Non-additive, identity-dependent effects of detrital species mixing on soft-sediment communities

Accelerating rates of species extinction and invasion have sparked recent interest in how changes in plant community composition can be propagated through food webs. Research in this area has, however, been largely restricted to considerations of how detrital species mixing affects litter decay processes. The consequences of changing detrital resources for whole assemblages of sediment‐dwelling invertebrates remain largely unknown. We manipulated the availability of three detrital sources, Avicennia marina leaves, Posidonia australis blades and Sargassum sp. thalli, on an Australian mudflat to test hypotheses about how changes in the type and number of macrophytes contributing to detrital resources might impact benthic invertebrate assemblages of estuarine soft‐sediments. By controlling for changes in total detrital biomass and ensuring that each detrital source was present in two‐ and three‐species mixes as well as monocultures, our experimental design was able to distinguish among effects of mixing, identity and biomass. Three months after detrital manipulation, macroinvertebrate abundance and species richness differed among treatments according to the biomass of detritus added and non‐additive effects of detrital species mixing. Whereas the mixing of two detrital species generally had an antagonistic effect on macroinvertebrate abundance and richness, faunal assemblages did not appreciably differ between three‐species mixes and monocultures. Generally negative effects of two‐species mixes on macroinvertebrates were opposed by positive effects on microphytobenthos, an important food‐source for many of the animals. Non‐additive effects on sediment communities were particularly apparent when Sargassum sp., the most labile of the three detrital sources considered, was included in two‐species mixes. This demonstration of non‐additive and identity‐dependent effects of detrital species mixing on soft‐sediment communities suggests that predicted compositional changes to aquatic macrophyte communities, resulting from coastal development and climate change, will flow on to effect other components of the estuarine food‐web.     

Detrital diversity influences estuarine ecosystem performance

Global losses of seagrasses and mangroves, eutrophication‐driven increases in ephemeral algae, and macrophyte invasions have impacted estuarine detrital resources. To understand the implications of these changes on benthic ecosystem processes, we tested the hypotheses that detrital source richness, mix identity, and biomass influence benthic primary production, metabolism, and nutrient fluxes. On an estuarine muddy sandflat, we manipulated the availability of eight detrital sources, including mangrove, seagrass, and invasive and native algal species that have undergone substantial changes in distribution. Mixes of these detrital sources were randomly assigned to one of 12 treatments and dried detrital material was added to seventy‐two 0.25 m² plots (n = 6 plots). The treatments included combinations of either two or four detrital sources and high (60 g) or low (40 g) levels of enrichments. After 2 months, the dark, light, and net uptake of NH₄⁺, dissolved inorganic nitrogen, and the dark efflux of dissolved organic nitrogen were each significantly influenced by the identity of detrital mixes, rather than detrital source richness or biomass. However, gross and net primary productivity, average oxygen flux, and net NO_X and dissolved inorganic phosphorous fluxes were significantly greater in treatments with low than with high detrital source richness. These results demonstrate that changes in detrital source richness and mix identity may be important drivers of estuarine ecosystem performance. Continued impacts to estuarine macrophytes may, therefore, further alter detritus‐fueled productivity and processes in estuaries. Specific tests that address predicted future changes to detrital resources are required to determine the consequences of this significant environmental problem.     

