatric cryptic species increases beta diversity and supports niche models. We tested these predictions using morphological and molecular data, coupled with environmental niche modelling analyses, of a fig wasp community along its 2700‐km latitudinal range. Molecular methods increased previous species diversity estimates from eight to eleven species, revealing morphologically cryptic species in each of the four wasp genera studied. Congeneric species pairs that were differentiated by a key morphological functional trait (ovipositor length) coexisted sympatrically over large areas. In contrast, morphologically similar species, with similar ovipositor lengths, typically showed parapatric ranges with very little overlap. Despite parapatric ranges, environmental niche models of cryptic congeneric pairs indicate large regions of potential sympatry, suggesting that competitive processes are important in determining the distributions of ecologically similar species. Niche processes appear to structure this insect community, and cryptic diversity may typically contribute mostly to beta rather than alpha diversity.     

Reconciling the contribution of environmental and stochastic structuring of tropical forest diversity through the lens of imaging spectroscopy

Both niche and stochastic dispersal processes structure the extraordinary diversity of tropical plants, but determining their relative contributions has proven challenging. We address this question using airborne imaging spectroscopy to estimate canopy β‐diversity for an extensive region of a Bornean rainforest and challenge these data with models incorporating niches and dispersal. We show that remotely sensed and field‐derived estimates of pairwise dissimilarity in community composition are closely matched, proving the applicability of imaging spectroscopy to provide β‐diversity data for entire landscapes of over 1000 ha containing contrasting forest types. Our model reproduces the empirical data well and shows that the ecological processes maintaining tropical forest diversity are scale dependent. Patterns of β‐diversity are shaped by stochastic dispersal processes acting locally whilst environmental processes act over a wider range of scales.     

Geographical ecology of dry forest tree communities in the West Indies

Aim

Seasonally dry tropical forest (SDTF) of the Caribbean Islands (primarily West Indies) is floristically distinct from Neotropical SDTF in Central and South America. We evaluate whether tree species composition was associated with climatic gradients or geographical distance. Turnover (dissimilarity) in species composition of different islands or among more distant sites would suggest communities structured by speciation and dispersal limitations. A nested pattern would be consistent with a steep resource gradient. Correlation of species composition with climatic variation would suggest communities structured by broad‐scale environmental filtering.

Location

The West Indies (The Bahamas, Cuba, Hispaniola, Jamaica, Puerto Rico, US Virgin Islands, Guadeloupe, Martinique, St. Lucia), Providencia (Colombia), south Florida (USA) and Florida Keys (USA).

Taxon

Seed plants—woody taxa (primarily trees).

Methods

We compiled 572 plots from 23 surveys conducted between 1969 and 2016. Hierarchical clustering of species in plots, and indicator species analysis for the resulting groups of sites, identified geographical patterns of turnover in species composition. Nonparametric analysis of variance, applied to principal components of bioclimatic variables, determined the degree of covariation in climate with location. Nestedness versus turnover in species composition was evaluated using beta diversity partitioning. Generalized dissimilarity modelling partitioned the effect of climate versus geographical distance on species composition.

Results

Despite a set of commonly occurring species, SDTF tree community composition was distinct among islands and was characterized by spatial turnover on climatic gradients that covaried with geographical gradients. Greater Antillean islands were characterized by endemic indicator species. Northern subtropical areas supported distinct, rather than nested, SDTF communities in spite of low levels of endemism.

Main conclusions

The SDTF species composition was correlated with climatic variation. SDTF on large Greater Antillean islands (Hispaniola, Jamaica and Cuba) was characterized by endemic species, consistent with their geological history and the biogeography of plant lineages. These results suggest that both environmental filtering and speciation shape Caribbean SDTF tree communities.     

Changes in assembly processes in soil bacterial communities following a wildfire disturbance

Although recent work has shown that both deterministic and stochastic processes are important in structuring microbial communities, the factors that affect the relative contributions of niche and neutral processes are poorly understood. The macrobiological literature indicates that ecological disturbances can influence assembly processes. Thus, we sampled bacterial communities at 4 and 16 weeks following a wildfire and used null deviation analysis to examine the role that time since disturbance has in community assembly. Fire dramatically altered bacterial community structure and diversity as well as soil chemistry for both time-points. Community structure shifted between 4 and 16 weeks for both burned and unburned communities. Community assembly in burned sites 4 weeks after fire was significantly more stochastic than in unburned sites. After 16 weeks, however, burned communities were significantly less stochastic than unburned communities. Thus, we propose a three-phase model featuring shifts in the relative importance of niche and neutral processes as a function of time since disturbance. Because neutral processes are characterized by a decoupling between environmental parameters and community structure, we hypothesize that a better understanding of community assembly may be important in determining where and when detailed studies of community composition are valuable for predicting ecosystem function.     

