Processes of community assembly in an environmentally heterogeneous,high biodiversity region

Despite decades of study, the relative importance of niche‐based versus neutral processes in community assembly remains largely ambiguous. Recent work suggests niche‐based processes are more easily detectable at coarser spatial scales, while neutrality dominates at finer scales. Analyses of functional traits with multi‐year multi‐site biodiversity inventories may provide deeper insights into assembly processes and the effects of spatial scale. We examined associations between community composition, species functional traits, and environmental conditions for plant communities in the Kouga‐Baviaanskloof region, an area within South Africa's Cape Floristic Region (CFR) containing high α and β diversity. This region contains strong climatic gradients and topographic heterogeneity, and is comprised of distinct vegetation classes with varying fire histories, making it an ideal location to assess the role of niche‐based environmental filtering on community composition by examining how traits vary with environment. We combined functional trait measurements for over 300 species with observations from vegetation surveys carried out in 1991/1992 and repeated in 2011/2012. We applied redundancy analysis, quantile regression, and null model tests to examine trends in species turnover and functional traits along environmental gradients in space and through time. Functional trait values were weakly associated with most spatial environmental gradients and only showed trends with respect to vegetation class and time since fire. However, survey plots showed greater compositional and functional stability through time than expected based on null models. Taken together, we found clear evidence for functional distinctions between vegetation classes, suggesting strong environmental filtering at this scale, most likely driven by fire dynamics. In contrast, there was little evidence of filtering effects along environmental gradients within vegetation classes, suggesting that assembly processes are largely neutral at this scale, likely the result of very high functional redundancy among species in the regional species pool.     

Functional potential and assembly of microbes from sediments in a lake bay and adjoining river ecosystem for polycyclic aromatic hydrocarbon biodegradation

Lake and adjoining river ecosystems are ecologically and economically valuable and are heavily threatened by anthropogenic activities. Determining the inherent capacity of ecosystems for polycyclic aromatic hydrocarbon (PAH) biodegradation can help quantify environmental impacts on the functioning of ecosystems, especially on that of the microbial community. Here, PAH biodegradation potential was compared between sediments collected from a lake bay (LS) and an adjoining river (RS) ecosystem. Microbial community composition, function, and their co-occurrence patterns were also explored. In the RS, the biodegradation rates (K_D) of pyrene or PAH were almost two orders of magnitude higher than those in the LS. Sediment functional community structure and network interactions were dramatically different between the LS and RS. Although PAH degradation genes (p450aro, quinoline, and qorl) were detected in the LS, the community activity of these genes needed to be biostimulated for accelerated bioremediation. In contrast, functional communities in the RS were capable of spontaneous natural attenuation of PAH. The degradation of PAH in the RS also required coordinated response of the complex functional community. Taken together, elucidating functions and network interactions in sediment microbial communities and their responses to environmental changes are very important for the bioremediation of anthropogenic toxic contaminants.     

Microbial Endpoints: The Rationale for their Exclusion as Ecological Assessment Endpoints

The functional importance of bacteria and fungi in terrestrial systems is recognized widely. However, microbial population, community, and functional measurement endpoints change rapidly and across very short spatial scales. Measurement endpoints of microbes tend to be highly responsive to typical fluxes of temperature, moisture, oxygen, and many other noncontaminant factors. Functional redundancy across broad taxonomic groups enables wild swings in community composition without remarkable change in rates of decomposition or community respiration. Consequently, it is exceedingly difficult to relate specific microbial activities with indications of adverse and unacceptable environmental conditions. Moreover, changes in microbial processes do not necessarily result in consequences to plant and animal populations or communities, which in the end are the resources most commonly identified as those to be protected. Therefore, unless more definitive linkages are made between specific microbial effects and an adverse condition for typical assessment endpoint species, microbial endpoints will continue to have limited use in risk assessments; they will not drive the process as primary assessment endpoints.     

