The balance between deterministic and stochastic processes in structuring lake bacterioplankton community over time

One major goal in microbial ecology is to establish the importance of deterministic and stochastic processes for community assembly. This is relevant to explain and predict how diversity changes at different temporal scales. However, understanding of the relative quantitative contribution of these processes and particularly of how they may change over time is limited. Here, we assessed the importance of deterministic and stochastic processes based on the analysis of the bacterial microbiome in one alpine oligotrophic and in one subalpine mesotrophic lake, which were sampled over two consecutive years at different time scales. We found that in both lakes, homogeneous selection (i.e., a deterministic process) was the main assembly process at the annual scale and explained 66.7% of the bacterial community turnover, despite differences in diversity and temporal variability patterns between ecosystems. However, in the alpine lake, homogenizing dispersal (i.e., a stochastic process) was the most important assembly process at the short‐term (daily and weekly) sampling scale and explained 55% of the community turnover. Alpha diversity differed between lakes, and seasonal stability of the bacterial community was more evident in the oligotrophic lake than in the mesotrophic one. Our results demonstrate how important forces that govern temporal changes in bacterial communities act at different time scales. Overall, our study validates on a quantitative basis, the importance and dominance of deterministic processes in structuring bacterial communities in freshwater environments over long time scales.     

Species abundance distributions as a proxy for the niche-neutrality continuum

Aims Species abundance distributions (SADs) are often used to verify mechanistic theories underlying community assembly. However, it is now accepted that SADs alone are not sufficient to reveal biological mechanisms. Recent attention focuses on the relative importance of stochastic dispersal processes versus deterministic processes such as interspecific competition and environmental filtering. Here, we combine a study of the commonness and rarity of species (i.e. the SAD) with mechanistic processes underlying community composition. By comparing the occurrence frequencies of each and every species with its abundance, we quantify the relative contributions of common and rare species to the maintenance of community structure. Essentially, we relate the continuum between commonness and rarity with that of niches and neutrality.Methods An individual-based, spatially explicit model was used to simulate local communities in niche spaces with the same parameters. We generated sets of assemblages from which species were eliminated in opposing sequences: from common to rare and from rare to common, and investigated the relationship between the abundance and frequency of species. We tested the predictions of our model with empirical data from a field experiment in the environmentally homogeneous alpine meadows of the Qinghai–Tibetan plateau.Important findings Our simulations support the widespread notion that common species maintain community structure, while rare species maintain species diversity, in both local and regional communities. Our results, both from theoretical simulations and from empirical observations, revealed positive correlations between the abundance of a particular species and its occurrence frequency. SAD curves describe a continuum between commonness and rarity. Removing species from the 'rare' end of this continuum has little effect on the similarity of communities, but removing species from the 'common' end of the continuum causes significant increases in beta diversity, or species turnover, between communities. In local communities distributed in a homogenous habitat, species located at the 'common' end of the continuum should be selected by environmental filtering, with niche space partitioning governed by interspecific competition. Conversely, species located at the 'rare' end of the continuum are most likely subject to stochastic dispersal processes. Species situated at intermediate locations on this continuum are therefore determined by niche and neutral processes acting together. Our results suggest that, in homogeneous habitats, SAD curves describing the common-rare continuum may also be used to describe the continuum between niches and neutrality.     

The scale dependence of fungal community distribution in paddy soil driven by stochastic and deterministic processes

Fungal communities play important roles in terrestrial ecosystem functioning. Unraveling the relative importance of stochastic versus deterministic processes in shaping biogeographic patterns of fungal communities has long been a challenge in microbial ecology, owing to high biodiversity and difficulties in identifying fungal taxa. Using a unique anthropogenic system of geographically isolated paddy ‘islands’, we collected 198 soil samples with a spatially explicit design to examine how ecological processes shaped fungal biogeographic patterns. Fungal community structure showed scale-dependent distance-decay relationships. Stochastic processes (dispersal and drift) contributed more to community assembly than deterministic processes (selection) at the local scale, which was largely attributed to drift. In contrast, deterministic processes contributed more to community assembly than stochastic processes at the regional scale, with soil dissolved organic carbon being the most important measured factor. Collectively, scale dependence of fungal biogeographical patterns in paddy soils is influenced by differential contribution of deterministic and stochastic processes.     