Consumer and resource diversity effects on marine macroalgal decomposition

Few manipulative experiments that explicitly test the relationship between biodiversity and ecosystem function have focussed on regenerative processes such as decomposition and nutrient cycling. Of the studies that have taken place, most have concentrated on the effects of leaf litter diversity rather than the effects of consumer diversity on decomposition. In the present study, we established an in‐situ mesocosm experiment on an intertidal mudflat in the Ythan Estuary, Scotland, to investigate the interactive effects of consumer diversity, resource diversity and microbial activity on algal consumption and decomposition. We assembled communities of three commonly occurring macrofaunal species (Hediste diversicolor, Hydrobia ulvae and Littorina littorea) in monoculture and all possible combinations of two and three species mixtures and supplied them with single vs two‐species mixtures of the algae Fucus spiralis and Ulva intestinalis. Further, we also investigated whether algal decomposition changes nutrient remineralisation within the sediment by determining the C:N ratio of the surficial sediment. Data were analysed using extended linear regression with generalized least squares estimation to characterise the variance structure. We found that consumer species diversity effects are best explained by compositional effects and that species richness per se may not be the single most important determinant of resource use and decomposition in this community. Algal identity and invertebrate identity effects underpin the observed response and reflect species‐specific traits associated with algal consumption and processing. The role of the microbial community is comparatively weak, but strongly linked to faunal activities and behaviour. The C:N ratio of the sediment increased with consumer species richness, indicating increased mineralisation in more diverse communities. Overall, our results suggest that although consumer species richness effects per se are weak, decomposition and subsequent incorporation of resources is nevertheless dependent on the composition of the decomposer community, which, in turn, has important implications for biogeochemical nutrient cycling in marine coastal habitats.     

Effects of stream predator richness on the prey community and ecosystem attributes

It is important to understand the role that different predators can have to be able to predict how changes in the predator assemblage may affect the prey community and ecosystem attributes. We tested the effects of different stream predators on macroinvertebrates and ecosystem attributes, in terms of benthic algal biomass and accumulation of detritus, in artificial stream channels. Predator richness was manipulated from zero to three predators, using two fish and one crayfish species, while density was kept equal (n = 6) in all treatments with predators. Predators differed in their foraging strategies (benthic vs. drift feeding fish and omnivorous crayfish) but had overlapping food preferences. We found effects of both predator species richness and identity, but the direction of effects differed depending on the response variable. While there was no effect on macroinvertebrate biomass, diversity of predatory macroinvertebrates decreased with increasing predator species richness, which suggests complementarity between predators for this functional feeding group. Moreover, the accumulation of detritus was affected by both predator species richness and predator identity. Increasing predator species richness decreased detritus accumulation and presence of the benthic fish resulted in the lowest amounts of detritus. Predator identity (the benthic fish), but not predator species richness had a positive effect on benthic algal biomass. Furthermore, the results indicate indirect negative effects between the two ecosystem attributes, with a negative correlation between the amount of detritus and algal biomass. Hence, interactions between different predators directly affected stream community structure, while predator identity had the strongest impact on ecosystem attributes.     

Aggregation of Cricket Activity in Response to Resource Addition Increases Local Diversity

Crickets are often found feeding on fallen fruits among forest litter. Fruits and other sugar-rich resources are not homogeneously distributed, nor are they always available. We therefore expect that crickets dwelling in forest litter have a limited supply of sugar-rich resource, and will perceive this and displace towards resource-supplemented sites. Here we evaluate how sugar availability affects cricket species richness and abundance in old-growth Atlantic forest by spraying sugarcane syrup on leaf litter, simulating increasing availability, and collecting crickets via pitfall trapping. We found an asymptotic positive association between resource addition and species richness, and an interaction between resource addition and species identity on cricket abundance, which indicates differential effects of resource addition among cricket species. Our results indicate that 12 of the 13 cricket species present in forest litter are maintained at low densities by resource scarcity; this highlights sugar-rich resource as a short-term driver of litter cricket community structure in tropical forests. When resource was experimentally increased, species richness increased due to behavioral displacement. We present evidence that the density of many species is limited by resource scarcity and, when resources are added, behavioral displacement promotes increased species packing and alters species composition. Further, our findings have technical applicability for increasing sampling efficiency of local cricket diversity in studies aiming to estimate species richness, but with no regard to local environmental drivers or species-abundance characteristics.     

