Tree species identity determines wood decomposition via microclimatic effects

Empirical evidence suggests that the rich set of ecosystem functions and nature's contributions to people provided by forests depends on tree diversity. Biodiversity–ecosystem functioning research revealed that not only species richness per se but also other facets of tree diversity, such as tree identity, have to be considered to understand the underlying mechanisms. One important ecosystem function in forests is the decomposition of deadwood that plays a vital role in carbon and nutrient cycling and is assumed to be determined by above‐ and belowground interactions. However, the actual influence of tree diversity on wood decay in forests remains inconclusive. Recent studies suggest an important role of microclimate and advocate a systematical consideration of small‐scale environmental conditions. We studied the influence of tree species richness, tree species identity, and microclimatic conditions on wood decomposition in a 12‐year‐old tree diversity experiment in Germany, containing six native species within a tree species richness gradient. We assessed wood mass loss, soil microbial properties, and soil surface temperature in high temporal resolution. Our study shows a significant influence of tree species identity on all three variables. The presence of Scots pine strongly increased wood mass loss, while the presence of Norway spruce decreased it. This could be attributed to structural differences in the litter layer that were modifying the capability of plots to hold the soil surface temperature at night, consequently leading to enhanced decomposition rates in plots with higher nighttime surface temperatures. Therefore, our study confirmed the critical role of microclimate for wood decomposition in forests and showed that soil microbial properties alone were not sufficient to predict wood decay. We conclude that tree diversity effects on ecosystem functions may include different biodiversity facets, such as tree identity, tree traits, and functional and structural diversity, in influencing the abiotic and biotic soil properties.     

Neighbour identity hardly affects litter-mixture effects on decomposition rates of New Zealand forest species

The mass loss of litter mixtures is often different than expected based on the mass loss of the component species. We investigated if the identity of neighbour species affects these litter-mixing effects. To achieve this, we compared decomposition rates in monoculture and in all possible two-species combinations of eight tree species, widely differing in litter chemistry, set out in two contrasting New Zealand forest types. Litter from the mixed-species litter bags was separated into its component species, which allowed us to quantify the importance of litter-mixing effects and neighbour identity, relative to the effects of species identity, litter chemistry and litter incubation environment. Controlling factors on litter decomposition rate decreased in importance in the order: species identity (litter quality) >> forest type >> neighbour species. Species identity had the strongest influence on decomposition rate. Interspecific differences in initial litter lignin concentration explained a large proportion of the interspecific differences in litter decomposition rate. Litter mass loss was higher and litter-mixture effects were stronger on the younger, more fertile alluvial soils than on the older, less-fertile marine terrace soils. Litter-mixture effects only shifted percentage mass loss within the range of 1.5%. There was no evidence that certain litter mixtures consistently showed interactive effects. Contrary to common theory, adding a relatively fast-decomposing species generally slowed down the decomposition of the slower decomposing species in the mixture. This study shows that: (1) species identity, litter chemistry and forest type are quantitatively the most important drivers of litter decomposition in a New Zealand rain forest; (2) litter-mixture effects—although statistically significant—are far less important and hardly depend on the identity and the chemical characteristics of the neighbour species; (3) additive effects predominate in this ecosystem, so that mass dynamics of the mixtures can be predicted from the monocultures.     

Wood decomposition is more strongly controlled by temperature than by tree species and decomposer diversity in highly species rich subtropical forests