Environmental changes affect the assembly of soil bacterial community primarily by mediating stochastic processes

Both ‘species fitness difference’‐based deterministic processes, such as competitive exclusion and environmental filtering, and ‘species fitness difference’‐independent stochastic processes, such as birth/death and dispersal/colonization, can influence the assembly of soil microbial communities. However, how both types of processes are mediated by anthropogenic environmental changes has rarely been explored. Here we report a novel and general pattern that almost all anthropogenic environmental changes that took place in a grassland ecosystem affected soil bacterial community assembly primarily through promoting or restraining stochastic processes. We performed four experiments mimicking 16 types of environmental changes and separated the compositional variation of soil bacterial communities caused by each environmental change into deterministic and stochastic components, with a recently developed method. Briefly, because the difference between control and treatment communities is primarily caused by deterministic processes, the deterministic change was quantified as (mean compositional variation between treatment and control) – (mean compositional variation within control). The difference among replicate treatment communities is primarily caused by stochastic processes, so the stochastic change was estimated as (mean compositional variation within treatment) – (mean compositional variation within control). The absolute of the stochastic change was greater than that of the deterministic change across almost all environmental changes, which was robust for both taxonomic and functional‐based criterion. Although the deterministic change may become more important as environmental changes last longer, our findings showed that changes usually occurred through mediating stochastic processes over 5 years, challenging the traditional determinism‐dominated view.     

Point process models,the dimensions of biodiversity and the importance of small-scale biotic interactions

Aims Recent mechanistic explanations for community assembly focus on the debates surrounding niche-based deterministic and dispersal-based stochastic models. This body of work has emphasized the importance of both habitat filtering and dispersal limitation, and many of these works have utilized the assumption of species spatial independence to simplify the complexity of the spatial modeling in natural communities when given dispersal limitation and/or habitat filtering. One potential drawback of this simplification is that it does not consider species interactions and how they may influence the spatial distribution of species, phylogenetic and functional diversity. Here, we assess the validity of the assumption of species spatial independence using data from a subtropical forest plot in southeastern China.Methods We use the four most commonly employed spatial statistical models—the homogeneous Poisson process representing pure random effect, the heterogeneous Poisson process for the effect of habitat heterogeneity, the homogenous Thomas process for sole dispersal limitation and the heterogeneous Thomas process for joint effect of habitat heterogeneity and dispersal limitation—to investigate the contribution of different mechanisms in shaping the species, phylogenetic and functional structures of communities.Important findings Our evidence from species, phylogenetic and functional diversity demonstrates that the habitat filtering and/or dispersal-based models perform well and the assumption of species spatial independence is relatively valid at larger scales (50×50 m). Conversely, at local scales (10×10 and 20×20 m), the models often fail to predict the species, phylogenetic and functional diversity, suggesting that the assumption of species spatial independence is invalid and that biotic interactions are increasingly important at these spatial scales.     

Stochastic and deterministic processes drive wetland community assembly across a gradient of environmental filtering

The role of deterministic and stochastic processes in community assembly is a key question in community ecology. We evaluated the effect of an abiotic filter (hydroperiod) on the partitioned diversity of three taxonomic groups (birds, vegetation, macroinvertebrates) from prairie pothole wetlands in Alberta, Canada, which naturally vary in water permanence. We observed that alpha and gamma diversity were higher in permanent than temporary wetlands (16–25% and 34–47% respectively, depending on the taxon). This suggests an influence of deterministic constraints on the number of species a wetland can support. Taxa which cannot persist in shallow, temporary wetlands are excluded by the deterministic constraints that a shortened hydroperiod imposes. In contrast, we observed that beta diversity was significantly higher (2–12%) in temporary wetlands than permanent ones, and temporary wetlands supported more unique combinations of community composition than permanent wetlands, despite having a smaller regional species pool. This observation contradicts prior mesocosm studies that found beta diversity mirrored the pattern in gamma diversity along an environmental filtering gradient. We conclude that deterministic processes are more influential in more stable permanent wetlands, whereas stochastic processes play a more important role in assembly in dynamic temporary wetlands that must disassemble and re‐establish annually. Considering three distinct taxonomic groups differing in their relative mobility, our large‐scale field study demonstrates that both stochastic and deterministic processes act together to influence the assembly of multiple communities and that the relative importance of the two processes varies consistently along a gradient of environmental filtering.     