Detrital traits affect substitutability of a range‐expanding foundation species across latitude

Climate‐driven range shifts of foundation species could alter ecosystem processes and community composition by providing different resources than resident foundation species. Along the US Atlantic coast, the northward expanding foundation species, black mangrove Avicennia germinans, is replacing the dominant salt marsh foundation species, marsh cordgrass Spartina alterniflora. These species have distinct detrital attributes that ostensibly provide different resources to epifauna. We experimentally examined how detritus of these species affects decomposition and community composition in different habitat contexts at regional and local scales. First, we manipulated detritus identity (Avicennia, Spartina) at 13 sites across a 5° latitudinal gradient spanning mangrove, mixed marsh‐mangrove and salt marsh habitats. Across latitude, we found that Avicennia detritus decomposed 2–4 times faster than Spartina detritus, suggesting that detrital turnover will increase with mangrove expansion. Epifaunal abundance and richness increased 2–7 times from south to north (mangrove to salt marsh) and were equivalent between Avicennia and Spartina detritus except for crabs, a dominant taxonomic group that preferred Spartina detritus. Second, to examine the whether changing habitat context affected regional patterns, we manipulated detritus identity and surrounding habitat type (mangrove, salt marsh) at a single mixed site, also including inert mimics to separate structural and nutritional roles of detritus. Epifaunal richness was similar between the two detrital types, but crabs were 2–7 times more abundant in Spartina detritus due to its structural attributes. Surrounding habitat type did not influence decomposition rate or community patterns, which suggests that latitudinal influences, not surrounding habitat, drove the regional community patterns in the first experiment. Overall, mangrove expansion could alter epifaunal communities due to the lower structural value and faster turnover of mangrove detritus. As species shift with changing climate, understanding foundation species substitutability is critical to predict community change, but we must account for concomitant environmental changes that also modify communities.     

Biodiversity and tallgrass prairie decomposition: the relative importance of species identity, evenness, richness, and micro-topography

Biodiversity has been declining in many areas, and there is great interest in determining whether this decline affects ecosystem functioning. Most biodiversity—ecosystem functioning studies have focused on the effects of species richness on net primary productivity. However, biodiversity encompasses both species richness and evenness, ecosystem functioning includes other important processes such as decomposition, and the effects of richness on ecosystem functioning may change at different levels of evenness. Here, we present two experiments on the effects of litter species evenness and richness on litter decomposition. In the first experiment, we varied the species evenness (three levels), identity of the dominant species (three species), and micro-topographic position (low points [gilgais] or high points between gilgais) of litter in three-species mixtures in a prairie in Texas, USA. In a second experiment, we varied the species evenness (three levels), richness (one, two, or four species per bag), and composition (random draws) of litter in a prairie in Iowa, USA. Greater species evenness significantly increased decomposition, but this effect was dependent on the environmental context. Higher evenness increased decomposition rates only under conditions of higher water availability (in gilgais in the first experiment) or during the earliest stages of decomposition (second experiment). Species richness had no significant effect on decomposition, nor did it interact with evenness. Micro-topographic position and species identity and composition had larger effects on decomposition than species evenness. These results suggest that the effects of litter species diversity on decomposition are more likely to be manifested through the evenness component of diversity than the richness component, and that diversity effects are likely to be environmentally context dependent.     

Phylogenetic distance among beneficiary species in a cushion plant species explains interaction outcome

Determining which drivers lead to a specific species assemblage is a central issue in community ecology. Although many processes are involved, plant–plant interactions are among the most important. The phylogenetic limiting similarity hypothesis states that closely related species tend to compete stronger than distantly related species, although evidence is inconclusive. We used ecological and phylogenetic data on alpine plant communities along an environmental severity gradient to assess the importance of phylogenetic relatedness in affecting the interaction between cushion plants and the whole community, and how these interactions may affect community assemblage and diversity. We first measured species richness and individual biomass of species growing within and outside the nurse cushion species, Arenaria tetraquetra. We then assembled the phylogenetic tree of species present in both communities and calculated the phylogenetic distance between the cushion species and its beneficiary species, as well as the phylogenetic community structure. We also estimated changes in species richness at the local level due to the presence of cushions. The effects of cushions on closely related species changed from negative to positive as environmental conditions became more severe, while the interaction with distantly related species did not change along the environmental gradient. Overall, we found an environmental context‐dependence in patterns of phylogenetic similarity, as the interaction outcome between nurses and their close and distantly‐related species showed an opposite pattern with environmental severity.     