大理河沉积物微生物碳源利用特征及其影响因素

沉积物是河流生态系统中氮磷等物质循环的重要场所,而微生物是河流生态系统的重要组成部分,探究沉积物中微生物群落碳源利用特征和功能多样性对于河流生态环境保护具有重要意义。利用Biolog Eco微平板法、基于主成分分析、冗余分析阐明了大理河流域沉积物中微生物群落碳源利用强度和功能多样性变化特征及其影响因素。结果表明：（1）从流域上游到流域下游,沉积物中微生物碳源利用强度逐渐降低,与上游相比,支流、中游、下游沉积物中微生物碳源利用强度分别降低了13.4%、30.5%、30.7%。（2）沉积物中微生物群落功能多样性存在差异,沉积物中微生物群落功能多样性（Shannon-Wiener多样性指数）表现为上游 > 支流 > 中游 > 下游,常见物种优势度（Simpson多样性指数）则表现为下游 > 支流 > 中游 > 上游。（3）与微生物代谢活动相关性较高碳源为糖类,其次是氨基酸类,聚合物类、羧酸类、胺类、酚酸类与微生物代谢活动相关性较低。（4）沉积物中全氮、氨氮、硝氮、有机碳含量是影响微生物群落功能多样性和碳源利用特征差异的主要因素。流域沉积物中合适的碳、氮水平对维持河流水生态健康具有重要的意义。     

Fish assemblages respond to altered flow regimes via ecological filtering of life history strategies

1. In riverine ecosystems, streamflow determines the physical template upon which the life history strategies of biota are forged. Human freshwater needs and activities have resulted in widespread alteration of the variability, predictability and timing of streamflow, and anticipating the biotic consequences of anthropogenic streamflow alteration is critical for successful environmental flow management. In this study, we examined relationships between dam characteristics, metrics of flow alteration and fish functional community composition according to life history strategies by coupling stream flow records and fish survey data in paired flow‐regulated and free‐flowing rivers across the conterminous United States. 2. Dam operations have generally reduced flow variability and increased flow constancy based on a comparison of pre‐ and post‐dam flow records (respective mean record lengths 26.2 and 43.1 years). In agreement with ecological theory, fish assemblages downstream of dams were characterised by a lower proportion of opportunistic species (a strategy favoured in environmental settings dominated by unpredictable environmental change) and a higher proportion of equilibrium species (a strategy favoured in more stable, predictable environments) compared to free‐flowing, neighbouring locations. 3. Multiple linear regression models provided modest support for links between alteration of specific flow attributes and differential life history representation below dams, and they provided strong support for life history associations with dam attributes (age and release type). We also found support for a relationship of both reduced flow variability and dam age with higher representation of non‐native species below dams. 4. Our study demonstrated that river regulation by large dams has significant hydrological and biological consequences across the United States. We showed that on ecological time scales (i.e. the order of years to decades), dams are effectively changing the functional composition of communities that have established over millennia. Furthermore, the changes are directional and indicate a filtering by dams for some life histories (equilibrium strategists) and against other life histories (opportunists). Finally, our study highlights that dependence upon long‐term flow records and availability of biotic surveys extracted from national survey efforts limit our ability to guide environmental flow standards particularly in data‐poor regions.     

Responses of the functional structure of soil microbial community to livestock grazing in the Tibetan alpine grassland

Microbes play key roles in various biogeochemical processes, including carbon (C) and nitrogen (N) cycling. However, changes of microbial community at the functional gene level by livestock grazing, which is a global land‐use activity, remain unclear. Here we use a functional gene array, GeoChip 4.0, to examine the effects of free livestock grazing on the microbial community at an experimental site of Tibet, a region known to be very sensitive to anthropogenic perturbation and global warming. Our results showed that grazing changed microbial community functional structure, in addition to aboveground vegetation and soil geochemical properties. Further statistical tests showed that microbial community functional structures were closely correlated with environmental variables, and variations in microbial community functional structures were mainly controlled by aboveground vegetation, soil C/N ratio, and NH₄⁺‐N. In‐depth examination of N cycling genes showed that abundances of N mineralization and nitrification genes were increased at grazed sites, but denitrification and N‐reduction genes were decreased, suggesting that functional potentials of relevant bioprocesses were changed. Meanwhile, abundances of genes involved in methane cycling, C fixation, and degradation were decreased, which might be caused by vegetation removal and hence decrease in litter accumulation at grazed sites. In contrast, abundances of virulence, stress, and antibiotics resistance genes were increased because of the presence of livestock. In conclusion, these results indicated that soil microbial community functional structure was very sensitive to the impact of livestock grazing and revealed microbial functional potentials in regulating soil N and C cycling, supporting the necessity to include microbial components in evaluating the consequence of land‐use and/or climate changes.     

