The structure and stability of model ecosystems assembled in a variable environment

To explore how environmental variability may create non‐random community structure, we simulated the assembly of model communities under varying levels of environmental variability. We assembled communities by creating a large pool of randomly constructed species, and then added species from this pool sequentially, allowing extinctions of invading and resident species to occur until the community became saturated. Because much current research on community structure focuses on single trophic levels, we constructed species pools consisting only of competitors. To compare with more realistic communities, we also created species pools with multiple trophic levels. For both types of communities, following assembly we calculated a variety of metrics of community structure, and five measures of community stability. Communities assembled under high environmental variability had fewer species, fewer and weaker interactions among species, and greater evenness in abundance of persisting species. For single trophic‐level communities, community size was dictated primarily by competitive exclusion. In contrast, for multiple trophic‐level communities, community size was increasingly limited by dynamical instabilities as environmental variability increased. Differences in community structure resulting from assembly under high environmental variability led to differences in community stability. According to two measures of stability related to population variability – the characteristic return rate to equilibrium and the coefficient of variation in individual species densities – stability increased for communities assembled under high environmental variability. In contrast, three additional measures of stability that are not directly related to population variability showed a variety of patterns, either increasing, decreasing, or remaining constant. Thus, communities assembled in highly variable environments are not necessarily generically more stable. Our results demonstrate that environmental variability can structure communities and affect their stability properties in non‐trivial ways. Thus, when making predictions about the response of communities to future extinctions or environmental degradation, account should be given to the forces responsible for community structure.     

Population and community variability in randomly fluctuating environments

The prediction that environmental fluctuations may destabilise populations and yet stabilise aggregate community properties has remained largely untested. We examined population and community stability under constant and fluctuating temperatures in simple planktonic assemblages of differing algal richness. Temperature dependent resource competition produced a highly asymmetric community structure where algal community biomass was dominated by one species. For a given level of species richness, temperature fluctuations induced lower community covariance and thus stabilised community biomass. However, increasing algal species richness increased the variability of population abundance and growth rates, as well as population and community variability. Consumer dynamics were directly destabilised by environmental fluctuations. These results confirm recent theoretical studies suggesting a stabilising effect of environmental fluctuations at the community level. However, they also support the theoretical prediction that increasing species richness may be of limited value for community stability, most especially in asymmetric communities, when competition directly affects population variability.     

Direct evidence that density-dependent regulation underpins the temporal stability of abundant species in a diverse animal community

To understand how ecosystems are structured and stabilized, and to identify when communities are at risk of damage or collapse, we need to know how the abundances of the taxa in the entire assemblage vary over ecologically meaningful timescales. Here, we present an analysis of species temporal variability within a single large vertebrate community. Using an exceptionally complete 33-year monthly time series following the dynamics of 81 species of fishes, we show that the most abundant species are least variable in terms of temporal biomass, because they are under density-dependent (negative feedback) regulation. At the other extreme, a relatively large number of low abundance transient species exhibit the greatest population variability. The high stability of the consistently common high abundance species—a result of density-dependence—is reflected in the observation that they consistently represent over 98% of total fish biomass. This leads to steady ecosystem nutrient and energy flux irrespective of the changes in species number and abundance among the large number of low abundance transient species. While the density-dependence of the core species ensures stability under the existing environmental regime, the pool of transient species may support long-term stability by replacing core species should environmental conditions change.     

邙山提灌站浮游植物群落结构空间变化对环境因子的响应

河南邙山提灌站作为黄河流域的重要水源地之一,已列入全国重要饮用水水源地名录。邙山提灌站从黄河取水、沉沙至水厂过程中,沿途生境变化剧烈。为更好地了解这种人工行为造成不同生境中浮游植物群落结构及变化特征,于2014年6月对提灌站的黄河取水口至水厂入口沿途设置的7个采样点开展了浮游植物调查,并同步开展水温、溶氧、总磷等水质因子监测。采用极点排序(POA)和冗余分析(RDA)两种多元统计方法研究了浮游植物群落结构变化及其与环境因子的关系。结果表明:从黄河取水口至水厂取水口,浮游植物群落结构、密度与生物量沿途发生了明显变化,其中浮游植物生物量和生物多样性先升高后降低。POA分析表明7个采样断点被分成3类生境,反映了沿途的生境变化过程,而RDA分析进一步表明总磷(TP)、溶解氧(DO)和氧化还原位(ORP)等水环境因子对浮游植物群结构存在显著的影响,呈现了与湖泊等天然水体不同的变化。     