空间和环境因子对黄河口自然和淡水恢复湿地底栖动物群落的差异影响

生物多样性的形成和维持机制是生态学研究的核心问题,其中环境和空间因子在群落构建中的相对重要性是生态学家面临的重要挑战。为探究黄河口湿地底栖动物群落的关键影响因子,及环境和空间因子对底栖动物群落结构的相对调控作用。于2017年10月与2018年5月对黄河口湿地32个样点（淡水恢复湿地19个和自然湿地13个）的底栖动物和水体理化指标进行采集分析。非度量多维标度排序（NMDS）结果显示,黄河口淡水恢复湿地和自然湿地的底栖动物群落结构显著不同。典范对应分析（CCA）表明,影响淡水恢复湿地底栖动物群落结构的环境因子主要为电导率、盐度和氧化还原电位;而自然湿地底栖动物群落结构主要受pH和无机碳的影响;盐度是两类湿地底栖动物群落组成差异的关键因子。变差分解（VPA）结果显示,环境过滤对淡水恢复湿地底栖动物群落起主导作用;在自然湿地中,空间因子对底栖动物群落具有主要的调控作用,同时环境和空间因子的相互作用也至关重要。本研究明确了黄河口的自然和恢复湿地中环境和空间因素对底栖动物群落特征的相对作用,对黄河三角洲河口湿地中生物多样性的保护和生态系统管理提供参考。     

Replacement of native seagrass with invasive algal detritus: impacts to estuarine sediment communities

Biological invasions modify the quality and supply of detrital subsidies to aquatic and terrestrial ecosystems. Where the invader has very different traits to native species, major changes in associated consumer communities may result, as a consequence of differences in their nutritional value and effects on the sedimentary habitat. We assessed how the replacement of seagrasses with the invasive alga Caulerpa taxifolia in modified Australian estuaries influences invertebrate communities of mudflats that are subsidized by detritus from submerged aquatic vegetation. Two months after experimental enrichment of sediments with high (60?g dry weight per 0.25?m² plot) or low (30?g dry weight) quantities of either non-native C. taxifolia or native Posidonia australis or Zostera capricorni detritus, there were positive effects of detrital addition on invertebrate abundance that occurred irrespective of the resource added. By 4?months after addition, however, detritus from invasive C. taxifolia had produced effects on benthic communities that could not be replicated by detritus from either of the native seagrasses. Plots receiving the high loading of C. taxifolia detritus contained fewer invertebrates than plots of the other treatments, perhaps due to the induction of sediment hypoxia. The pattern, however, reversed at low detrital loading, with the plots receiving 30?g of C. taxifolia containing more invertebrates and more taxa than the other plots, presumably due to the greater resource availability for detritivores. Our results demonstrate that replacement of native seagrass with invasive algal detritus can have large impacts on sediment-dwelling communities.     

Species traits and environmental characteristics together regulate ant‐associated biodiversity