Human and Environmental Impacts on River Sediment Microbial Communities

Sediment microbial communities are responsible for a majority of the metabolic activity in river and stream ecosystems. Understanding the dynamics in community structure and function across freshwater environments will help us to predict how these ecosystems will change in response to human land-use practices. Here we present a spatiotemporal study of sediments in the Tongue River (Montana, USA), comprising six sites along 134 km of river sampled in both spring and fall for two years. Sequencing of 16S rRNA amplicons and shotgun metagenomes revealed that these sediments are the richest (∼65,000 microbial ‘species’ identified) and most novel (93% of OTUs do not match known microbial diversity) ecosystems analyzed by the Earth Microbiome Project to date, and display more functional diversity than was detected in a recent review of global soil metagenomes. Community structure and functional potential have been significantly altered by anthropogenic drivers, including increased pathogenicity and antibiotic metabolism markers near towns and metabolic signatures of coal and coalbed methane extraction byproducts. The core (OTUs shared across all samples) and the overall microbial community exhibited highly similar structure, and phylogeny was weakly coupled with functional potential. Together, these results suggest that microbial community structure is shaped by environmental drivers and niche filtering, though stochastic assembly processes likely play a role as well. These results indicate that sediment microbial communities are highly complex and sensitive to changes in land use practices.     

Comparative Resistance and Resilience of Soil Microbial Communities and Enzyme Activities in Adjacent Native Forest and Agricultural Soils

Degradation of soil properties following deforestation and long-term soil cultivation may lead to decreases in soil microbial diversity and functional stability. In this study, we investigated the differences in the stability (resistance and resilience) of microbial community composition and enzyme activities in adjacent soils under either native tropical forest (FST) or in agricultural cropping use for 14 years (AGR). Mineral soil samples (0 to 5 cm) from both areas were incubated at 40°C, 50°C, 60°C, or 70°C for 15 min in order to successively reduce the microbial biomass. Three and 30 days after the heat shocks, fluorescein diacetate (FDA) hydrolysis, cellulase and laccase activities, and phospholipid-derived fatty acids-based microbial community composition were measured. Microbial biomass was reduced up to 25% in both soils 3 days after the heat shocks. The higher initial values of microbial biomass, enzyme activity, total and particulate soil organic carbon, and aggregate stability in the FST soil coincided with higher enzymatic stability after heat shocks. FDA hydrolysis activity was less affected (more resistance) and cellulase and laccase activities recovered more rapidly (more resilience) in the FST soil relative to the AGR counterpart. In the AGR soil, laccase activity did not show resilience to any heat shock level up to 30 days after the disturbance. Within each soil type, the microbial community composition did not differ between heat shock and control samples at day 3. However, at day 30, FST soil samples treated at 60°C and 70°C contained a microbial community significantly different from the control and with lower biomass regardless of high enzyme resilience. Results of this study show that deforestation followed by long-term cultivation changed microbial community composition and had differential effects on microbial functional stability. Both soils displayed similar resilience to FDA hydrolysis, a composite measure of a broad range of hydrolases, supporting the concept of high functional redundancy in soil microbial communities. In contrast, the resilience of the substrate-specific activities of laccase and cellulase were lower in AGR soils, indicating a less diverse community of microorganisms capable of producing these enzymes and confirming that specific microbial functions are more sensitive measurements for evaluating change in the ecological stability of soils.     