On one invariant in the centuries-old dynamics of the species structure in freshwater zooplankton

Dynamics of the species structure of Cladocera based on the materials on bottom sediments has been studied for 3000- and 6000-year periods in two lakes. The community of one of these lakes has reached a dynamic climax, and the community of the other lake is at the stage of directing succession. The species structure of both communities is approximated well by the Motomura–Whittaker equation (i.e., model of geometrical series). For each of the studied lakes, the main parameter of this model (the coefficient of geometrical progression) does not differ significantly from its theoretically derived value equaled to 1–exp(–1) = 0.632. A stable relationship is found between species diversity of the community, the rate of ecological succession, and the coefficient of geometrical progression.     

The impact of environmental variability and species composition on the stability of experimental microbial populations and communities

Understanding how environmental fluctuations affect the stability of populations and communities is complex, for example, because direct effects of environmental variability on populations may be modified and propagated across communities by species interactions. One way to explore and further understand these complexities is via a factorial manipulation of community composition and environmental conditions. Using laboratory based aquatic microcosms we manipulated environmental fluctuation by creating two environments; one with variable light and one with constant light. Within these environments, community composition was manipulated by constructing communities from all possible combinations of three species that vary in their reliance on light for growth (an autotroph: a diatom completely reliant on light, a heterotroph: a Paramecium species not reliant on light, and a mixotroph: a Paramecium species somewhat reliant on light). Community composition was predicted to affect populations and communities by introducing and altering competitive interactions between species and affecting the degree of niche differentiation between species. We found that population stability was predominantly influenced by an interaction between community composition and environmental variability, whereby the effect of environmental variability synergistically combined with effects of community composition to reduce population stability. Covariance of populations was determined by an interaction between community composition and environmental variability, though this did not result from the effect of niche differentiation between species. Species interactions drove correlations between population biomass and the environment which otherwise did not exist. Our results demonstrate the complex and interrelated effects of abiotic and biotic factors on population and community stability, and suggest the need to consider aspects of community composition when predicting the impact of environmental fluctuations.     

长江口潮下带春季大型底栖动物的群落结构

2005年4月对长江口全区域潮下带共10个采样站位的大型底栖动物进行了调查。调查采获大型底栖动物38种,分属5个生态类型,种类数较少,河口外缘站位种类数多于口内站位。各站位大型底栖动物的平均丰度为32.9个/m2、平均生物量为5.035g/m2(湿重);与20世纪七八十年代相比,平均生物量显著降低;口外缘站位的总丰度和总生物量均高于口内站位。环境因子相关分析表明,盐度是决定长江口大型底栖动物种类分布最重要的环境因子。群落聚类、标序分析显示,春季长江口潮下带大型底栖动物群落结构空间分异明显,完全符合目前长江口支、港、槽“三级分汊”的空间格局。其中,北支的大型底栖动物以混合高盐水种类为主,而南支则以淡水和半咸水种类为主。南支的南北槽分界处内外站位的群落差异也由盐度决定,因为靠近口内的群落均受长江冲淡水影响较大;而口外站位群落则受咸淡水影响。南支的南北港分界点内外的群落差异则主要受长江来水的影响,原因在于处在港分界点以内的群落所在区域,直接受长江来水的冲刷,底质环境极不稳定;而港、槽分界点之间的群落所在区域由于河口上段的诸多明暗沙体的阻挡,水势较为稳定,所以底质环境较稳定,从而使得港、槽分界点之间的群落出现了更多的沙蚕等底质环境类型种类。     

Extrinsic environmental factors,not resident diversity itself,lead to invasion of <Emphasis Type="Italic">Ageratum conyzoides</Emphasis> L. in diverse communities