Host‐associated organisms (e.g., parasites, commensals, and mutualists) may rely on their hosts for only a portion of their life cycle. The life‐history traits and physiology of hosts are well‐known determinants of the biodiversity of their associated organisms. The environmental context may strongly influence this interaction, but the relative roles of host traits and the environment are poorly known for host‐associated communities. We studied the roles of host traits and environmental characteristics affecting ant‐associated mites in semi‐natural constructed grasslands in agricultural landscapes of the Midwest USA. Mites are frequently found in ant nests and also riding on ants in a commensal dispersal relationship known as phoresy. During nonphoretic stages of their development, ant‐associated mites rely on soil or nest resources, which may vary depending on host traits and the environmental context of the colony. We hypothesized that mite diversity is determined by availability of suitable host ant species, soil detrital resources and texture, and habitat disturbance. Results showed that that large‐bodied and widely distributed ant species within grasslands support the most diverse mite assemblages. Mite richness and abundance were predicted by overall ant richness and grassland area, but host traits and environmental predictors varied among ant hosts: mites associated with Aphaenogaster rudis depended on litter depth, while Myrmica americana associates were predicted by host frequency and grassland age. Multivariate ordinations of mite community composition constructed with host ant species as predictors demonstrated host specialization at both the ant species and genus levels, while ordinations with environmental variables showed that ant richness, soil texture, and grassland age also contributed to mite community structure. Our results demonstrate that large‐bodied, locally abundant, and cosmopolitan ant species are especially important regulators of phoretic mite diversity and that their role as hosts is also dependent on the context of the interaction, especially soil resources, texture, site age, and area.     

Partitioning the effects of environmental and spatial heterogeneity on distribution of plant diversity in the Yellow River Estuary

For successful conservation and restoration of biodiversity, it is important to understand how diversity is regulated. In the ecological research community, a current topic of interest is how much of the variation in plant species richness and composition is explained by environmental variation (niche-based model), relative to spatial processes (neutral theory). The Yellow River Estuary (YRE) is a newly formed and fragile wetland ecosystem influenced by both the Yellow River and Bohai Bay. Here, we applied variance partitioning techniques to assess the relative effects of spatial and environmental variables on species richness and composition in the YRE. We also conducted a species indicator analysis to identify characteristic species for three subestuaries within the YRE. Partial redundancy analysis showed that the variations in species richness and composition were explained by both environmental and spatial factors. The majority of explained variation in species richness and composition was attributable to local environmental factors. Among the environmental variables, soil salinity made the greatest contribution to species abundance and composition. Soil salinity was the most important factor in the Diaokou subestuary, while soil moisture was the most important factor influencing species richness in the Qingshui and Chahe subestuaries. The combined effects of soil salinity and moisture determined species richness and composition in the wetlands. These results increase our understanding of the organization and assembly of estuarine plant communities.     

A biological trait approach to assess the functional composition of subtidal benthic communities in an estuarine ecosystem

Within transitional/estuarine environments ‘ecosystem functioning’ has been mostly investigated with “traditional” taxonomic analysis, based on the taxonomic composition of benthic invertebrate communities. However, ‘ecosystem functioning’ depends also greatly on the functional characteristics (biological traits) of organisms.It was a priori suggested that the biological traits of the subtidal benthic invertebrate communities within an estuarine environment would respond to the high variability of environmental pressures (natural and human induced) within this type of ecosystem.For this study, traditional taxonomic analysis (species richness, species density and Shannon–Wiener diversity) as well as biological trait analysis were used together for the first time to investigate the response of the subtidal benthic invertebrate communities to the environmental pressures within the Mondego estuary (Portugal).Biological trait analysis, in addition to traditional taxonomic analysis provided a more comprehensive understanding of the functioning within this type of ecosystem. Some of the most important outcomes are: (i) the trait “salinity preference” was the most important trait that distributed the species along the estuary, (ii) the central part of the estuary appeared to be under higher environmental stress levels than the other areas, as suggested by a dominance of some “opportunistic” traits (e.g. small short-lived species), (iii) the ratio between functional diversity (FD) and Shannon–Wiener diversity (H′) indicated lower functional redundancy at the upper reaches of the estuary. Our results, suggest that the ratio (FD/H′) might be a helpful tool to visualize this functional attribute and could potentially be applied to different communities from distinct environments. Using the traditional taxonomic analysis alone, this last functional aspect would not be detectable. Therefore, the inclusion of biological traits analysis is recommendable for estuarine ecological studies.     

