Placing biodiversity and ecosystem functioning in context: environmental perturbations and the effects of species richness in a stream field experiment

Greater biodiversity is often associated with increased ecosystem process rates, and is expected to enhance the stability of ecosystem functioning under abiotic stress. However, these relationships might themselves be altered by environmental factors, complicating prediction of the effects of species loss in ecosystems subjected to abiotic stress. In boreal streams, we investigated effects of biodiversity and two abiotic perturbations on three related indices of ecosystem functioning: leaf decomposition, detritivore leaf processing efficiency (LPE) and detritivore growth. Replicate field enclosures containing leaves and detritivore assemblages were exposed to liming and nutrient enrichment, raising pH and nutrient levels. Both treatments constitute perturbations for our naturally acidic and nutrient-poor streams. We also varied detritivore species richness and density. The effects of the abiotic and diversity manipulations were similar in magnitude, but whereas leaf decomposition increased by 18% and 8% following liming and nutrient enrichment, respectively, increased detritivore richness reduced leaf decomposition (6%), detritivore LPE (19%) and detritivore growth (12%). The detritivore richness effect on growth was associated with negative trait-independent complementarity, indicating interspecific interference competition. These interactions were apparently alleviated in both enriched and limed enclosures, as trait-independent complementarity became less negative. LPE increased with detritivore density in the monocultures, indicating benefits of intra-specific aggregation that outweighed the costs of intra-specific competition, and dilution of these benefits probably contributed to lowered leaf decomposition in the species mixtures. Finally, the effects of liming were reduced in most species mixtures relative to the monocultures. These results demonstrate how environmental changes might regulate the consequences of species loss for functioning in anthropogenically perturbed ecosystems, and highlight potential influences of biodiversity on functional stability. Additionally, the negative effects of richness and positive effects of density in our field study were opposite to previous laboratory observations, further illustrating the importance of environmental context for biodiversity–ecosystem functioning relationships.     

How biological diversity influences ecosystem function: a test with a tropical stream detritivore guild

We investigated the relationship between diversity and ecosystem function, which is controversial and has rarely been examined for consumer assemblages, for the process of leaf breakdown by the shredder guild in a tropical stream. We manipulated species richness, evenness and identity of four macroinvertebrate shredder species (three caddisflies and one mayfly) in microcosms and tested their effect on leaf breakdown rates measured as leaf mass loss per capita and per milligram of animal. Species richness, evenness and species identity all affected leaf breakdown rates. Breakdown rates tended to increase with higher richness, but only for the three caddisflies, probably through a release of intraspecific interference, although other mechanisms such as niche complementarity or facilitation cannot be discarded. Leaf breakdown by the caddisflies was reduced in the presence of the mayfly, possibly because of its mode of movement by swimming instead of crawling and its similarity to some predators that are common in leaf litter. Species identity was more important than species richness in determining leaf breakdown rates, indicating that some species within the shredder guild are not redundant, and suggesting important consequences of particular species loss for the functioning of the ecosystem.     

Dominant species can produce a negative relationship between species diversity and ecosystem function

Several studies have reported a positive relationship between species richness and ecosystem functioning. However, if much of a particular ecosystem function is performed by one species (i.e. a functionally dominant species) and this species is also a competitive dominant that excludes other taxa from a habitat, then it is possible to obtain a negative relationship between richness and ecosystem functioning. Results of a leaf pack breakdown experiment in a small stream suggested that the caddisfly Pycnopsyche gentilis , a common detritivorous insect in North American headwater streams, was both a functional and competitive dominant. In a second experiment we compared the effect of Pycnopsyche on leaf breakdown to that of other detritivore taxa by enclosing them with leaf packs in a section of headwater stream in which they were uncommon ( Pycnopsyche transplant experiment). Final leaf pack mass was significantly lower in the Pycnopsyche enclosure treatment; leaves exposed to a greater diversity of detritivores displayed little reduction in leaf mass. These results demonstrated that Pycnopsyche was a functionally dominant detritivore. In a third experiment ( Pycnopsyche density experiment) we found that Pycnopsyche was also a competitively dominant species. Leaf packs and large Pycnopsyche were placed in enclosures that were permeable to the majority of other detritivores but not Pycnopsyche . Leaf mass lost increased with increasing Pycnopsyche density. Leaf packs exposed to Pycnopsyche , however, contained fewer detritivore taxa which suggested that Pycnopsyche was also a competitive dominant. There was a negative relationship between three measures of diversity and leaf litter breakdown in the Pycnopsyche density experiment. Experiments conducted in natural communities that incorporate important species interactions may produce diversity-ecosystem function relationships other than the positive ones that are commonly reported.     