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.     

The legacy of human use in Amazonian palm communities along environmental and accessibility gradients

Aim

Palms are iconic and dominant elements of neotropical forests. In the Amazon region, palms have been used and managed by humans for food, material, medicine and other purposes for millennia. It is, however, debated to what extent the structure of modern palm communities reflects long-term human modification. Here, we investigate the complex interplay of ecological and societal factors that influence the distributions of both human-used and non-used palms in western Amazonia.

Location

Amazonia.

Time period

Present.

Major taxa studied

Palms (Arecaceae).

Methods

We used Bayesian hierarchical joint species distribution models to predict the distributions and environmental niche dimensions of 78 western Amazonian species, and to explore their relationships with their diversity of human uses and with specific uses (food, construction and medicine). The models were parameterized with a comprehensive set of field- and satellite-derived environmental predictors.

Results

Our results suggest that a combination of ecological and anthropogenic factors drive the present-day distributions of Amazonian palms. The modelled ecological niches of the species revealed use-related species-sorting along soil, climatic, accessibility and drainage gradients. We found peaks in the proportions of useful palms and their diversity of uses in fertile soils, close to rivers, and on floodplains. These are habitats favourable for human settlement, although they harbour naturally restricted palm species pools. We also found a negative correlation between predicted palm species richness and number of human uses across western Amazonia.

Main conclusions

Soil characteristics, accessibility, and species pool size all contribute to defining palm–human relationships. At the basin scale, the signature of human use on palm communities was predicted to be stronger in the species-poor south-west than in central-western Amazonia. Overall, we conclude that environmental conditions have influenced modern Amazonian palm distributions both directly and indirectly, by regulating human settlement patterns and natural resource use over extended time periods.     

Strong influence of regional species pools on continent-wide structuring of local communities

There is a long tradition in ecology of evaluating the relative contribution of the regional species pool and local interactions on the structure of local communities. Similarly, a growing number of studies assess the phylogenetic structure of communities, relative to that in the regional species pool, to examine the interplay between broad-scale evolutionary and fine-scale ecological processes. Finally, a renewed interest in the influence of species source pools on communities has shown that the definition of the source pool influences interpretations of patterns of community structure. We use a continent-wide dataset of local ant communities and implement ecologically explicit source pool definitions to examine the relative importance of regional species pools and local interactions for shaping community structure. Then we assess which factors underlie systematic variation in the structure of communities along climatic gradients. We find that the average phylogenetic relatedness of species in ant communities decreases from tropical to temperate regions, but the strength of this relationship depends on the level of ecological realism in the definition of source pools. We conclude that the evolution of climatic niches influences the phylogenetic structure of regional source pools and that the influence of regional source pools on local community structure is strong.     

Niche assembly of epiphytic bryophyte communities in the Guianas: a regional approach

Aim Epiphytic bryophyte communities of tropical forests show a gradient in species composition from the base to the top of the host trees, indicating a strong role of niche assembly. This pattern, however, has never been tested at a regional scale. The aim of this study was to test whether niche assembly, rather than dispersal limitation, predominantly drives species composition of bryophyte communities across large spatial scales. Location Three lowland forests in the Guianas: one near Saul, French Guiana; and two near Mabura Hill, Guyana. Methods Communities of epiphytic bryophytes were sampled from six different height zones of several trees in three lowland forests. We analysed the composition of these communities using detrended correspondence analysis in order to find the best explanatory variable for the variation in community composition. A multi‐response permutation procedure was used to test the significance of grouping communities by height zone. We conducted an indicator species analysis to classify species as specialists or generalists and then tested, through weighted averaging, if specialists would indeed maintain their preferred height zone across the Guianas. Results Community composition was explained mainly by height zone. The similarity among communities inhabiting the same height zone of trees, across a distance of up to 640 km, was higher than the similarity among communities established along the vertical gradient of a single standing tree (30–50 m). More than half (57%) of the species had a preferred height zone, and the preference was consistent: species occupied roughly the same height zone on host trees in the different localities. The three local communities investigated were found to belong to the same regional species pool. Main conclusions Throughout the Guianas, epiphytic bryophyte communities are drawn from the same regional species pool, and their composition is shaped by micro‐environmental conditions. The predominance of niche assembly over dispersal assembly rules is consistently found at both local and regional scales.     