While the number of studies on the role of biodiversity on ecosystem functioning is steadily increasing, a key component of biogeochemical cycling in forests, dead wood decay, has been largely neglected. It remains widely unknown whether and how dead wood decay is affected by diversity loss in forests. We studied the hierarchical effects of tree species diversity on wood decay rates in a subtropical forest landscape in southeast China via its influence on fungal OTU richness and invertebrate diversity using piecewise structural equation models. The experiment was conducted in natural forest plots that span a wide gradient of tree species diversity embedded in a heterogeneous topography. To account for interactions between macro‐invertebrates and fungi, that potentially modify the influence of tree biodiversity and climate on dead wood decay, we compared a macro‐invertebrate exclusion treatment with a control treatment that allowed access to all types of decomposers. Diversity effects of trees on wood decay rates were mostly negative and mediated by the diversity of macro‐invertebrates. However, the effects of tree species diversity or fungal OTU richness and macro‐invertebrate diversity on wood decay rates were comparatively weak. Temperature affected decay rates positively and had the strongest influence in all treatments. While the exclusion of macro‐invertebrates did not lead to a reduction of wood decay rates, our results suggest that they may however have a mediating role in the process. In the presence of invertebrates the predictability of wood decay rates was higher and we observed a tendency of a stronger temperature control. Our results suggest that there is evidence for diversity effects on wood decomposition, but the temperature control is still more important. Thus, an increase in mean annual temperature will increase carbon and nutrient turnover through wood decomposition in subtropical forest irrespective of biotic composition.     

Multi-scale spatial patterns of three seagrass species with different growth dynamics

Knowledge of landscape spatial patterns of seagrasses and their rates of loss and natural colonization is critical for understanding the ecology of this group of submerged aquatic plants. Seagrasses form extensive meadows that occupy sheltered coastal seas of the world. In this paper, we examine the multi-scale variability of three seagrass species over a large near-shore region (42 km²) in Western Australia. Geostatistical non-parametric methods were used to explore spatial variation in presence of Amphibolis griffithii , Posidonia coriacea and P. sinuosa , and to identify the spatial scales at which distinct patterns in the species distributions occur: <50, 50–610 and >610 m. Each species showed unique variance structure across local (<50 and 50–610 m) and regional scales (>610 m), suggesting differences in species biology, environmental requirements, inter-species interactions, and their ability to modify their environment. These observations reflect that 1) seagrass landscapes are created by processes that independently act on each seagrass species at different spatial scales; 2) the species' distributions differ in their hydrodynamic forcing, and; 3) seagrass species distributions reflect colonization history such that related species are separated in space because they have different places in the successional sequence. This cross-scale study demonstrates that shoot studies only partly address the spatial structure of seagrass landscapes and further large-scale spatially-explicit research is required before we can interpret the driving processes.     

Decomposition and nutrient dynamics in mixed litter of Mediterranean species

In the last decade a great research effort addressed the effects of litter diversity on ecosystem functions, reporting both synergistic and antagonistic effects for decomposition dynamics. Four coexisting Mediterranean species, representing a range of litter quality, were used to arrange litter mixtures at three diversity levels for a litterbag decomposition experiment. Species identity appeared as the major determinant for litter mass loss (Coronilla emerus～Hedera helix>Festuca drymeia>Quercus ilex) and nutrient release, with rates for all leaf litter types following the sequence K>N>Mg≥Ca>>Fe. Additive diversity effects were prevalent pooling together all data but also for nutrients separately. Antagonistic interactions were more common than synergistic in the cases of mass loss, N and Ca contents, but not for K, Mg and Fe dynamics. The number of species in the litterbag significantly affected the outcome of non-additive interactions, which were mostly antagonistic for two-species mixtures, and synergistic for the combined 4 species. Litter quality appears to be the most important factor affecting mass loss and nutrient dynamics, while litter diversity, influencing the rates of these processes, plays an important role in reducing their variability, thus suggesting a greater stability of ecosystems properties in presence of mixed litter.     

Aquatic hyphomycete diversity and identity affect leaf litter decomposition in microcosms

We conducted a microcosm experiment with monocultures and all possible combinations of four aquatic hyphomycete species, Articulospora tetracladia, Flagellospora curta, Geniculospora grandis and Heliscus submersus, to examine the potential effects of species richness on three functional aspects: leaf litter decomposition (leaf mass loss), fungal production (ergosterol buildup) and reproductive effort (released spores). Both species richness and identity significantly affected fungal biomass and conidial production (number and biomass of released spores), whereas only species identity had a significant effect on leaf mass loss. In mixed cultures, all measures of fungal functions were greater than expected from the weighted performances of participating species in monoculture. Mixed cultures outperformed the most active monoculture for biomass accumulation but not for leaf mass loss and conidial production. The three examined aspects of aquatic hyphomycete activity tended to increase with species richness, and a complementary effect was unequivocally demonstrated for fungal biomass. Our results also suggest that specific traits of certain species may have a greater influence on ecosystem functioning than species number.     