Spatial Patterns of Benthic Macroinvertebrates in Intertidal Areas of a Southern European Estuary: The Tagus, Portugal

This study characterizes the composition and spatial distribution patterns of the benthic macrofauna in the intertidal mudflats of the Tagus estuary, western Portugal. A total of 68 species, more than 226,000 specimens with a total wet weight biomass of approximately 1170 g were identified in 380 sites. The species Streblospio shrubsolii, Cyathura carinata, Tharyx sp., Hydrobia ulvae and Tubificids were the most common and abundant. Scrobicularia plana strongly dominated the biomass. The invertebrate macrofauna of the Tagus estuary shows similarities to what is known from other temperate mudflats. The diversity of species, their overall abundance and the ratio of Molluscs plus Crustaceans to Polychaete species corroborate the distinctiveness between temperate and tropical mudflats and sandflats. The spatial distribution of the fauna reflects the sediment characteristics but the relationship between the environmental and the biological data is not as strong as obtained for sublittoral areas. This relationship diminishes from the sublittoral shelf to sublittoral estuarine areas, showing minimum values in this study, suggesting that such a relationship is less straightforward as natural disturbance increases. Nevertheless, a mixture of grain-size, elevation (inundation time) and particular habitats (relic oyster beds) form the best explanatory factors for the spatial distribution patterns of the intertidal benthic macrofauna of the Tagus estuary.     

Genetic and community similarities are correlated in endemic-rich springs of the northern Chihuahuan Desert

Aim Species diversity and genetic diversity within a taxon are intrinsic parts of global biodiversity. These two levels of biodiversity can show strong correlation due to a variety of reasons (i.e. parallel processes affecting both communities and populations, genotypes of a numerically or functionally dominant species affecting community composition, a species assemblage selecting for a particular genotype by affecting its selection regime). We examined correlations between species and genetic biodiversity in four isolated endemic‐rich spring systems in a hot desert and their potential link to environmental variables and physical isolation. Location Chihuahuan Desert spring systems in the Pecos River basin of New Mexico and Texas, USA. Methods We compared species richness of fish and benthic macroinvertebrate communities to within‐population allelic richness of amphipods (monophyletic Gammarus spp.) and Pecos gambusia (Gambusia nobilis) using Mantel tests. We also compared pairwise community similarities with pairwise genetic identities of populations among the same groups. We tested correlations among diversity, similarity and environmental variables after controlling for the effects of spatial distances using partial Mantel tests. We partitioned genetic and species diversity into three spatial scales (i.e. individual springs, individual spring systems, the entire region) using AMOVA and partition . Results We found strong correlations between invertebrate species richness and mosquitofish allelic richness. We found even stronger correlations of amphipod and gambusia genetic identities with fish and invertebrate community similarities; these were best explained by geographic distance rather than abiotic environmental factors. Most of the taxa and communities exhibited the largest proportion of diversity at the regional level. Main conclusions Our results suggest that drift and migration are the mechanisms that best explain our observations, and although α‐diversity among genes and species may not be strongly correlated, the pattern of species and allelic complementarity among these groups seems to be concordant at the regional level.     

Limited evidence that mixing leaf litter accelerates decomposition or increases diversity of decomposers in streams of eastern Canada

Most studies of terrestrial litter decomposition in streams and rivers have used leaves from a single tree species, but leaf packs in streams in eastern North America are usually mixtures of two or more species. Litter mixtures may decay more quickly than either of the component species. If so, estimates of stream energy and nutrient budgets may be inaccurate. In northern Nova Scotia, Canada, we measured mass loss from binary mixtures (1:1 mass ratio) of leaf litter in mesh bags, using freshly fallen or air-dried litter from five species of canopy trees. We repeated the experiment eight times, in summer and fall, in two streams and a small river, over 3 years. In some trials we enumerated benthic invertebrate and fungal colonization of decaying litter. Although there were marked differences in mass loss rates among litter types, decomposition was accelerated in mixtures relative to the mean of the component species in only three of eight trials, and only in mixtures containing N-rich speckled alder leaves. Mixing yellow birch and red maple leaves inhibited decomposition. Diversity (Shannon–Weaver Index), species richness, and abundance of aquatic hyphomycete fungi, as indexed by conidial production, were never greater (and sometimes less) on litter mixtures than on the component species. Total numbers, taxonomic richness and diversity of benthic invertebrates generally, and litter-feeding species in particular, were not augmented by mixing litter types. Litter mixtures appear to dilute a preferred substrate with patches of a less preferred substrate. Our results provide only weak support for the contention that combining two litter types leads to acceleration of decomposition rates. Handling editor: K. Martens     