Ecosystem functioning in stream assemblages from different regions: contrasting responses to variation in detritivore richness, evenness and density

1. The diversity of species traits in a biological assemblage varies not only with species richness, but also with species evenness and organism density, which together influence the concentration of traits within functional guilds. Potential trait diversity at local scales is also constrained by the regional species pool. Implications of such variation for spatio-temporal variability in biodiversity-ecosystem functioning relationships are likely to be complex, but are poorly understood. 2. In microcosm experiments conducted at laboratories in Sweden, Ireland and Romania, we investigated effects of species richness, evenness and density of stream-living detritivores on two related processes: detritivore leaf-processing efficiency (LPE) and growth. Assemblage composition varied among laboratories: one taxonomic order (Plecoptera) was studied in Sweden, whereas two orders, encompassing wider trait variation, were studied in Romania (Trichoptera and Plecoptera) and Ireland (Trichoptera and Isopoda). 3. Relationships between density and both LPE and growth ranged from negative to positive across the study species, highlighting the potential for density-dependent variation in process rates to alter ecosystem functioning, but indicating that such effects depend on species identity. 4. LPE varied with species diversity in the two more heterogeneous assemblages, but whereas LPE in the Romanian study was generally enhanced as richness increased, LPE in the Irish study increased only in less-even polycultures dominated by particular species. Transgressive overyielding was detected in the Irish experiment, indicating complementary resource use and/or facilitation (complementarity). These mechanisms could not be distinguished from the selection effect in the Romanian study. 5. Growth was elevated in Romanian species mixtures, reflecting positive complementarity, but lower than expected growth in some Swedish mixtures was associated with negative complementarity, indicating interspecific interference competition. 6. Our results emphasize the potential importance of detritivore diversity for stream ecosystem functioning, but both the effects of diversity on the studied processes, and the mechanisms underlying those effects, were specific to each assemblage and process. Such variability highlights challenges in generalizing impacts of diversity change for functional integrity in streams and other ecosystems in which the occurrence of important species traits fluctuates over relatively small spatio-temporal scales.     

Biodiversity and ecosystem functioning in a species-poor guild: a test using tropical stream detritivores

The relationship between biodiversity and ecosystem functioning (B–EF) was investigated by examining top-down effects of aquatic detritivore diversity on the functional process of leaf-litter breakdown. This study was undertaken in tropical Hong Kong where the stream detritivore guild is depauperate and loss of one or a few species might have strong effects on processes. Effects of detritivore richness and composition were investigated by comparing feeding rates of three species of detritivores with their two- and three-species mixtures in laboratory trials. The detritivores were a caddisfly larva (Anisocentropus maculatus: Calamoceratidae), a snail (Brotia hainanensis: Pachychilidae) and a shrimp (Caridina cantonensis: Atyidae). Liquidambar formosana (Hamamelidaceae) litter was provided as food. All three detritivore species had positive non-additive effects on litter processing. Per capita and mass-specific feeding rates of each species were faster in mixtures than when they were alone, although the non-additive effects of shrimps and snails were larger than those attributable to caddisflies, and thus, litter processing was strongly influenced by the composition of detritivore mixtures. The compositional effect appears to be evidence of facilitation indicating a lack of functional redundancy amongst these detritivores, probably due to their evolutionary distinctness, implying that extinction consequences in this species-poor guild will depend on the identity of the species lost.     