Effects of species evenness and dominant species identity on multiple ecosystem functions in model grassland communities

Ecosystems provide multiple services upon which humans depend. Understanding the drivers of the ecosystem functions that support these services is therefore important. Much research has investigated how species richness influences functioning, but we lack knowledge of how other community attributes affect ecosystem functioning. Species evenness, species spatial arrangement, and the identity of dominant species are three attributes that could affect ecosystem functioning, by altering the relative abundance of functional traits and the probability of synergistic species interactions such as facilitation and complementary resource use. We tested the effect of these three community attributes and their interactions on ecosystem functions over a growing season, using model grassland communities consisting of three plant species from three functional groups: a grass (Anthoxanthum odoratum), a forb (Plantago lanceolata), and a N-fixing forb (Lotus corniculatus). We measured multiple ecosystem functions that support ecosystem services, including ecosystem gas exchange, water retention, C and N loss in leachates, and plant biomass production. Species evenness and dominant species identity strongly influenced the ecosystem functions measured, but spatial arrangement had few effects. By the end of the growing season, evenness consistently enhanced ecosystem functioning and this effect occurred regardless of dominant species identity. The identity of the dominant species under which the highest level of functioning was attained varied across the growing season. Spatial arrangement had the weakest effect on functioning, but interacted with dominant species identity to affect some functions. Our results highlight the importance of understanding the role of multiple community attributes in driving ecosystem functioning.     

Plant genotype identity and diversity interact with mesograzer species diversity to influence detrital consumption in eelgrass meadows

Seagrass meadows are among the world's most productive ecosystems, and as in many other systems, genetic diversity is correlated with increased production. However, only a small fraction of seagrass production is directly consumed, and instead much of the secondary production is fueled by the detrital food web. Here, we study the roles of plant genetic diversity and grazer species diversity on detrital consumption in California eelgrass Zostera marina meadows. We used three common mesograzers—an amphipod, Ampithoe lacertosa, an isopod, Idotea resecata, and a polychaete, Platynereis bicanaliculata. Each grazer consumed eelgrass detritus at rates greater than live eelgrass or macroalgae. This detrital consumption, however, was not spread evenly over leaves shed from different eelgrass clones. Palatability and consumption varied because of genotype specific differences in leaf texture, secondary metabolites (phenolics), and nutritional quality (nitrogen). Further, detritus derived from some eelgrass genotypes was palatable to all grazers, while detritus from other genotypes was preferentially consumed by only one grazer species. Under monospecific grazer assemblages, plant genetic identity but not diversity influenced detritus consumption. However, more realistic, diverse mesoconsumer communities combined with high plant‐detrital genotypic diversity resulted in greater consumption and grazer survival. These results provide a mechanism for field observations of increased mesograzer density and diversity in genetically diverse seagrass assemblages and offer a potential explanation for variation in results of resource diversity– detrital processing experiments in the literature, which often exclude macroinvertebrate taxa. More broadly, our findings support the emerging principle that biodiversity effects are strongest when diversity in both consumer and resource taxa are present.     

Simulating species loss following perturbation: assessing the effects on process rates

We removed stream-living macroinvertebrate shredder species in the sequences in which they are predicted to disappear, in response to two common types of anthropogenic disturbances: acidification and organic pollution, and analysed the effects on leaf breakdown rates. The experiment was performed in field microcosms using three shredder species. Species identity significantly affected leaf breakdown rates, while species richness per se was non-significant. The simulated sequential species loss showed large effects on leaf breakdown rates, with observed rates being significantly higher than expected from single-species treatments in two, out of four, two-species, and in all four three-species treatments. The invertebrates used in this study were taxonomically distinct (Insecta: Plecoptera and Trichoptera; Crustacea: Amphipoda), and of different sizes, hence a high degree of complementarity was probably present. A method to study the effects of species loss, characteristic of perturbation type, could be more useful than a random approach when investigating the impact of perturbation. Our results may have general applicability for investigations on the effects of diversity loss on ecosystem functioning in any ecosystem exposed to human perturbations, given that the order of extinction is known or can easily be assessed.     