The relationship between diversity and invasibility might be confounded by extrinsic environmental factors and the evolutionary structure of the resident community. To examine the role of extrinsic environmental factors, species and phylogenetic diversity in regulating community susceptibility to invasion, we established 109 plots either with or without Ageratum conyzoides L. in Liandu, China. We identified all the species in our samples, weighed the aboveground biomass of each species, and measured environmental variables. For all species recorded in our survey, we constructed a community phylogeny using PhytoPhylo mega-phylogeny as a backbone. We selected the best-fit environment model based on the minimum corrected Akaike information criteria score to examine the effect of extrinsic environmental variables on the relative abundance of A. conyzoides. Relationship between biodiversity and invasion of A. conyzoides was examined by a multiple regression, in which extrinsic ecological factors and biodiversity were combined to predict the relative abundance of A. conyzoides. To reduce the number of extrinsic variables, the first six components produced by a principal component analysis of environmental variables were used as predictive variables in the multiple regression. The best-fit environment model indicated that the relative abundance of A. conyzoides was higher in summer and in communities with lower total organic matter and higher total nitrogen in the soil. The multiple regression indicated that only the positive relationship between the Shannon–Wiener diversity of exotics and the relative abundance of A. conyzoides was significant. This result challenges the importance of diversity–resistance to plant invasion. Generalist facilitation might exist between A. conyzoides and other exotic species, although mechanisms for such facilitation are unclear. Overall, our finding suggests the extrinsic factors covarying with diversity are more important than diversity itself in regulating community susceptibility to invasion.     

Distribution of fish and the biotopic structure of the basin of the small salmon river (the Nachilova River,Western Kamchatka)

For one of the small Western Kamchatka rivers, according to the combination of several characteristics (watershed, channel slope, water abundance of the flow, pattern of bottom sediments), the classification of channel processes was performed, and on its basis, six biotopes successively replacing each other from the effluents to the mouth were separated. It was shown that against the background of relatively stable conditions of the summer-autumn period, an independent population of Salmonoidei, an assembly, is formed within each biotope. This is manifested in specific features of species composition, size and age structure, and the density of populating biotopes. The stability of the distribution of fish population is supported by constant species composition in particular biotopes over three years and by a comparatively high level of residence of the dominant salmon species.     

Testate amoebae communities in the southern tundra and forest-tundra of Western Siberia

The species composition and community structure of soil-inhabiting testate amoebae communities have been studied in biotopes of different types in the southern tundra and forest-tundra of the Tazovskaya Lowland, Western Siberia. A total of 93 species and forms have been identified. It has been found that the species richness of testate amoebae is much lower in dry than in moist biotopes due to a lower level of beta-diversity, with alpha diversity being the same (on average, 16.9 and 17.1 species per sample, respectively). Factors acting at the microbiotope level (biotope type and moisture) play the most important role in the formation of species richness; biotope features (soils and vegetation) are second in importance. In moist habitats, local communities of testate amoebae from different microbiotopes (mosses, lichens, or litter) are fairly similar in species structure, and communities from different moist biotopes are heterogeneous. In dry areas, the opposite situation is observed: local communities differ at the microbiotope level but are similar at the biotope level. The abundance of testate amoebae in moist biotopes reaches 200 × 10³ ind./g dry soil, being an order of magnitude lower in dry biotopes.     

一座南亚热带小型贫营养水库浮游植物群落结构及季节变化

库容大小是影响水体水动力学过程的一个重要变量,它能在很大程度决定生态系统的结构,特别是浮游植物的群落结构.为了解小型贫营养水库浮游植物的群落特点,于2006年4、8、12月对位珠海市的贫营养小型水库-吉大水库的浮游植物群落结构进行采样和计数分析.3次采样共检出浮游植物32种,浮游植物细胞丰度的变化范围在69～342 cells·mL^-1,生物量的变化范围在1.34～3.69 mg·L^-1,夏季浮游植物的丰度和生物量明显高于冬季.甲藻是最主要的优势种类,且相对优势度较为稳定.夏季,隐藻门的隐藻(Cryptomonas sp.)和绿藻门的鼓藻(Cosmarium spp.)大量出现,甲藻的相对优势度有所降低.冬季,隐藻数量急剧下降,但硅藻门的颗粒直链藻(Aulacoseira granulata)大量出现,与甲藻共同成为水体中的优势种.由降雨引起的营养盐浓度增加是浮游植物变化的主要影响因子,而透明度全年维持较高的水平为浮游植物的生长形成了有利条件,此外,较为稳定的水体和甲藻利用营养盐的能力使得甲藻成为浮游植物中的最主要的优势种.     