Riparian plant species loss alters trophic dynamics in detritus-based stream ecosystems

Riparian vegetation is closely connected to stream food webs through input of leaf detritus as a primary energy supply, and therefore, any alteration of plant diversity may influence aquatic ecosystem functioning. We measured leaf litter breakdown rate and associated biological parameters in mesh bags in eight headwater streams bordered either with mixed deciduous forest or with beech forest. The variety of leaf litter types in mixed forest results in higher food quality for large-particle invertebrate detritivores (‘shredders’) than in beech forest, which is dominated by a single leaf species of low quality. Breakdown rate of low quality (oak) leaf litter in coarse mesh bags was lower in beech forest streams than in mixed forest streams, a consequence of lower shredder biomass. In contrast, high quality (alder) leaf litter broke down at similar rates in both stream categories as a result of similar shredder biomass in coarse mesh bags. Microbial breakdown rate of oak and alder leaves, determined in fine mesh bags, did not differ between the stream categories. We found however aquatic hyphomycete species richness on leaf litter to positively co-vary with riparian plant species richness. Fungal species richness may enhance leaf litter breakdown rate through positive effects on resource quality for shredders. A feeding experiment established a positive relationship between fungal species richness per se and leaf litter consumption rate by an amphipod shredder (Gammarus fossarum). Our results show therefore that plant species richness may indirectly govern ecosystem functioning through complex trophic interactions. Integrating microbial diversity and trophic dynamics would considerably improve the prediction of the consequences of species loss.     

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.     

Intra-specific leaf trait responses to species richness at two different local scales

Plant functional traits, especially leaf traits, are accepted proxies for ecosystem properties. Typically, they are measured at the species level, neglecting within-species variation. While there is extensive knowledge about functional trait changes (both within and across species) along abiotic gradients, little is known about biotic influences, in particular at local scales. Here, we used a large biodiversity-ecosystem functioning experiment in subtropical China to investigate intra-specific trait changes of 16 tree species as a response to species richness of the local neighbourhood. We hypothesized that because of positive complementarity effects, species shift their leaf traits towards a more acquisitive growth strategy, when species richness of the local neighbourhood is higher. The trait shift should be most pronounced, when a focal tree's closest neighbour is from a different species, but should still be detectable as a response to species richness of the directly surrounding tree community. Consequently, we expected that trees with a con-specific closest neighbour have the strongest response to species richness of the surrounding tree community, i.e., the steepest increase of acquisitive traits. Our results indicate that species diversity promoted reduced competition and complementarity in resource use at both spatial scales considered. In addition, the closest neighbour had considerably stronger effects than the surrounding tree community. As expected, trees with a con-specific nearest neighbour showed the strongest trait shifts. However, the predicted positive effect of local hetero-specificity disappeared at the highest diversity levels of the surrounding tree community, potentially resulting from a higher probability to meet a strong competitor in a diverse environment. Our findings show that leaf traits within the same species vary not only in response to changing abiotic conditions, but also in response to local species richness. This highlights the benefit of including within-species trait variation when analysing relationships between plant functional traits and ecosystem functions.     

Species richness matters for the quality of ecosystem services: a test using seed dispersal by frugivorous birds

The positive link between biodiversity and ecosystem functioning is a current paradigm in ecological science. However, little is known of how different attributes of species assemblages condition the quality of many services in real ecosystems affected by human impact. We explore the links between the attributes of a frugivore assemblage and the quantitative and qualitative components of its derived ecosystem service, seed dispersal, along a landscape-scale gradient of anthropogenic forest loss. Both the number and the richness of seeds being dispersed were positively related to frugivore abundance and richness. Seed dispersal quality, determined by the fine-scale spatial patterns of seed deposition, mostly depended on frugivore richness. In fact, richness was the only attribute of the frugivore assemblage affecting the probability of seed dispersal into deforested areas of the landscape. The positive relationships between frugivore richness per se (i.e. independent of frugivore abundance and composition) and all components of seed dispersal suggest the existence of functional complementarity and/or facilitation between frugivores. These links also point to the whole assemblage of frugivores as a conservation target, if we aim to preserve a complete seed dispersal service and, hence, the potential for vegetation regeneration and recovery, in human-impacted landscapes.     

