Phylogenetic beta diversity in bacterial assemblages across ecosystems: deterministic versus stochastic processes

Increasing evidence has emerged for non-random spatial distributions of microbes, but knowledge of the processes that cause variation in microbial assemblage among ecosystems is lacking. For instance, some studies showed that deterministic processes such as habitat specialization are important, while other studies hold that bacterial communities are assembled by stochastic forces. Here we examine the relative influence of deterministic and stochastic processes for bacterial communities from subsurface environments, stream biofilm, lake water, lake sediment and soil using pyrosequencing of the 16S ribosomal RNA gene. We show that there is a general pattern in phylogenetic signal in species ecological niches across recent evolutionary time for all studied habitats, enabling us to infer the influences of community assembly processes from patterns of phylogenetic turnover in community composition. The phylogenetic dissimilarities among-habitat types were significantly higher than within them, and the communities were clustered according to their original habitat types. For communities within-habitat types, the highest phylogenetic turnover rate through space was observed in subsurface environments, followed by stream biofilm on mountainsides, whereas the sediment assemblages across regional scales showed the lowest turnover rate. Quantifying phylogenetic turnover as the deviation from a null expectation suggested that measured environmental variables imposed strong selection on bacterial communities for nearly all sample groups. For three sample groups, spatial distance reflected unmeasured environmental variables that impose selection, as opposed to spatial isolation. Such characterization of spatial and environmental variables proved essential for proper interpretation of partial Mantel results based on observed beta diversity metrics. In summary, our results clearly indicate a dominant role of deterministic processes on bacterial assemblages and highlight that bacteria show strong habitat associations that have likely emerged through evolutionary adaptation.     

Dissecting spatiotemporal patterns of functional diversity through the lens of Darwin's naturalization conundrum

Darwin's naturalization conundrum describes the paradigm that community assembly is regulated by two opposing processes, environmental filtering and competitive interactions, which predict both similarity and distinctiveness of species to be important for establishment. Our goal is to use long‐term, large‐scale, and high‐resolution temporal data to examine diversity patterns over time and assess whether environmental filtering or competition plays a larger role in regulating community assembly processes. We evaluated Darwin's naturalization conundrum and how functional diversity has changed in the Laurentian Great Lakes fish community from 1870 to 2010, which has experienced frequent introductions of non‐native species and extirpations of native species. We analyzed how functional diversity has changed over time by decade from 1870 to 2010 at three spatial scales (regional, lake, and habitat) to account for potential noninteractions between species at the regional and lake level. We also determined which process, environmental filtering or competitive interactions, is more important in regulating community assembly and maintenance by comparing observed patterns to what we would expect in the absence of an ecological mechanism. With the exception of one community, all analyses show that functional diversity and species richness has increased over time and that environmental filtering regulates community assembly at the regional level. When examining functional diversity at the lake and habitat level, the regulating processes become more context dependent. This study is the first to examine diversity patterns and Darwin's conundrum by integrating long‐term, large‐scale, and high‐resolution temporal data at multiple spatial scales. Our results confirm that Darwin's conundrum is highly context dependent.     

Phylogenetic distance–decay patterns are not explained by local community assembly processes in freshwater lake microbial communities

While water and sediment microbial communities exhibit pronounced spatio-temporal patterns in freshwater lakes, the underlying drivers are yet poorly understood. Here, we evaluated the importance of spatial and temporal variation in abiotic environmental factors for bacterial and microeukaryotic community assembly and distance–decay relationships in water and sediment niches in Hongze Lake. By sampling across the whole lake during both Autumn and Spring sampling time points, we show that only bacterial sediment communities were governed by deterministic community assembly processes due to abiotic environmental drivers. Nevertheless, consistent distance–decay relationships were found with both bacterial and microeukaryotic communities, which were relatively stable with both sampling time points. Our results suggest that spatio-temporal variation in environmental factors was important in explaining mainly bacterial community assembly in the sediment, possibly due lesser disturbance. However, clear distance–decay patterns emerged also when the community assembly was stochastic. Together, these results suggest that abiotic environmental factors do not clearly drive the spatial structuring of lake microbial communities, highlighting the need to understand the role of other potential drivers, such as spatial heterogeneity and biotic species interactions.     