Protozoan communities of the Flint River-Lake Blackshear ecosystem (Georgia,USA)

Freshwater protozoa are poorly characterized in river ecosystems. We report here the richness of the protozoan biotas in relation to environmental gradients from an ecosystematic survey of a large, coastal plain river. Communities were collected from natural and artificial substrates concurrent with water chemistry analysis at 11 sites along the Flint River and Lake Blackshear impoundment. Community similarity, the distribution of collected taxa in functional feeding groups, and the relation of communities to environmental gradients were evaluated. Two principal compenents determined from water chemistry data showed important downstream gradients of decreasing water hardness and increasing nutrient levels. Taxonomic richness was high; 200 to 450 taxa were collected depending on season and collecting technique. Artificial substrates provided the richest collections. Bactivorous species were the vast majority of all taxa collected. Community composition showed an orderly transition from upstream to downstream, and photosynthetic forms were enhanced at nutrient enriched sites. Communities were strongly influenced by increasing nutrient levels. Protozoan community analysis showed that microbial community composition reflects human influences on river ecosystems. Since microbial species exploit detrital resources and respond sensitively to human influences, they can provide important information regarding ecosystem conditions.     

Temporal variability in the spatial and environmental determinants of functional metacommunity organization – stream fish in a human‐modified landscape

1. Quantifying the relative importance of environmental filtering versus regional spatial structuring has become an intensively studied area in the context of metacommunity ecology. However, most studies have evaluated the role of environmental and spatial processes using taxonomic data sets of single snapshot surveys. 2. Here, we examined temporal changes in patterns and possible processes behind the functional metacommunity organization of stream fishes in a human‐modified landscape. Specifically, we (i) studied general changes in the functional composition of fish assemblages among 40 wadeable stream sites during a 3‐year study period in the catchment area of Lake Balaton, Hungary, (ii) quantified the relative importance of spatial and environmental factors as determinants of metacommunity structure and (iii) examined temporal variability in the relative role of spatial and environmental processes for this metacommunity. 3. Partial triadic analysis showed that assemblages could be effectively ordered along a functional gradient from invertebrate consuming species dominated by the opportunistic life‐history strategy, to assemblages with a diverse array of functional attributes. The analysis also revealed that functional fish assemblage structure was moderately stable among the sites between the sampling periods. 4. Despite moderate stability, variance partitioning using redundancy analyses (RDA) showed considerable temporal variability in the contribution of environmental and spatial factors to this pattern. The analyses also showed that environmental variables were, in general, more important than spatial ones in determining metacommunity structure. Of these, natural environmental variables (e.g. altitude, velocity) proved to be more influential than human‐related effects (e.g. pond area, % inhabited area above the site, nutrient enrichment), even in this landscape with relatively low variation in altitude and stream size. 5. Pond area was, however, the most important human stressor variable that was positively associated with the abundance of non‐native species with diverse functional attributes. The temporal variability in the relative importance of environmental and spatial factors was probably shaped by the release of non‐native fish from fish ponds to the stream system during flood events. 6. To conclude, both spatial processes and environmental control shape the functional metacommunity organization of stream fish assemblages in human‐modified landscapes, but their importance can vary in time. We argue, therefore, that metacommunity studies should better consider temporal variability in the ecological mechanisms (e.g. dispersal limitation, species sorting) that determine the dynamics of landscape‐level community organization.     

Fish species introductions provide novel insights into the patterns and drivers of phylogenetic structure in freshwaters