Fine‐scale substrate heterogeneity in green roof plant communities: The constraint of size

Heterogeneity–diversity relationship (HDR) is commonly shown to be positive in accordance with classic niche processes. However, recent soil‐based studies have often found neutral and even negative HDRs. Some of the suggested reasons for this discrepancy include the lack of resemblance between manipulated substrate and natural settings, the treated areas not being large enough to contain species' root span, and finally limited‐sized plots may not sustain focal species’ populations over time. Vegetated green roofs are a growing phenomenon in many cities that could be an ideal testing ground for this problem. Recent studies have focused on the ability of these roofs to sustain stable and diverse plant communities and substrate heterogeneity that would increase niches on the roof has been proposed as a method to attain this goal. We constructed an experimental design using green roof experimental modules (4 m²) where we manipulated mineral and organic substrate component heterogeneity in different subplots (0.25 m²) within the experimental module while maintaining the total sum of mineral and organic components. A local annual plant community was seeded in the modules and monitored over three growing seasons. We found that plant diversity and biomass were not affected by experimentally created substrate heterogeneity. In addition, we found that different treatments, as well as specific subplot substrates, had an effect on plant community assemblages during the first year but not during the second and third years. Substrate heterogeneity levels were mostly unchanged over time. The inability to retain plant community composition over the years despite the maintenance of substrate differences supports the hypothesis that maintenance of diversity is constrained at these spatial scales by unfavorable dispersal and increased stochastic events as opposed to predictions of classic niche processes.     

The influence of historical dispersal on the phylogenetic structure of tree communities in the tropical Andes

We test for evidence of the Tropical Niche Conservatism or the Out of The Tropics hypotheses in structuring patterns of tree community composition along a 2000 + meter elevational gradient in the northern tropical Andes. By collecting and integrating data on the presence–absence of tree species within plots with phylogenetic information, we analyzed the following: (a) patterns of phylogenetic dispersion and species diversity along the elevational gradient based on indexes of net relatedness, nearest taxon relatedness, and species richness (α‐diversity); and (b) the replacement of lineages along the gradient using the PhyloSorensen metric (β‐diversity). More specifically, we established 20 0.25‐ha permanent tree inventory plots between 750 and 2,802 m asl where all individuals with diameter at breast height (DBH) ≥ 10 cm were measured and identified. We then used a series of linear models to test for changes in α and β diversity between plots in relation to elevation. Neither the net relatedness index nor the nearest taxon index showed a significant relationship with elevation. However, there was greater phylogenetic overdispersion at intermediate elevations; this likely reflects the mixing of species with contrasting origins from tropical and temperate lineages. β‐diversity between plots was negatively related to the corresponding difference in elevation, indicating that closely related lineages occupy similar ranges of elevation and temperature. We conclude that the immigration of lineages from extra‐tropical regions has significant effects in determining the phylogenetic structure of tree communities in tropical Andean forests. Abstract in Spanish is available with online material.     

Beyond climate control on species range: The importance of soil data to predict distribution of Amazonian plant species

Aim

To evaluate the relative importance of climatic versus soil data when predicting species distributions for Amazonian plants and to gain understanding of potential range shifts under climate change.

Location

Amazon rain forest.

Methods

We produced species distribution models (SDM) at 5‐km spatial resolution for 42 plant species (trees, palms, lianas, monocot herbs and ferns) using species occurrence data from herbarium records and plot‐based inventories. We modelled species distribution with Bayesian logistic regression using either climate data only, soil data only or climate and soil data together to estimate their relative predictive powers. For areas defined as unsuitable to species occurrence, we mapped the difference between the suitability predictions obtained with climate‐only versus soil‐only models to identify regions where climate and soil might restrict species ranges independently or jointly.

Results

For 40 out of the 42 species, the best models included both climate and soil predictors. The models including only soil predictors performed better than the models including only climate predictors, but we still detected a drought‐sensitive response for most of the species. Edaphic conditions were predicted to restrict species occurrence in the centre, the north‐west and in the north‐east of Amazonia, while the climatic conditions were identified as the restricting factor in the eastern Amazonia, at the border of Roraima and Venezuela and in the Andean foothills.

Main conclusions

Our results revealed that soil data are a more important predictor than climate of plant species range in Amazonia. The strong control of species ranges by edaphic features might reduce species’ abilities to track suitable climate conditions under a drought‐increase scenario. Future challenges are to improve the quality of soil data and couple them with process‐based models to better predict species range dynamics under climate change.