Distance decay of similarity,effects of environmental noise and ecological heterogeneity among species in the spatio‐temporal dynamics of a dispersal‐limited community

The joint spatial and temporal fluctuations in community structure may be due to dispersal, variation in environmental conditions, ecological heterogeneity among species and demographic stochasticity. These factors are not mutually exclusive, and their relative contribution towards shaping species abundance distributions and in causing species fluctuations have been hard to disentangle. To better understand community dynamics when the exchange of individuals between localities is very low, we studied the dynamics of the freshwater zooplankton communities in 17 lakes located in independent catchment areas, sampled at end of summer from 2002 to 2008 in Norway. We analysed the joint spatial and temporal fluctuations in the community structure by fitting the two‐dimensional Poisson lognormal model under a two‐stage sampling scheme. We partitioned the variance of the distribution of log abundance for a random species at a random time and location into components of demographic stochasticity, ecological heterogeneity among species, and independent environmental noise components for the different species. Non‐neutral mechanisms such as ecological heterogeneity among species (20%) and spatiotemporal variation in the environment (75%) explained the majority of the variance in log abundances. Overdispersion relative to Poisson sampling and demographic stochasticity had a small contribution to the variance (5%). Among a set of environmental variables, lake acidity was the environmental variable that was most strongly related to decay of community similarity in space and time.     

Distribution of heterotrophic flagellates at the littoral of estuary of Chernaya River (Kandalaksha Bay, White Sea)

Species composition, distribution, and the character of structural changes in the heterotrophic flagellate community were studied along environmental gradients in the Chernaya River estuary. There were 99 species and forms of heterotrophic flagellates, subdivided into three groups: prevalently marine species and euryhaline species preferring biotopes either of higher or decreased salinity. The heterotrophic flagellate community of the estuary was continuously divided into two distinct variants: (1) cenosis of halophilic species, prevalently of sea forms and euryhaline species preferring biotopes of increased salinity; (2) cenosis of halophobic species with prevalence of euryhaline forms gravitating to fresh biotopes. The arbitrary and indistinct boundary between the variants of the community ran at a salinity of 9–10‰. The response of estuarine communities of heterotrophic flagellates and infusorians to variation of abiotic factors was similar and differed from response of communities of microphyto-, meiozoo-and macrozoobenthos; this implied similarity of the response mechanism to environmental factors in organisms of one level of organization.     

Habitat size determine algae biomass in tank-bromeliads

The primary goal of this study was to evaluate the relative importance of habitat physical properties, bottom-up and top-down factors, and their interaction on algae biomass in tank-bromeliads. We sampled algae biomass (chlorophyll-a concentration), micro-metazoan density, mosquito abundance, and several environmental variables, including nutrient concentrations and characteristics of the habitat physical structure, in a survey of 64 tank-bromeliads of four different species (Aechmea nudicaulis, Aechmea lingulata, Neoregelia cruenta, and Vriesea neoglutinosa). We analyzed the complete and individual bromeliad species datasets using an information-theoretic model selection approach (Akaike’s information criterion). Bromeliad species, maximum water volume, and bromeliad diameter comprised the best model for determining chlorophyll-a concentrations for the complete dataset. The maximum water volume also comprised the best model to explain chlorophyll-a concentrations in three of four bromeliad species datasets. Interactions between consumers and nutrient concentration were included in the subsequent models, but they were not statistically significant. Taken together, these results demonstrate that the impact of habitat size on the associated autotrophic biomass occurs possibly via changes in community susceptibility to disturbances, particularly drought. We can conclude that habitat size is more important than resource availability or herbivory on phytotelm autotrophic biomass regulation in these natural microcosms.     

Scale-Dependence of Processes Structuring Dung Beetle Metacommunities Using Functional Diversity and Community Deconstruction Approaches

Community structure is driven by mechanisms linked to environmental, spatial and temporal processes, which have been successfully addressed using metacommunity framework. The relative importance of processes shaping community structure can be identified using several different approaches. Two approaches that are increasingly being used are functional diversity and community deconstruction. Functional diversity is measured using various indices that incorporate distinct community attributes. Community deconstruction is a way to disentangle species responses to ecological processes by grouping species with similar traits. We used these two approaches to determine whether they are improvements over traditional measures (e.g., species composition, abundance, biomass) for identification of the main processes driving dung beetle (Scarabaeinae) community structure in a fragmented mainland-island landscape in southern Brazilian Atlantic Forest. We sampled five sites in each of four large forest areas, two on the mainland and two on the island. Sampling was performed in 2012 and 2013. We collected abundance and biomass data from 100 sampling points distributed over 20 sampling sites. We studied environmental, spatial and temporal effects on dung beetle community across three spatial scales, i.e., between sites, between areas and mainland-island. The γ-diversity based on species abundance was mainly attributed to β-diversity as a consequence of the increase in mean α- and β-diversity between areas. Variation partitioning on abundance, biomass and functional diversity showed scale-dependence of processes structuring dung beetle metacommunities. We identified two major groups of responses among 17 functional groups. In general, environmental filters were important at both local and regional scales. Spatial factors were important at the intermediate scale. Our study supports the notion of scale-dependence of environmental, spatial and temporal processes in the distribution and functional organization of Scarabaeinae beetles. We conclude that functional diversity may be used as a complementary approach to traditional measures, and that community deconstruction allows sufficient disentangling of responses of different trait-based groups.     