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.     

Nearly a decade‐long repeatable seasonal diversity patterns of bacterioplankton communities in the eutrophic Lake Donghu (Wuhan,China)

Uncovering which environmental factors govern community diversity patterns and how ecological processes drive community turnover are key questions related to understand the community assembly. However, the ecological mechanisms regulating long‐term variations of bacterioplankton communities in lake ecosystems remain poorly understood. Here we present nearly a decade‐long study of bacterioplankton communities from the eutrophic Lake Donghu (Wuhan, China) using 16S rRNA gene amplicon sequencing with MiSeq platform. We found strong repeatable seasonal diversity patterns in terms of both common (detected in more than 50% samples) and dominant (relative abundance >1%) bacterial taxa turnover. Moreover, community composition tracked the seasonal temperature gradient, indicating that temperature is a key environmental factor controlling observed diversity patterns. Total phosphorus also contributed significantly to the seasonal shifts in bacterioplankton composition. However, any spatial pattern of bacterioplankton communities across the main lake areas within season was overwhelmed by their temporal variabilities. Phylogenetic analysis further indicated that 75%–82% of community turnover was governed by homogeneous selection due to consistent environmental conditions within seasons, suggesting that the microbial communities in Lake Donghu are mainly controlled by niche‐based processes. Therefore, dominant niches available within seasons might be occupied by similar combinations of bacterial taxa with modest dispersal rates throughout different lake areas.     

Phylogenetic turnover during subtropical forest succession across environmental and phylogenetic scales

Although spatial and temporal patterns of phylogenetic community structure during succession are inherently interlinked and assembly processes vary with environmental and phylogenetic scales, successional studies of community assembly have yet to integrate spatial and temporal components of community structure, while accounting for scaling issues. To gain insight into the processes that generate biodiversity after disturbance, we combine analyses of spatial and temporal phylogenetic turnover across phylogenetic scales, accounting for covariation with environmental differences. We compared phylogenetic turnover, at the species‐ and individual‐level, within and between five successional stages, representing woody plant communities in a subtropical forest chronosequence. We decomposed turnover at different phylogenetic depths and assessed its covariation with between‐plot abiotic differences. Phylogenetic turnover between stages was low relative to species turnover and was not explained by abiotic differences. However, within the late‐successional stages, there was high presence‐/absence‐based turnover (clustering) that occurred deep in the phylogeny and covaried with environmental differentiation. Our results support a deterministic model of community assembly where (i) phylogenetic composition is constrained through successional time, but (ii) toward late succession, species sorting into preferred habitats according to niche traits that are conserved deep in phylogeny, becomes increasingly important.     

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.     

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.     

Patterns of temporal community turnover are spatially synchronous across boreal lakes

1. Ecosystems are often exposed to broad‐scale environmental change, which can potentially synchronise community dynamics and biodiversity trends. Detection of temporal coherence may, however, depend on the metrics used and their sensitivity to detect change, requiring several lines of evidence to elucidate the full range of temporal responses to environmental change. 2. Here, we tested whether the patterns of synchrony among littoral invertebrate communities of Swedish lakes over 20 years (1988–2007) differed when analysed using univariate (taxon richness, evenness, Shannon diversity and total abundance) or multivariate (temporal turnover in community composition) metrics. We included both culturally acidified and circumneutral lakes to examine whether anthropogenic stress influenced the patterns of synchrony. 3. Average total abundance, taxon richness and temporal turnover in community composition changed monotonically with time, while evenness and Shannon diversity fluctuated around a long‐term mean. However, among‐lake variability was high, resulting in a weak temporal coherence. Only trends of temporal turnover changed synchronously across lakes, irrespective of their acidification history. 4. Spatially synchronous trends in turnover across lakes were correlated with increasing water colour and decreasing sulphate concentrations, showing the importance of regional drivers of spatiotemporal coherence. 5. Our results underpin an increasing body of evidence that the detection of diversity patterns varies among metrics that ignore (taxon richness, evenness, Shannon diversity) or consider (turnover) species identities. More generally, our results suggest that community‐level studies of synchrony are suitable for elucidating the role of intrinsic versus extrinsic factors in mediating complex community assembly processes in the long term. This, in turn, contributes to our understanding of temporal patterns of biodiversity.     