The short-term influence of herbivory near patch reefs varies between seagrass species

The coexistence of multiple species within a trophic level can be regulated by consumer preferences and nutrient supply, but the influence of these factors on the co-occurrence of seagrass species is not well understood. We examined the biomass and density responses of two seagrass species in the Florida Keys Reef Tract to grazing pressure near patch reefs, and evaluated how nutrient enrichment impacted herbivory dynamics. We transplanted Halodule wrightii (shoalgrass) sprigs into caged and uncaged plots in a Thalassia testudinum (turtlegrass) bed near a patch reef. Nutrients (N and P) were added to half of the experimental plots. We recorded changes in seagrass shoot density, and after three months, we measured above- and belowground biomass and tissue nutrient content of both species. Herbivory immediately and strongly impacted H. wrightii. Within six days of transplantation, herbivory reduced the density of uncaged H. wrightii by over 80%, resulting in a decrease in above- and belowground biomass of nearly an order of magnitude. T. testudinum shoot density and belowground biomass were not affected by herbivory, but aboveground biomass and leaf surface area were higher within cages, suggesting that although herbivory influenced both seagrass species, T. testudinum was more resistant to herbivory pressure than H. wrightii. Nutrient addition did not alter herbivory rates or the biomass of either species over the short-term duration of this study. In both species, nutrient addition had little effect on the tissue nutrient content of seagrass leaves, and N:P was near the 30:1 threshold that suggested a balance between N and P. The different impacts of grazing on these two seagrass species suggest that herbivory may be an important regulator of the distribution of multiple seagrass species near herbivore refuges like patch reefs in the Caribbean.     

Evaluation of aboveground and belowground biomass recovery in physically disturbed seagrass beds

Several studies addressed aboveground biomass recovery in tropical and subtropical seagrass systems following physical disturbance. However, there are few studies documenting belowground biomass recovery despite the important functional and ecological role of roots and rhizomes for seagrass ecosystems. In this study, we compared the recovery of biomass (g dry weight m(-2)) as well as the biomass recovery rates in ten severely disturbed multi-species seagrass meadows, after the sediments were excavated and the seagrasses removed. Three sites were located in the tropics (Puerto Rico) and seven in the subtropics (Florida Keys), and all were originally dominated by Thalassia testudinum. Total aboveground biomass reached reference values at four out of ten sites studied, two in the Florida Keys and two in Puerto Rico. Total belowground biomass was lower at the disturbed locations compared to the references at all sites, apart from two sites in the Florida Keys where the compensatory effect of opportunistic species (Syringodium filiforme and Halodule wrightii) was observed. The results revealed large variation among sites in aboveground and belowground biomass for all species, with higher aboveground recovery than belowground for T. testudinum. Recovery rates for T. testudinum were highly variable across sites, but a general trend of faster aboveground than belowground recovery was observed. Equal rates between aboveground and belowground biomass were found for opportunistic species at several sites in the Florida Keys. These results indicate the importance of belowground biomass when assessing seagrass recovery and suggest that the appropriate metric to assess seagrass recovery should address belowground biomass as well as aboveground biomass in order to evaluate the full recovery of ecological services and functions performed by seagrasses. We point out regional differences in species composition and species shifts following severe disturbance events and discuss ecological implications of gap dynamics in multi-species seagrass meadows.     

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.     