Genotypic richness and dissimilarity opposingly affect ecosystem functioning

Biodiversity is an essential determinant of ecosystem functioning. Numerous studies described positive effects of diversity on the functioning of communities arising from complementary resource use and facilitation. However, high biodiversity may also increase competitive interactions, fostering antagonism and negatively affecting community performance. Using experimental bacterial communities we differentiated diversity effects based on genotypic richness and dissimilarity. We show that these diversity characteristics have opposite effects on ecosystem functioning. Genotypic dissimilarity governed complementary resource use, improving ecosystem functioning in complex resource environments. Contrastingly, genotypic richness drove allelopathic interactions, mostly reducing ecosystem functioning. The net biodiversity effect on community performance resulted from the interplay between the genetic structure of the community and resource complexity. These results demonstrate that increasing richness, without concomitantly increasing dissimilarity, can decrease ecosystem functioning in simple environments due to antagonistic interactions, an effect insufficiently considered so far in mechanistic models of the biodiversity-ecosystem functioning relationship.     

Composition of speciose leaf litter alters stream detritivore growth, feeding activity and leaf breakdown

Leaf litter derived from riparian trees can control secondary production of detritivores in forested streams. Species-rich assemblages of leaf litter reflect riparian plant species richness and represent a heterogeneous resource for stream consumers. Such variation in resource quality may alter consumer growth and thus the feedback on leaf breakdown rate via changes in feeding activity. To assess the consequences of this type of resource heterogeneity on both consumer growth and subsequent litter breakdown, we performed a laboratory experiment where we offered a leaf-shredding stream detritivore (the stonefly Tallaperla maria, Peltoperlidae) ten treatments of either single- or mixed-species leaf litter. We measured consumer growth rate, breakdown rate and feeding activity both with and without consumers for each treatment and showed that all three variables responded to speciose leaf litter. However, the number of leaf species was not responsible for these results, but leaf species composition explained the apparent non-additive effects. T. maria growth responded both positively and negatively to litter composition, and growth on mixed-litter could not always be predicted by averaging estimates of growth in single-species treatments. Furthermore, breakdown and feeding rates in mixed litter treatments could not always be predicted from estimates of single-species rates. Given that species richness and composition of senesced leaves in streams reflects riparian plant species richness, in-stream secondary production of detritivores and organic matter dynamics may be related to species loss of trees in the riparian zone. Loss of key species may be more critical to maintaining such processes than species richness per se.     

Species richness–decomposition relationships depend on species dominance

Human disturbances both decrease the number of species in ecosystems and change their relative abundances. Here we present field evidence demonstrating that shifts in species abundances can have effects on ecosystem functioning that are as great as those from shifts in species richness. We investigated spatial and temporal variability of leaf decomposition rates and community metrics of leaf‐eating invertebrates (shredders) in streams. The shredder community composition dramatically influenced the diversity–function relationship; decomposition was much higher for a given species richness at sites with high species dominance than at sites where dominance was low. Decomposition rates also markedly depended on the identity of the dominant species. Further, dominance effects on decomposition varied seasonally and the number of species required for maintaining decomposition increased with increasing evenness. These findings reveal important but less obvious aspects of the biodiversity–ecosystem functioning relationship.     

Functional diversity predicts overyielding effect of species combination on primary productivity

The effect of biodiversity on ecosystem functioning has proven variable both within and among manipulative studies. Species richness is the most commonly used measure of biodiversity in such studies, but the range of species’ functional traits (functional diversity), not the number of species per se, likely underpins a key mechanistic link between species richness and ecosystem functioning. However, the majority of experiments that have examined the effect of functional diversity have manipulated functional group richness, an approach recognised to suffer numerous limitations. Continuous measures of functional diversity avoid many of these limitations, but the relationship between continuous functional diversity and the magnitude of ecosystem processes has been largely untested. Using one vs two‐species mixtures of rock pool macroalgae as a model, we conducted a field experiment to determine the effect of a continuous measure of functional diversity (functional attribute diversity, FAD, the degree of functional differentiation based on four functional traits) on the magnitude of net primary productivity and overyielding, based upon two alternative null‐models. The total magnitude of productivity was largely determined by the identity of species present, not FAD. However, FAD proved to be a good predictor of overyielding (variation in productivity after the dominant effects of species identity had been accounted for). Furthermore, despite differences in the mean magnitude of the effect of combining species, the positive relationship between FAD and overyielding was consistent according to both additive and substitutive null‐models. Our findings imply that whilst knowledge of species’ independent contributions remains indispensable in the prediction of biotic effects on ecosystem functioning within a trophic level, continuous measures of functional diversity should be used as a supplementary tool to predict the magnitude of overyielding, thereby refining predictions.     