Distinct successional patterns and processes of free-living and particle-attached bacterial communities throughout a phytoplankton bloom

1. Understanding the successional patterns of microbial communities during a phytoplankton bloom is crucial for predicting the compositional and functional stability of lake ecosystems in response to the disturbance of a bloom. Previous studies on bacterial communities associated with blooms have rarely studied the dynamics of these communities. The successional patterns of bacterial communities within different micro-habitats (i.e. water column versus particles) and mechanisms that shape these communities that differ in composition and structure remain unclear.
2. We selected a eutrophic urban lake to investigate the succession of bacterial communities during a bloom. We divided the bacterial communities into free-living (FL) and particle-attached (PA) groups based on their different lifestyles. The amplicon-based 16S rRNA gene high-throughput sequencing technology was used to obtain bacterial community composition and phylogenetic structure.
3. Our study showed distinct successional patterns between FL and PA bacterial communities, and the two bacterial lifestyles showed different responses and resilience to the bloom, in terms of diversity and relative abundance of bacterial taxa. Alpha-diversity of the PA bacterial community decreased during the bloom, whereas that of the FL bacterial community increased. More taxa in the FL bacterial community showed resilience after the disturbance than in the PA bacterial community.
4. The influence of phytoplankton blooms on the assembly of the bacterial community can be viewed as niche selection that led to the decrease in the relative importance of stochastic processes in shaping both FL and PA bacterial communities. This study shows the ecological significance of the bacterial community response to bloom events in lakes. It also shows that assembly processes differ for bacterial communities that have different lifestyles in lake ecosystems disturbed by phytoplankton blooms.

     

荒漠草原盐沼湿地苦豆子土壤细菌群落构建机制及其影响因素

【背景】荒漠草原盐沼湿地是陆地生态系统的重要组成部分,土壤水分和盐分变化是影响该生态系统土壤细菌群落构建的重要因素。【目的】土壤细菌群落构建是由确定性和随机性主导的连续生态过程,阐明荒漠草原盐沼湿地土壤细菌群落的构建机制对于加深微生物作为关键生态系统因子重要性的理解具有积极意义。【方法】以宁夏中部典型荒漠草原盐沼苦水湖湿地为研究对象,对近湖边（near the lake,NL）和远离湖边（far from the lake,FL）苦豆子群落土壤理化特性进行测定并结合土壤细菌高通量测序分析。【结果】NL和FL样地具有明显的水盐梯度变化,NL样地土壤pH、含水量和电导率均显著高于FL样地;变形菌门、放线菌门、厚壁菌门、拟杆菌门和黏菌门是研究区域土壤细菌群落的优势菌门,变形菌门相对丰度随水盐梯度上升而升高,放线菌门和厚壁菌门相对丰度则随之下降,门下成员大多与水盐变化具有明显的相关性;此外,FL样地土壤细菌网络则具有稳定的网络关系;随着NL样地向FL样地的延伸,土壤细菌群落由随机过程主导,并且受pH、电导率和环境变量的影响。【结论】荒漠草原盐沼湿地水分和盐分的变化改变了土壤细菌群落结构;土壤细菌群落通过生态位占据等策略提高逆境下的生存能力;细菌群落构建是随机过程和确定性过程组成的连续统一体,同样受环境变化的影响。本结果揭示了荒漠草原盐沼湿地细菌群落结构和相互关系对环境变化的响应特征,同时阐明了该区土壤细菌群落的构建机制及影响因素,也为相关科学研究提供了一定理论参考。     

Depth-Dependent Patterns of Bacterial Communities and Assembly Processes in a Typical Red Soil Critical Zone

Abstract

Depth patterns of soil microbial distribution have not been well characterized and little is known about the balance between stochastic and deterministic processes in shaping microbial community through soil profiles. We examined how the composition, diversity, and assembly processes of bacterial communities change to a depth of 4 m along three cores sampled through a typical upland red soil Critical Zone in subtropical China. The sampled soils at the center of the Critical Zone were acidic, highly weathered, and clay loam to clay in texture, and developed from Quaternary red clay. Bacterial richness and phylogenetic diversity decreased with depth in the upper soil zone (0–90?cm), but was constant in the deeper zone (90–420?cm). In both the upper and deeper zones, richness was strongly and positively correlated with total organic carbon, dissolved organic carbon and total nitrogen. Depth-dependent changes of community structure were observed above the upper but not deeper zones. Stochastic processes were more important in the upper zone, whereas deterministic processes were dominant in the deeper zone. The distinct depth-dependent patterns of bacterial communities and assembly processes exist through deep soil profiles and are influenced by both contemporary and historical pedogenetic factors and processes.     