Despite long-standing interest of terrestrial ecologists, freshwater ecosystems are a fertile, yet unappreciated, testing ground for applying community phylogenetics to uncover mechanisms of species assembly. We quantify phylogenetic clustering and overdispersion of native and non-native fishes of a large river basin in the American Southwest to test for the mechanisms (environmental filtering versus competitive exclusion) and spatial scales influencing community structure. Contrary to expectations, non-native species were phylogenetically clustered and related to natural environmental conditions, whereas native species were not phylogenetically structured, likely reflecting human-related changes to the basin. The species that are most invasive (in terms of ecological impacts) tended to be the most phylogenetically divergent from natives across watersheds, but not within watersheds, supporting the hypothesis that Darwin''s naturalization conundrum is driven by the spatial scale. Phylogenetic distinctiveness may facilitate non-native establishment at regional scales, but environmental filtering restricts local membership to closely related species with physiological tolerances for current environments. By contrast, native species may have been phylogenetically clustered in historical times, but species loss from contemporary populations by anthropogenic activities has likely shaped the phylogenetic signal. Our study implies that fundamental mechanisms of community assembly have changed, with fundamental consequences for the biogeography of both native and non-native species.     

Seasonal shifts in the assembly dynamics of benthic macroinvertebrate and diatom communities in a subtropical river

Identifying seasonal shifts in community assembly for multiple biological groups is important to help enhance our understanding of their ecological dynamics. However, such knowledge on lotic assemblages is still limited. In this study, we used biological traits and functional diversity indices in association with null model analyses to detect seasonal shifts in the community assembly mechanisms of lotic macroinvertebrates and diatoms in an unregulated subtropical river in China. We found that functional composition and functional diversity (FRic, FEve, FDis, MNN, and SDNN) showed seasonal variation for macroinvertebrate and diatom assemblages. Null models suggested that environmental filtering, competitive exclusion, and neutral process were all important community assembly mechanisms for both biological groups. However, environmental filtering had a stronger effect on spring macroinvertebrate assemblages than autumn assemblages, but the effect on diatom assemblages was the same in both seasons. Moreover, macroinvertebrate and diatom assemblages were shaped by different environmental factors. Macroinvertebrates were filtered mainly by substrate types, velocity, and COD_Mn, while diatoms were mainly shaped by altitude, substrate types, and water quality. Therefore, our study showed (a) that different biological assemblages in a river system presented similarities and differences in community assembly mechanisms, (b) that multiple processes play important roles in maintaining benthic community structure, and (c) that these patterns and underlying mechanisms are seasonally variable. Thus, we highlight the importance of exploring the community assembly mechanisms of multiple biological groups, especially in different seasons, as this is crucial to improve the understanding of river community changes and their responses to environmental degradation.     

Altitudinal patterns of diversity and functional traits of metabolically active microorganisms in stream biofilms

Resources structure ecological communities and potentially link biodiversity to energy flow. It is commonly believed that functional traits (generalists versus specialists) involved in the exploitation of resources depend on resource availability and environmental fluctuations. The longitudinal nature of stream ecosystems provides changing resources to stream biota with yet unknown effects on microbial functional traits and community structure. We investigated the impact of autochthonous (algal extract) and allochthonous (spruce extract) resources, as they change along alpine streams from above to below the treeline, on microbial diversity, community composition and functions of benthic biofilms. Combining bromodeoxyuridine labelling and 454 pyrosequencing, we showed that diversity was lower upstream than downstream of the treeline and that community composition changed along the altitudinal gradient. We also found that, especially for allochthonous resources, specialisation by biofilm bacteria increased along that same gradient. Our results suggest that in streams below the treeline biofilm diversity, specialisation and functioning are associated with increasing niche differentiation as potentially modulated by divers allochthonous and autochthonous constituents contributing to resources. These findings expand our current understanding on biofilm structure and function in alpine streams.     

Microbial minorities modulate methane consumption through niche partitioning

Microbes catalyze all major geochemical cycles on earth. However, the role of microbial traits and community composition in biogeochemical cycles is still poorly understood mainly due to the inability to assess the community members that are actually performing biogeochemical conversions in complex environmental samples. Here we applied a polyphasic approach to assess the role of microbial community composition in modulating methane emission from a riparian floodplain. We show that the dynamics and intensity of methane consumption in riparian wetlands coincide with relative abundance and activity of specific subgroups of methane-oxidizing bacteria (MOB), which can be considered as a minor component of the microbial community in this ecosystem. Microarray-based community composition analyses demonstrated linear relationships of MOB diversity parameters and in vitro methane consumption. Incubations using intact cores in combination with stable isotope labeling of lipids and proteins corroborated the correlative evidence from in vitro incubations demonstrating γ-proteobacterial MOB subgroups to be responsible for methane oxidation. The results obtained within the riparian flooding gradient collectively demonstrate that niche partitioning of MOB within a community comprised of a very limited amount of active species modulates methane consumption and emission from this wetland. The implications of the results obtained for biodiversity–ecosystem functioning are discussed with special reference to the role of spatial and temporal heterogeneity and functional redundancy.     