Relative influence of deterministic processes on structuring marsh plant communities varies across an abiotic gradient

Understanding the processes determining community structure is one of the major goals of ecological research. Both deterministic and stochastic processes may shape community structure. The challenge is to understand the relative influence of each type of process across different environmental conditions. We investigated the influence of deterministic and stochastic processes on plant community assembly in tidal marshes across a strong abiotic (salinity) gradient in three estuaries in Georgia, USA using probabilistic Raup–Crick community dissimilarity. Our results indicated that deterministic processes had an increasingly important influence on structuring plant communities in salt and brackish marshes, probably due to high heterogeneity of microhabitats produced by the interplay between abiotic stress and biotic interactions. In contrast, the influence of deterministic processes on plant community assembly decreased in tidal freshwater marshes, suggesting an increasingly important role of stochastic processes in plant community assembly in tidal freshwater marshes, probably due to the higher species richness, higher recruitment from seed, and lower levels of abiotic stress in these habitats. At the estuarine scale (across tidal freshwater, brackish and salt marshes in each estuary), our results suggested that deterministic processes also had a relatively important influence on shaping plant community structure. Our results illustrated that plant community structure in tidal marshes is influenced by both deterministic and stochastic processes, but that the relative influence of these two types of processes varies across estuarine landscapes.     

Balance of Neutral and Deterministic Components in the Dynamics of Activated Sludge Floc Assembly

Understanding the processes that generate patterns of community structure is a central focus of ecological research. With that aim, we manipulated the structure of bacterial activated sludge to test the influence of the species richness and composition of bacterial communities on the dynamics of activated sludge floc assembly in lab-scale bioreactors. Bacterial community structure was analyzed using denaturing gradient gel electrophoresis of RT-PCR amplified 16S rRNA. Fingerprinting of four parallel reactors, started with the same source communities added in different proportions, converged to patterns that were more similar than expected by chance, suggesting a deterministic selection in floc development. Evidence for neutral dynamics was suggested by the dependence of the rate of replacement of species (bacterial taxa–time relationships) on the number of available species in the source community. Further indication of stochastic dynamics was obtained by the application of the Sloan neutral model for prokaryotes. The fitting of the observed data to the model predictions revealed that the importance of the stochastic component increased with the size of the reservoir of species richness from which the community is drawn. Taken together, the results illustrate how both neutral and deterministic dynamics operate simultaneously in the assembly of the bacterial floc and show that the balance of the two depends on the richness of the source community.     

Disentangling the importance of ecological niches from stochastic processes across scales

Deterministic theories in community ecology suggest that local, niche-based processes, such as environmental filtering, biotic interactions and interspecific trade-offs largely determine patterns of species diversity and composition. In contrast, more stochastic theories emphasize the importance of chance colonization, random extinction and ecological drift. The schisms between deterministic and stochastic perspectives, which date back to the earliest days of ecology, continue to fuel contemporary debates (e.g. niches versus neutrality). As illustrated by the pioneering studies of Robert H. MacArthur and co-workers, resolution to these debates requires consideration of how the importance of local processes changes across scales. Here, we develop a framework for disentangling the relative importance of deterministic and stochastic processes in generating site-to-site variation in species composition (β-diversity) along ecological gradients (disturbance, productivity and biotic interactions) and among biogeographic regions that differ in the size of the regional species pool. We illustrate how to discern the importance of deterministic processes using null-model approaches that explicitly account for local and regional factors that inherently create stochastic turnover. By embracing processes across scales, we can build a more synthetic framework for understanding how niches structure patterns of biodiversity in the face of stochastic processes that emerge from local and biogeographic factors.     