Interacting effects of leaf litter species and macrofauna on decomposition in different litter environments

The leaf litter environment (single species versus mixed species), and interactions between litter diversity and macrofauna are thought to be important in influencing decomposition rates. However, the role of soil macrofauna in the breakdown of different species of leaf litter is poorly understood. In this study we examine the multiple biotic controls of decomposition – litter quality, soil macrofauna and litter environment and their interactions. The influence of soil macrofauna and litter environment on the decomposition of six deciduous tree species (Fraxinus excelsior L., Acer pseudoplatanus L., Acer campestre L., Corylus avellana L., Quercus robur L., Fagus sylvatica L.) was investigated in a temperate forest, Wytham Woods, Southern England. We used litterbags that selectively excluded macrofauna to assess the relative importance of macrofauna versus microbial, micro and mesofauna decomposition, and placed single species bags in either conspecific single species or mixed species litter environments. The study was designed to separate plant species composition effects on litter decomposition rates, allowing us to evaluate whether mixed species litter environments affect decomposition rates compared to single species litter environments, and if so whether the effects vary among litter species, over time, and with regard to the presence of soil macrofauna. All species had faster rates of decomposition when macrofauna were present, with 22–41% of the total mass loss attributed to macrofauna. Macrofauna were most important for easily decomposable species as soon as the leaves were placed on the ground, but were most important for recalcitrant species after nine months in the field. The mass loss rates did not differ between mixed and single species litter environments, indicating that observed differences between single species and mixed species litterbags in previous field studies are due to the direct contact of neighbouring species inside the litterbag rather than the litter environment in which they are placed.     

Nonhost species reduce parasite infection in a focal host species within experimental fish communities

The dilution effect describes the negative association between host biodiversity and the risk of infectious disease. Tests designed to understand the relative roles of host species richness, host species identity, and rates of exposure within experimental host communities would help resolve ongoing contention regarding the importance and generality of dilution effects. We exposed fathead minnows to infective larvae of the trematode, Ornithodiplostomum ptychocheilus in minnow‐only containers and in mixed containers that held 1–3 other species of fish. Parasite infection was estimated as the number of encysted worms (i.e., brainworms) present in minnows following exposure. The results of exposure trials showed that nonminnow fish species were incompatible with O. ptychocheilus larvae. There was no reduction in mean brainworm counts in minnows in mixed containers with brook sticklebacks or longnose dace. In contrast, brainworm counts in minnows declined by 51% and 27% in mesocosms and aquaria, respectively, when they co‐occurred with emerald shiners. Dilution within minnow + shiner containers may arise from shiner‐induced alterations in minnow or parasite behaviors that reduced encounter rates between minnows and parasite larvae. Alternatively, shiners may act as parasite sinks for parasite larvae. These results highlight the role of host species identity in the dilution effect. Our results also emphasize the complex and idiosyncratic effects of host community composition on rates of parasite infection within contemporary host communities that contain combinations of introduced and native species.     

The influence of interspecific interactions on species range expansion rates

Ongoing and predicted global change makes understanding and predicting species' range shifts an urgent scientific priority. Here, we provide a synthetic perspective on the so far poorly understood effects of interspecific interactions on range expansion rates. We present theoretical foundations for how interspecific interactions may modulate range expansion rates, consider examples from empirical studies of biological invasions and natural range expansions as well as process‐based simulations, and discuss how interspecific interactions can be more broadly represented in process‐based, spatiotemporally explicit range forecasts. Theory tells us that interspecific interactions affect expansion rates via alteration of local population growth rates and spatial displacement rates, but also via effects on other demographic parameters. The best empirical evidence for interspecific effects on expansion rates comes from studies of biological invasions. Notably, invasion studies indicate that competitive dominance and release from specialized enemies can enhance expansion rates. Studies of natural range expansions especially point to the potential for competition from resident species to reduce expansion rates. Overall, it is clear that interspecific interactions may have important consequences for range dynamics, but also that their effects have received too little attention to robustly generalize on their importance. We then discuss how interspecific interactions effects can be more widely incorporated in dynamic modeling of range expansions. Importantly, models must describe spatiotemporal variation in both local population dynamics and dispersal. Finally, we derive the following guidelines for when it is particularly important to explicitly represent interspecific interactions in dynamic range expansion forecasts: if most interacting species show correlated spatial or temporal trends in their effects on the target species, if the number of interacting species is low, and if the abundance of one or more strongly interacting species is not closely linked to the abundance of the target species.     