Environmental associations of abundance-weighted functional traits in Australian plant communities

Predictions of how vegetation responds to spatial and temporal differences in climate rely on established links with plant functional traits and vegetation types that can be encoded into Dynamic Global Vegetation Models. Individual traits have been linked to climate at species level and at community level within regions. However, a recent global assessment of aggregated community level traits found unexpectedly weak links with macroclimate, bringing into question broadscale trait–climate associations and implicating local-scale environmental differences in the filtering of communities. To further evaluate patterns in light of these somewhat contradictory results, we quantified the power of macro-environmental variables to explain aggregated plant community traits, taking advantage of new trait data for leaf area, plant height and seed mass combined with a national survey that records cover-abundance using consistent methods for a large number of plots across Australia. In contrast to the global study, we found that abundance-weighted community mean and variance of leaf area and maximum height were correlated with macroclimate. Height and leaf area were highest in wet (especially warm, wet) environments, with actual evapotranspiration explaining 30% of variation in leaf area and 26% in maximum height. Seed mass was weakly related to environment, with no variable explaining more than 5% of variance. Considering all three traits together in a redundancy analysis, the complete set of environmental variables explained 43% of variation in site-mean traits and 29% of within-site trait variance. While significant trait variation remains unexplained, the trait–environment relationships reported here suggest climatically-driven filtering plays a strong role in assembling these vegetation communities. Regional assessments using standardised species abundances can therefore be used to predict aspects of vegetation function. Our quantification of plant community trait patterns along macroclimatic gradients at continental scale thereby contributes a much-needed functional basis for Australian vegetation.     

Phylogenetic community structure in Minnesota oak savanna is influenced by spatial extent and environmental variation

The relative importance of environmental filtering, biotic interactions and neutral processes in community assembly remains an openly debated question and one that is increasingly addressed using phylogenetic approaches. Closely related species may occur together more frequently than expected (phylogenetic clustering) if environmental filtering operates on traits with significant phylogenetic signal. Recent studies show that phylogenetic clustering tends to increase with spatial scale, presumably because greater environmental variation is encompassed at larger spatial scales, providing opportunities for species to sort across environmental gradients. However, if environmental filtering is the cause of species sorting along environmental gradients, then environmental variation rather than spatial scale per se should drive the processes governing community assembly. Using species abundance and light availability data from a long‐term experiment in Minnesota oak savanna understory communities, we explicitly test the hypothesis that greater environmental variation results in greater phylogenetic clustering when spatial scale is held constant. Concordant with previous studies, we found that phylogenetic community structure varied with spatial extent. At the landscape scale (~1000 ha), communities were phylogenetically clustered. At the local scale (0.375ha), phylogenetic community structure varied among plots. As hypothesized, plots encompassing the greatest environmental variation in light availability exhibited the strongest phylogenetic clustering. We also found strong correlations between species functional traits, particularly specific leaf area (SLA) and perimeter per area (PA), and species light availability niche. There was also a phylogenetic signal in both functional traits and species light availability niche, providing a mechanistic explanation for phylogenetic clustering in relation to light availability. We conclude that the pattern of increased phylogenetic clustering with increased environmental variation is a consequence of environmental filtering acting on phylogenetically conserved functional traits. These results indicate that the importance of environmental filtering in community assembly depends not on spatial scale per se, but on the steepness of the environmental gradient.     