Measuring biodiversity to explain community assembly: a unified approach

One of the oldest challenges in ecology is to understand the processes that underpin the composition of communities. Historically, an obvious way in which to describe community compositions has been diversity in terms of the number and abundances of species. However, the failure to reject contradictory models has led to communities now being characterized by trait and phylogenetic diversities. Our objective here is to demonstrate how species, trait and phylogenetic diversity can be combined together from large to local spatial scales to reveal the historical, deterministic and stochastic processes that impact the compositions of local communities. Research in this area has recently been advanced by the development of mathematical measures that incorporate trait dissimilarities and phylogenetic relatedness between species. However, measures of trait diversity have been developed independently of phylogenetic measures and conversely most of the phylogenetic diversity measures have been developed independently of trait diversity measures. This has led to semantic confusions particularly when classical ecological and evolutionary approaches are integrated so closely together. Consequently, we propose a unified semantic framework and demonstrate the importance of the links among species, phylogenetic and trait diversity indices. Furthermore, species, trait and phylogenetic diversity indices differ in the ways they can be used across different spatial scales. The connections between large‐scale, regional and local processes allow the consideration of historical factors in addition to local ecological deterministic or stochastic processes. Phylogenetic and trait diversity have been used in large‐scale analyses to determine how historical and/or environmental factors affect both the formation of species assemblages and patterns in species richness across latitude or elevation gradients. Both phylogenetic and trait diversity have been used at different spatial scales to identify the relative impacts of ecological deterministic processes such as environmental filtering and limiting similarity from alternative processes such as random speciation and extinction, random dispersal and ecological drift. Measures of phylogenetic diversity combine phenotypic and genetic diversity and have the potential to reveal both the ecological and historical factors that impact local communities. Consequently, we demonstrate that, when used in a comparative way, species, trait and phylogenetic structures have the potential to reveal essential details that might act simultaneously in the assembly of species communities. We highlight potential directions for future research. These might include how variation in trait and phylogenetic diversity alters with spatial distances, the role of trait and phylogenetic diversity in global‐scale gradients, the connections between traits and phylogeny, the importance of trait rarity and independent evolutionary history in community assembly, the loss of trait and phylogenetic diversity due to human impacts, and the mathematical developments of biodiversity indices including within‐species variations.     

Convergence of community composition during secondary succession on Zokor rodent mounds on the Tibetan Plateau

Aims The community succession theory is much debated in ecology. We studied succession on Zokor rodent mounds on the Tibetan Plateau to address several fundamental questions, among them: (i) During secondary succession, does the community composition converge towards one community state or multiple states depending on the initial colonization? (ii) Do mound communities located in different background communities exhibit different assembly trajectories?Methods In a sub-alpine meadow, we investigated a total of 80 mound communities at several successional stages in three different background communities resulting from different management histories and compared their changes in species composition. The distribution of plant communities over time was analyzed with quantitative classification and ordination methods. The co-occurrence patterns of species were evaluated at each successional stage, and the degree of convergence/divergence among communities was obtained by calculating two beta-diversity indices.Important findings During secondary succession, species richness of mound communities changed over time, and this change was dependent on the background community. Five life-form groups exhibited different dynamic patterns in species richness and plant cover. Community composition and the degree of species co-occurrence between communities increased over time since disturbance. There was much variation in species composition at earlier stages of succession, but communities on older mounds became more similar to each other and to their surrounding vegetation over the course of secondary succession. Post-disturbance succession of Zokor mound communities transitioned from 'multiple alternative states' to 'background-based deterministic community assembly' over time. Tradeoffs between competition and colonization, as well as the characteristics of different life-forms and mass effects within a limited species pool are the mechanisms responsible for convergence of mound communities.     

Community assembly of a euryhaline fish microbiome during salinity acclimation

Microbiomes play a critical role in promoting a range of host functions. Microbiome function, in turn, is dependent on its community composition. Yet, how microbiome taxa are assembled from their regional species pool remains unclear. Many possible drivers have been hypothesized, including deterministic processes of competition, stochastic processes of colonization and migration, and physiological ‘host‐effect’ habitat filters. The contribution of each to assembly in nascent or perturbed microbiomes is important for understanding host–microbe interactions and host health. In this study, we characterized the bacterial communities in a euryhaline fish and the surrounding tank water during salinity acclimation. To assess the relative influence of stochastic versus deterministic processes in fish microbiome assembly, we manipulated the bacterial species pool around each fish by changing the salinity of aquarium water. Our results show a complete and repeatable turnover of dominant bacterial taxa in the microbiomes from individuals of the same species after acclimation to the same salinity. We show that changes in fish microbiomes are not correlated with corresponding changes to abundant taxa in tank water communities and that the dominant taxa in fish microbiomes are rare in the aquatic surroundings, and vice versa. Our results suggest that bacterial taxa best able to compete within the unique host environment at a given salinity appropriate the most niche space, independent of their relative abundance in tank water communities. In this experiment, deterministic processes appear to drive fish microbiome assembly, with little evidence for stochastic colonization.     