Effect of nitrogen addition,form and clipping on competitive interactions between grassland species

Aims Although the effects of N addition on plant biomass are well understood, we know a lot less about the importance of N form even though some studies have shown different impacts from reduced and oxidized forms of N. Furthermore, responses to grazing are likely to interact with the response to N addition. This experiment investigates the interactive effects of N addition and form with clipping on competition between three grassland species.Methods The three species (Anthoxanthum odoratum L., Plantago lanceolata L. and Prunella vulgaris L.) were grown alone and in combination with factorial additions of deionized water, sodium nitrate and ammonium chloride, and a clipping treatment. Above- and belowground biomass was harvested after 4 months.Important findings In monocultures, the results show increases in biomass with N addition, but clipping resulted in fewer changes with species displaying varying degrees of growth compensation. A. odoratum was the strongest competitor when grown with other species. In monocultures without clipping, N form was not important, but in the presence of clipping and in different species combinations, N form became important. Significant two- and three-way interactive effects were observed showing that complex interactions exist between N addition, clipping and species identity. The results have important implications when considering the effects of N deposition.     

Introduced species in seagrass ecosystems: Status and concerns

A literature review revealed that at least 56 non-native species, primarily invertebrates and seaweeds, have been introduced to seagrass beds, largely through shipping/boating activities and aquaculture. Four seagrass species also have been introduced. The introductions of the seaweeds Caulerpa taxifolia, C. racemosa v. cylindracea, Codium fragile ssp. tomentosoides, Sargassum muticum, the Asian mussel, Musculista senhousia, and the seagrass, Zostera japonica, are the best-known examples in seagrass beds. The ecological effects on seagrasses and associated communities have been examined for slightly less than half of the introduced species, which have predominantly negative effects. There is a paucity of experimental data for ecological effects, particularly for seagrass community structure and function. The exception to this finding is the introduction of the seagrass Z. japonica with oyster aquaculture to native eelgrass beds on the Pacific coasts of Canada and the USA. Recent experiments in several different seagrass ecosystems confirmed that disturbance contributes to the invasibility of seagrass beds. More definitive studies are required to elucidate the relative effects of nutrient pollution and introduced species in causing seagrass decline, particularly where reduced herbivory and boating activity also covary. Seagrass beds often are subject to multiple introduced species, but their cumulative effect has been virtually unstudied. The potential for compounded negative effects merits serious attention. Heightened attention to the issue of introduced species in seagrass beds is called for given the evidence that introduced species can contribute to seagrass decline, to biodiversity changes that could affect seagrass ecosystem functions, and that they can compromise seagrass restoration. Comprehensive surveys in seagrass beds, complemented by more stringent experimental and mensurative sampling designs, are needed. In the interim, conserving seagrass density and bed size can offer resistance to introduced species. Managing to prevent the introductions, including restricting transplantations of non-native biota during seagrass restorations, is likely to bear positive benefits for seagrass ecosystems.     

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.     

Scale and species numbers

One of the main tasks confronting community ecologists is to explain why a particular site harbours a certain number of species. The site might range from a drop of water to the whole Earth, and the species might be drawn from a very restricted taxon or include all living organisms. The common problem, however, is to understand the relative importance of speciation and extinction and, more locally, of immigration and loss. Speciation is the ultimate motor driving biodiversity and ecologists need to know the factors influencing rates of speciation, and whether there is a feedback, positive or negative, between species numbers and the generation of new taxa. However, the relative importance of speciation and other factors determining species numbers varies crucially across different scales of enquiry. Here, we explore some of these issues as we move from a macro- to microscale perspective, focusing on a limited number of studies that we believe make important advances in the field.