漓江河岸带植物功能性状变异与关联

研究植物功能性状随环境梯度的变异和关联格局, 对于认识不同环境梯度下群落构建和植物适应型具有重要意义。该研究以漓江河岸带不同河段植物群落为研究对象, 调查了研究区内36个样方的物种组成, 测量了样方内42种木本植物的叶面积(LA)、比叶面积(SLA)和木材密度(WD)的功能性状值, 并运用性状梯度分析法对3个功能性状进行群落内(α组分)和群落间(β组分)组分分解及相关性分析。结果表明: (1)群落平均LA表现为中游最小且和下游差异显著, 群落平均WD则表现为中上游显著高于下游, 群落平均SLA在两两河段间均差异显著。(2)不同河段的3个植物功能性状β组分差异显著且实际观测值均小于随机模拟的零模型分布, 但α组分在河岸带不同河段均差异不显著且3个功 能性状的α组分分布范围均小于β组分, 说明在河岸带不同河段的群落构建过程中环境筛选的作用要大于群落内种间的相互作用。(3)性状SLA与LA在群落间和群落内呈现出实际观测和随机模拟的相关性均较低, 暗示了LA和SLA各自代表了植物在不同生态策略上的维度; 但SLA和WD实际观测值和随机模拟值呈现出较强的负相关关系, 暗示这2个性状对于环境筛选表现出较高的整体趋同适应性, 体现了植物功能性状对群落间环境变异的依赖性大于群落内种间相互作用的依赖性。     

Vulnerability of macronutrients to the concurrent effects of enhanced temperature and atmospheric pCO<Subscript>2</Subscript> in representative shelf sea sediment habitats

Fundamental changes in seawater carbonate chemistry and sea surface temperatures associated with the ocean uptake of anthropogenic CO₂ are accelerating, but investigations of the susceptibility of biogeochemical processes to the simultaneous occurrence of multiple components of climate change are uncommon. Here, we quantify how concurrent changes in enhanced temperature and atmospheric pCO₂, coupled with an associated shift in macrofaunal community structure and behavior (sediment particle reworking and bioirrigation), modify net carbon and nutrient concentrations (NH₄-N, NO_x-N, PO₄-P) in representative shelf sea sediment habitats (mud, sandy-mud, muddy-sand and sand) of the Celtic Sea. We show that net concentrations of organic carbon, nitrogen and phosphate are, irrespective of sediment type, largely unaffected by a simultaneous increase in temperature and atmospheric pCO₂. However, our analyses also reveal that a reduction in macrofaunal species richness and total abundance occurs under future environmental conditions, varies across a gradient of cohesive to non-cohesive sediments, and negatively moderates biogeochemical processes, in particular nitrification. Our findings indicate that future environmental conditions are unlikely to have strong direct effects on biogeochemical processes but, particularly in muddy sands, the abundance, activity, composition and functional role of invertebrate communities are likely to be altered in ways that will be sufficient to regulate the function of the microbial community and the availability of nutrients in shelf sea waters.     

Quantifying structural redundancy in ecological communities

In multivariate analyses of the effects of both natural and anthropogenic environmental variability on community composition, many species are interchangeable in the way that they characterise the samples, giving rise to the concept of structural redundancy in community composition. Here, we develop a method of quantifying the extent of this redundancy by extracting a series of subsets of species, the multivariate response pattern of each of which closely matches that for the whole community. Structural redundancy is then reflected in the number of such subsets, which we term “response units”, that can be extracted without replacement. We have applied this technique to the effects of the Amoco-Cadiz oil-spill on marine macrobenthos in the Bay of Morlaix, France, and to the natural interannual variability of macrobenthos at two stations off the coast of Northumberland, England. Structural redundancy is shown to be remarkably high, with the number and sizes of subsets being comparable in all three examples. Taxonomic/functional groupings of species within the differing response units change in abundance in the same way over time. The response units are shown to possess a wide taxonomic spread and, using two different types of randomisation test, demonstrated to have a taxonomically and functionally coherent structure. The level of structural redundancy may therefore be an indirect measure of the resilience or compensation potential within an assemblage. Received: 23 January 1996 / Accepted: 14 July 1997