Mechanisms of soil bacterial and fungal community assembly differ among and within islands

The study of islands has made substantial contributions to the development of evolutionary and ecological theory. However, we know little about microbial community assembly on islands. Using soil microbial data collected from 29 lake islands and nearby mainland, we examined the assembly mechanisms of soil bacterial and fungal communities among and within islands. We found that deterministic processes, especially homogeneous selection, tended to be more important in shaping the assembly of soil bacterial communities among islands, while stochastic processes tended to be more important within islands. Moreover, increasing island area increased the importance of homogeneous selection, but reduced the importance of variable selection, for soil bacterial community assembly within islands. By contrast, stochastic processes tended to dominate soil fungal community assembly both among and within islands, with dispersal limitation playing a more important role within than among islands. Our results highlight the scale- and taxon-dependence of insular soil microbial community assembly, suggesting that spatial scale should be explicitly considered when evaluating the influences of habitat fragmentation on soil microbial communities.     

Deterministic control on community assembly peaks at intermediate levels of disturbance

Despite long-standing research, the processes that drive community assembly remain poorly understood. We censused macroinvertebrate communities and measured flood disturbance in 17 Scandinavian mountain streams to assess the hypothesis that communities are shaped by stochastic processes under stable conditions, and increasingly by deterministic processes as disturbance becomes more severe. Each study stream was categorized as being stable (n=5), intermediate (n=7), or disturbed (n=5) depending on the severity of scouring floods. Following spring floods, the number of potential colonisers decreased with increasing disturbance, suggesting that disturbance filtered out species unable to cope with the stress involved. Communities at stable sites had the highest beta diversity, indicating that stochastic processes of community assembly were most important under the least disturbed conditions. In partial contrast with our predictions, the lowest beta diversity occurred between intermediate (not disturbed) sites, suggesting that increasing disturbance first enhances determinism but then rekindles stochasticity at severity levels beyond intermediate. Macroinvertebrate communities were shaped by deterministic processes, which recruit potential regional colonists depending on niche differences and disturbance conditions and by stochastic processes, which distribute the selected species randomly among individual localities. Although often considered opposing, stochastic and deterministic processes interact hierarchically, with relative strength modified by disturbance.     

Ecological uniqueness of macroinvertebrate communities in high-latitude streams is a consequence of deterministic environmental filtering processes

Variation in biological communities is a consequence of stochastic and deterministic factors. Examining the relative importance of these factors helps to understand variation in the whole biodiversity in a region. We examined the roles of stochastic and deterministic factors in structuring macroinvertebrate communities in high-latitude streams across two seasons. We predicted that if communities are the result of deterministic environmental filtering processes, the communities should show strong association with environmental variables, as taxa would be selected according to stream environmental conditions. However, if communities are driven by stochastic factors, they should show strong association with spatial variables, as the distribution of taxa in communities would be driven by spatially related dispersal factors. We studied these predictions by calculating the degree of uniqueness of the streams in terms of their taxonomic and functional community compositions and by modelling the resulting index values using spatial and environmental variables. Our results supported the first prediction where the communities are more influenced by the environmental filtering processes, although indications of the effect of spatial processes in structuring the communities were present especially in autumn. High-latitude stream communities also seem to be sensitive to environmental changes, as even small changes in environment were enough to affect the ecological uniqueness of the streams. These findings highlight the vulnerability of northern streams in the face of the climate change. To maintain biodiversity in high-latitude catchments, it would be important to protect varying habitat conditions, which are the main forces affecting the ecological uniqueness of the streams.     

Predators temper the relative importance of stochastic processes in the assembly of prey metacommunities

Communities assemble through a combination of stochastic processes, which can make environmentally similar communities divergent (high β-diversity), and deterministic processes, which can make environmentally similar communities convergent (low β-diversity). Top predators can influence both stochasticity (e.g. colonization and extinction events) and determinism (e.g. size of the realized species pool), in community assembly, and thus their net effect is unknown. We investigated how predatory fish influenced the scaling of prey diversity in ponds at local and regional spatial scales. While fish reduced both local and regional richness, their effects were markedly more intense at the regional scale. Underlying this result was that the presence of fish made localities within metacommunities more similar in their community composition (lower β-diversity), suggesting that fish enhance the deterministic, relative to the stochastic, components of community assembly. Thus, the presence of predators can alter fundamental mechanisms of community assembly and the scaling of diversity within metacommunities.     