Diversity and composition of macrobenthic community associated with sandy shoals of the Louisiana continental shelf

Along the Louisiana, USA continental shelf, sandy shoals are shallow, possibly oxygen-rich “islands” surrounded by deeper muddy deposits prone to hypoxia. Shoals also contain significant quantities of fine sand that may be mined in the future for coastal restoration. The ecological role of shoals remains poorly understood and we hypothesized that shoals provide critical habitat for benthic invertebrates. Using Ship Shoal as a model system, we assessed the diversity and structure of macrobenthic assemblages and how community structure varies with season and environmental parameters. High biomass (averaging 26.7 g m⁻²) and high diversity (161 species) of macrobenthos was found in 2006. Polychaetes (45%—72 species) and crustaceans comprised most of the species (28%—46 species); spionids and amphipods dominated the polychaete and crustacean groups respectively, both in terms of number of species and abundances. Sharp decreases in diversity, abundance and biomass occurred from spring to autumn. Species diversity and total abundance significantly increased with decreasing sediment grain size and increasing bottom water dissolved oxygen. Across seasons, mole crabs Albunea paretii and amphioxus Branchiostoma floridae typified the community and contributed most of the biomass. The polychaetes Nephtys simoni, Neanthes micromma, Dispio uncinata, Mediomastus californiensis and Magelona sp. A, the amphipod Acanthohautorius sp. A and the burrowing shrimp Ogyrides alphaerostris also contributed to variation in community composition. Cluster analyses quantified seasonal variation, mainly based on sharp decreases in abundance, as well as spatial differences in species composition oriented along both east–west and north–south gradients. Variation in benthic assemblages was correlated with water depth and sediment characteristics (mean grain size and percentage of gravel-sized shell debris). We conclude that Ship Shoal is an unrecognized biodiversity hotspot and a hypoxia refuge compared to the immediate surrounding area where the benthic community is affected by seasonal hypoxia events and we discuss how sand-mining may influence this community.     

Trade-offs in community properties through time in a desert rodent community

Resource limitation represents an important constraint on ecological communities, which restricts the total abundance, biomass, and community energy flux a given community can support. However, the exact relationship among these three measures of biological activity remains unclear. Here we use a simple framework that links abundance and biomass with an energetic constraint. Under constant energetic availability, it is expected that changes in abundance and biomass can result from shifts in the distribution of individual masses. We test these predictions using long-term data from a desert rodent community. Total energy use for the community has not changed directionally for 25 years, but species composition has. As a result, the average body size has decreased by almost 50%, and average abundance has doubled. These results lend support to the idea of resource limitation on desert rodent communities and demonstrate that systems are able to maintain community energy flux in the face of environmental change, through changes in composition and structure.     

Sensitivity of three grassland communities to simulated extreme temperature and rainfall events

Three grassland communities in New Zealand with differing climates and proportions of C3 and C4 species were subjected to one‐off extreme heating (eight hours at 52.5°C) and rainfall (the equivalent of 100 mm) events. A novel experimental technique using portable computer‐controlled chambers simulated the extreme heating events. The productive, moist C3/C4 community was the most sensitive to the extreme events in terms of short‐term community composition compared with a dry C3/C4 community or an exclusively C3 community. An extreme heating event caused the greatest change to plant community species abundance by favouring the expansion of C4 species relative to C3 species, shifting C4 species abundance from 43% up to 84% at the productive, moist site. This was observed both in the presence and absence of added water. In the absence of C4 species, heating reduced community productivity by over 60%. The short‐term shifts in the abundance of C3 and C4 species in response to the single extreme climatic events did not have persistent effects on community structure or on soil nitrogen one year later. There was no consistent relationship between diversity and stability of biomass production of these plant communities, and species functional identity was the most effective explanation for the observed shifts in biomass production. The presence of C4 species resulted in an increased stability of productivity after extreme climatic events, but resulted in greater overall shifts in community composition. The presence of C4 species may buffer grassland community productivity against an increased frequency of extreme heating events associated with future global climate change.