Microbial biogeography of permafrost thaw ponds across the changing northern landscape

Microbial diversity varies at multiple spatial scales, but little is known about how climate change may influence this variation. Here we assessed the free‐living bacterioplankton composition of thaw ponds over a north‐south gradient of permafrost degradation in the eastern Canadian subarctic. Three nested spatial scales were compared: 1) among ponds within individual valleys 2) between two valleys within each landscape type, and 3) between landscape types (southern sporadic versus northern discontinuous permafrost). As a reference point, we sampled rock‐basin lakes whose formation was not related to permafrost thawing. β‐diversity was low at the smallest scale despite marked differences in limnological properties among neighboring ponds. β‐diversity was high among valleys, associated with greater environmental heterogeneity. The largest differences were between landscape types and appeared to reflect the concomitant effects of environmental filtering and dispersal limitation. Raup–Crick β‐diversity indicated that community assembly was driven by both stochastic (random extinction, dispersal, ecological drift) and deterministic (environmental filtering) processes. Communities sampled in the most degraded valley appeared primarily assembled through stochastic processes, while environmental filtering played a greater role at the other valleys. These results imply that climate warming and ongoing permafrost degradation will influence microbial community assembly, which in turn is likely to affect the functioning of thaw pond ecosystems.     

Intraspecific trait variation improves the detection of deterministic community assembly processes in early successional forests,but not in late successional forests

Aims Intraspecific trait variation (ITV) has been increasingly recognized to play an important role in understanding the underlying processes influencing community assembly. However, gaps remain in our understanding of how incorporating ITV will influence the relative importance of deterministic (e.g. habitat filtering, limiting similarity) and stochastic processes in driving community assembly at different successional stages.     

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.     

保安湖水体细菌群落结构时空变化特征及驱动因子

为了探究保安湖水体细菌群落结构时空变化特征及驱动因子,研究于2019年春、夏、秋、冬四季采集保安湖水样,使用宏基因组测序技术对保安湖水体细菌群落的组成及多样性变化进行了研究。结果表明:(1)保安湖不同湖区细菌群落组成无显著差异(P>0.05);夏、秋季细菌丰富度、均匀度、香农及辛普森多样性指数均显著高于春、冬季(P<0.05),夏、秋季优势类群为变形菌门(Proteobacteria)、放线菌门(Actinobactetiota)、蓝藻门(Cyanobacteria),而春、冬季变形菌门(Proteobacteria)和放线菌门(Actinobactetiota)占主导地位;(2)温度、透明度、pH、溶解氧、高锰酸盐指数、叶绿素a及总磷等因素是保安湖水体细菌群落结构变化的重要驱动因子;(3)保安湖细菌群落构建过程在春、夏、秋季由随机性过程主导,在冬季由确定性过程主导;(4)保安湖细菌网络互作具有明显的季节特征,从春季到冬季,保安湖细菌种间相互作用逐渐紧密复杂。综上所述,保安湖水体细菌群落结构具有明显的季节变化特征,温度、透明度、pH、溶解氧、高锰酸盐指数等因素具有驱动水体细...     

Phylogenetic and functional dissimilarity does not increase during temporal heathland succession

Succession has been a focal point of ecological research for over a century, but thus far has been poorly explored through the lens of modern phylogenetic and trait-based approaches to community assembly. The vast majority of studies conducted to date have comprised static analyses where communities are observed at a single snapshot in time. Long-term datasets present a vantage point to compare established and emerging theoretical predictions on the phylogenetic and functional trajectory of communities through succession. We investigated within, and between, community measures of phylogenetic and functional diversity in a fire-prone heathland along a 21 year time series. Contrary to widely held expectations that increased competition through succession should inhibit the coexistence of species with high niche overlap, plots became more phylogenetically and functionally clustered with time since fire. There were significant directional shifts in individual traits through time indicating deterministic successional processes associated with changing abiotic and/or biotic conditions. However, relative to the observed temporal rate of taxonomic turnover, both phylogenetic and functional turnover were comparatively low, suggesting a degree of functional redundancy among close relatives. These results contribute to an emerging body of evidence indicating that limits to the similarity of coexisting species are rarely observed at fine spatial scales.     