Dominant species identity regulates invasibility of old-field plant communities

Dominant species are known to exert strong influence over community dynamics, although little work has addressed how they affect invasibility. In this study, we examined whether dominant species identity and abundance affected invasibility of old-field plant communities. To quantify invasibility, we added seeds of 19 plant species into plots dominated by one of four different herbaceous perennial species ( Andropogon virginicus , Bromus inermis , Centaurea maculosa , or Solidago canadensis ) . We found that, independent of species richness and abiotic variables, plots dominated by Andropogon were the least invasible, while Bromus and Centaurea plots had the highest invasibility. We examined several potential mechanisms by which these dominant species might influence invasibility, and found invasion to increase with decreasing litter biomass and increasing community species richness. The abundance of the dominant species was not a significant predictor of invasion. These results indicate that dominant species identity plays an important role in determining invasibility of plant communities, though exact mechanisms underlying these effects still need to be explored.     

Consistent effects of consumer species loss across different habitats

Our knowledge of the effects of consumer species loss on ecosystem functioning is limited by a paucity of manipulative field studies, particularly those that incorporate inter‐trophic effects. Further, given the ongoing transformation of natural habitats by anthropogenic activities, studies should assess the relative importance of biodiversity for ecosystem processes across different environmental contexts by including multiple habitat types. We tested the context‐dependency of the effects of consumer species loss by conducting a 15‐month field experiment in two habitats (mussel beds and rock pools) on a temperate rocky shore, focussing on the responses of algal assemblages following the single and combined removals of key gastropod grazers (Patella vulgata, P. ulyssiponensis, Littorina littorea and Gibbula umbilicalis). In both habitats, the removal of limpets resulted in a larger increase in macroalgal richness than that of either L. littorea or G. umbilicalis. Further, by the end of the study, macroalgal cover and richness were greater following the removal of multiple grazer species compared to single species removals. Despite substantial differences in physical properties and the structure of benthic assemblages between mussel beds and rock pools, the effects of grazer loss on macroalgal cover, richness, evenness and assemblage structure were remarkably consistent across both habitats. There was, however, a transient habitat‐dependent effect of grazer removal on macroalgal assemblage structure that emerged after three months, which was replaced by non‐interactive effects of grazer removal and habitat after 15 months. This study shows that the effects of the loss of key consumers may transcend large abiotic and biotic differences between habitats in rocky intertidal systems. While it is clear that consumer diversity is a primary driver of ecosystem functioning, determining its relative importance across multiple contexts is necessary to understand the consequences of consumer species loss against a background of environmental change. Synthesis The roles of species may vary with environmental context, making it difficult to predict how biodiversity loss affects ecosystem functioning across multiple habitats. We tested how natural algal assemblages in two distinct intertidal habitats responded to the removal of different combinations of key consumer species. Despite an initial habitat‐dependent effect of consumer loss, habitat type did not modify the longer‐term responses of algal assemblages to either the identity or number of consumer species removed. Our findings show that, in certain systems, consumer diversity remains a primary driver of ecosystem functioning across widely different environmental contexts.     