Quantifying the role of deterministic assembly and stochastic drift in a natural community of Arctic mosses

The interpretation of natural plant communities frequently invokes species‐sorting controlled by niche differences along spatial environmental gradients. This process of niche structuring can be explained by reference to functional traits, which provide a mechanistic explanation for community structure. In contrast, models explaining species coexistence obviate the limiting effect of niche difference, by invoking processes which cause species‐level drift, e.g. demographic stochasticity. This paper investigates a simple habitat with strong gradients (moss communities in a patterned arctic wetland) to identify signature‐patterns under‐pinning the relative importance of deterministic assembly and stochastic drift in a natural community. First, ordination analysis was used to confirm community composition structured by a range of nine carefully selected functional traits. Second, to determine whether traits explaining community composition might also explain species richness, local species richness (sR) was compared to (1) observed trait diversity and (2) expected trait diversity based on permutation tests, which are used to simulate null community assembly for different values of sR. Traits explaining species composition, consistent with deterministic niche structuring, do not appear to maintain sR. This surprising result was explained by decomposing the community into individual pair‐wise comparisons, i.e. species niche‐differences and association (χ²). Results support deterministic processes via the sorting of species with similar and contrasting niches, at opposite ends of a composite environmental gradient. Nevertheless, stochastic drift is apparent in the random structure of a majority of pair‐wise associations; in addition, a species’ abundance was in general not related to environmental distance from response‐optima. We suggest therefore that spatial pattern in the moss community is a balance between deterministic forces with respect to species traits and controlling environmental gradients, and stochastic drift, which weakens this deterministic structure.     

Trait-based filtering from the regional species pool into local grassland communities

Aims For plants to establish in a local community from a pool of possible colonizers from the region, it must pass through a series of filters. Which of the filters is most important in this process has been much debated. In this study, we explored how species are filtered from the regional species pool into local communities. The aim was to determine if differences in species abundance and functional traits could explain which species from the regional species pool establish at the local scale and if the filtering differed between grassland communities.Methods This study took place in a cultivated landscape in southeastern Sweden. We estimated plant species abundance in 12 ex-arable field sites and 8 adjacent seminatural grassland sites and in a 100-m radius around the center of each site. We used Monte Carlo simulations to examine if species abundance and functional traits (height, seed mass, clonal abilities, specific leaf area and dispersal method) controlled the filtering of species from the regional pool into local communities.Important findings On average, only 28% of species found in the regional pool established in the ex-arable field sites and 45% in the seminatural grassland sites, indicating that the size of the regional species pool was not limiting local richness. For both grassland types, species abundance in the regional pool was positively correlated with species occurrence at the local scale. We found evidence for both species interaction filtering and dispersal limitation influencing the local assembly. Both local and regional processes were thus influencing the filtering of species from the regional species pool into local communities. In addition, the age of the communities influenced species filtering, indicating that community assembly and the importance of different filters in that process change over succession.     

海南岛热带低地雨林刀耕火种弃耕地自然恢复过程中的群落构建

研究群落构建机制是群落生态学的一个重要目标, 群落动态过程中的构建规律对于了解群落演替机理有重要的作用。该文以海南岛刀耕火种干扰后自然恢复的10 hm ²热带低地雨林为研究对象, 通过比较不同恢复阶段的次生林(15年、30年和60年)和老龄林在幼苗、幼树和成年树群落的物种组成, 揭示次生演替过程中的群落构建规律。研究结果表明, 老龄林中不同径级群落的物种多样性及不同径级间的物种相似度显著高于各恢复阶段的次生林, 但优势种在群落中的比例低于各恢复阶段的次生林。随着自然恢复过程的进行, 次生林群落物种组成与老龄林的相似性也逐渐增大, 支持演替平衡理论。所有恢复阶段样地中幼苗的个体、物种丰富度和基于多度涵盖估计量(ACE)都低于幼树和成年树群落, 幼苗层物种组成与幼树、成年树也有较大差异, 说明新增到幼苗群落可能是一个难于预测的过程。研究结果说明了确定过程和随机过程共同决定了次生演替的群落构建。