The macroecology of rapid evolutionary radiation

A long-standing debate in ecology addresses whether community composition is the result of stochastic factors or assembly rules. Non-random, over-dispersed patterns of species co-occurrence have commonly been attributed to competition--a particularly important force in adaptive radiation. We thus examined the macroecology of the recently radiated cichlid rock-fish assemblage in Lake Malawi, Africa at a spectrum of increasingly fine spatial scales (entire lake to depth within rock-reef sites). Along this range of spatial scales, we observed a signal of community structure (decreased co-occurrence of species) at the largest and smallest scales, but not in between. Evidence suggests that the lakewide signature of structure is driven by extreme endemism and micro-allopatric speciation, while patterns of reduced co-occurrence with depth are indicative of species interactions. We identified a 'core' set of rock-reef species, found in combination throughout the lake, whose depth profiles exhibited replicated positive and negative correlation. Our results provide insight into how ecological communities may be structured differently at distinct spatial scales, re-emphasize the importance of local species interactions in community assembly, and further elucidate the processes shaping speciation in this model adaptive radiation.     

pH Influences the Importance of Niche-Related and Neutral Processes in Lacustrine Bacterioplankton Assembly

pH is an important factor that shapes the structure of bacterial communities. However, we have very limited information about the patterns and processes by which overall bacterioplankton communities assemble across wide pH gradients in natural freshwater lakes. Here, we used pyrosequencing to analyze the bacterioplankton communities in 25 discrete freshwater lakes in Denmark with pH levels ranging from 3.8 to 8.8. We found that pH was the key factor impacting lacustrine bacterioplankton community assembly. More acidic lakes imposed stronger environmental filtering, which decreased the richness and evenness of bacterioplankton operational taxonomic units (OTUs) and largely shifted community composition. Although environmental filtering was determined to be the most important determinant of bacterioplankton community assembly, the importance of neutral assembly processes must also be considered, notably in acidic lakes, where the species (OTU) diversity was low. We observed that the strong effect of environmental filtering in more acidic lakes was weakened by the enhanced relative importance of neutral community assembly, and bacterioplankton communities tended to be less phylogenetically clustered in more acidic lakes. In summary, we propose that pH is a major environmental determinant in freshwater lakes, regulating the relative importance and interplay between niche-related and neutral processes and shaping the patterns of freshwater lake bacterioplankton biodiversity.     

Soil pH and temperature regulate assembly processes of abundant and rare bacterial communities in agricultural ecosystems

The factors determining stochastic and deterministic processes that drive microbial community structure, specifically the balance of abundant and rare bacterial taxa, remain underexplored. Here we examined biogeographic patterns of abundant and rare bacterial taxa and explored environmental factors influencing their community assembly processes in agricultural fields across eastern China. More phylogenetic turnover correlating with spatial distance was observed in abundant than rare sub-communities. Homogeneous selection was the main assembly process for both the abundant and rare sub-communities; however, the abundant sub-community was more tightly clustered phylogenetically and was more sensitive to dispersal limitations than the rare sub-community. Rare sub-community of rice fields and abundant sub-community of maize fields were more governed by stochastic assembly processes, which showed higher operational taxonomic unit richness. We propose a conceptual paradigm wherein soil pH and mean annual temperature mediate the assembly of the abundant and rare sub-communities respectively. A higher soil pH leads to deterministic assembly of the abundant sub-community. For the rare sub-community, the dominance of stochasticity in low-temperature regions indicates weaker niche-based exclusion and the arrival of more evolutionary lineages. These findings suggest that the community assembly processes for abundant and rare bacterial taxa are dependent on distinct environmental variables in agro-ecosystems.