Effects of Macrobrachium hainanense predation on benthic community functioning in tropical Asian streams

1. Macrobrachium hainanense is a large predatory palaemonid shrimp, present at high densities in pools of low‐order forested streams in Hong Kong. The present study investigated the impacts of M. hainanense on benthic community structure and functions in pools of two streams: Tai Po Kau Forest Stream and Tai Shing Stream. 2. Repeated whole‐pool experiments involving shrimp density manipulations (removal, control and addition) were conducted in both streams between October 2000 and April 2002, and included a wet (May to September) and two dry (October to April) seasons. The three objectives of the study were to determine the effects of M. hainanense predation on benthic macroinvertebrate abundance and species richness, rates of leaf litter breakdown because of effects on detritivores, and periphyton standing stocks by way of an effect on herbivores. 3. Wet season results showed consistent reductions in benthos densities and species richness following heavy rainfall, irrespective of shrimp manipulation. These results suggested that spate‐induced disturbances might override biotic effects and play a dominant role in structuring benthic communities in stream pools in Hong Kong. 4. No significant, reproducible effects on any of the response variables measured in either stream were found during the dry season. Litter breakdown was reduced in the absence of shrimps during one experiment only, suggesting it might be a type I error. These results signified no effect of shrimp removal on benthic communities, or on the functional processes of litter breakdown, or on periphyton accumulation. The large scale of the experimental units (8–40 m²), refuge availability, and the presence of benthic predatory fishes that cropped excess prey made available by removal of M. hainanense, may have contributed to the lack of any effect, despite the abundance of the predatory shrimps.     

Resource–consumer diversity: testing the effects of leaf litter species diversity on stream macroinvertebrate communities

1. Understanding relationships between resource and consumer diversity is essential to predicting how changes in resource diversity might affect several trophic levels and overall ecosystem functioning. 2. We tested for the effects of leaf litter species diversity (i.e. litter mixing) on litter mass remaining and macroinvertebrate communities (taxon diversity, abundance and biomass) during breakdown in a detritus‐based headwater stream (North Carolina, U.S.A.). We used full‐factorial analyses of single‐ and mixed‐species litter from dominant riparian tree species with distinct leaf chemistries [red maple (Acer rubrum), tulip poplar (Liriodendron tulipifera), chestnut oak (Quercus prinus) and rhododendron (Rhododendron maximum)] to test for additivity (single‐species litter presence/absence effects) and non‐additivity (emergent effects of litter species interactions). 3. Significant non‐additive effects of litter mixing on litter mass remaining were explained by species composition, but not richness, and litter‐mixing effects were variable throughout breakdown. Specifically, small differences in observed versus expected litter mass remaining were measured on day 14; whereas observed litter mass remaining in mixed‐species leaf packs was significantly higher on day 70 and lower on day 118 than expected from data for single‐species leaf packs. 4. Litter mixing had non‐additive effects on macroinvertebrate community structure. The number of species in litter mixtures (two to four), but not litter species composition, was a significant predictor of the dominance of particular macroinvertebrates (i.e. indicator taxa) within mixed‐species packs. 5. In addition, the presence/absence of high‐ (L. tulipifera) and low‐quality (R. maximum) litter had additive effects on macroinvertebrate taxon richness, abundance and biomass. The presence of L. tulipifera litter had both positive (synergistic) and negative (antagonistic) effects on invertebrate taxon richness, that varied during breakdown but were not related to litter chemistry. In contrast, the presence/absence of L. tulipifera had a negative relationship with total macroinvertebrate biomass (due to low leaf mass remaining when L. tulipifera was present and higher condensed and hydrolysable tannins associated with leaf packs lacking L. tulipifera). Macroinvertebrate abundance was consistently lower when R. maximum was present, which was partially explained by litter chemistry [e.g., high concentrations of lignin, condensed tannins, hydrolysable tannins and total phenolics and high carbon to nutrient (N and P) ratios]. 6. The bottom‐up effects of litter species diversity on stream macroinvertebrates and litter breakdown are different, which suggests that structural attributes of macroinvertebrate communities may only partially explain the effects of litter‐mixing on organic matter processing in streams. In addition, stream macroinvertebrates colonising decomposing litter are influenced by resource diversity as well as resource availability. Broad‐scale shifts in riparian tree species composition will alter litter inputs to streams, and our results suggest that changes in the diversity and availability of terrestrial litter may alter stream food webs and organic matter processing.