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.     

Detritivore stoichiometric diversity alters litter processing efficiency in a freshwater ecosystem

Many studies have estimated relationships between biodiversity and ecosystem functioning, and observed generally positive effects. Because detritus is a major food resource in stream ecosystems, decomposition of leaf litter is an important ecosystem process and many studies report the full range of positive, negative and no effects of diversity on decomposition. However, the mechanisms underlying decomposition processes in fresh water remain poorly understood. Organism body stoichiometry relates to consumption rates and tendencies, and decomposition processes of litter may therefore be affected by diversity in detritivore body stoichiometry. We predicted that the stoichiometric diversity of detritivores (differences in C: nutrient ratios among species) would increase the litter processing efficiency (litter mass loss per total capita metabolic capacity) in fresh water through complementation regarding different nutrient requirements. To test this prediction, we conducted a microcosm experiment wherein we manipulated the stoichiometric diversity of detritivores and quantified mass loss of leaf litter mixtures. We compared litter processing efficiency among litter species in each microcosm with single species detritivores, and observed detritivores with nutrient‐rich bodies tended to prefer litter with lower C: nutrient ratios over litter with higher C: nutrient ratios. Furthermore, litter processing efficiencies were significantly higher in the microcosms containing species of detritivores with both nutrient‐rich and ‐poor bodies than microcosms containing species of detritivores including only nutrient‐rich or ‐poor bodies. This might mean a higher stoichiometric diversity of detritivores increased litter processing efficiency. Our results suggest that ecological stoichiometry may improve understanding of links between biodiversity and ecosystem function in freshwater ecosystems.     

Leaf herbivory is more impacted by forest composition than by tree diversity or edge effects

Links between biodiversity and ecosystem functioning are well established. Beyond biodiversity per se, community composition can have strong effects on ecosystem functioning. Furthermore, spatial processes including edge effects, can impact the diversity-functioning relationship. These spatial processes are especially relevant within a food web context, such as the transfer of plant biomass across the food chain through herbivory. The relative importance of diversity, community composition and spatial context on herbivory pressure at the community and the species level is, however, poorly understood.To fill this gap in our understanding, we studied to what degree herbivory in temperate forest plots varies according to edge distance, tree diversity and forest composition. In contrast to the prevailing view of tree herbivory increasing at forest edges, we found that the effects of forest edge and tree diversity on leaf herbivory were masked by effects of forest composition, i.e. the specific contributions of the tree species. The strongest composition effect found was increased herbivory on Quercus robur in the presence of Fagus sylvatica.Our findings highlight that neither edge distance, tree diversity, nor the interaction affected one ecosystem function, namely herbivory, whilst tree community composition did. This warrants consideration of identity and composition effects in future studies if we are to deepen our understanding of the determinants of ecosystem functions across systems.     

Ecological stoichiometry explains larger-scale facilitation processes by shrubs on species coexistence among understory plants

Plant facilitation (positive plant–plant interactions) strongly influences biodiversity, structure, and dynamics in plant communities, and the topic has received considerable attention among ecologists. Most studies of facilitation processes by shrubs have been conducted at small spatial scales between shrubs and their neighboring species. Yet, we know little about whether facilitation processes by shrubs at a small scale (i.e., a patch scale) also work at a larger scale (i.e., a site scale) in terms of the maintenance of biodiversity. Here, we report that the facilitative effects of shrubs on plant diversity at a larger scale can be explained by changing ecological stoichiometry. The soil fertility showed unimodal shape along shrub cover gradient, suggesting that the facilitative effects of a shrub do not necessarily increase as the shrub develops. The unimodal shape of dependence of plant species richness on shrub cover probably was generated by the unimodal dependence of soil fertility on shrub cover. Soil nutrient enrichment by shrubs shifted low N:P ratios of plant communities with low levels of shrub cover to more balanced N:P ratios at intermediate levels of shrub cover. At the peak N:P ratio along the gradient in shrub cover, the maximum species richness and functional richness were observed, which was consistent with the unimodal relationship predicted by the resource balance hypothesis. Thus, our findings showed that facilitation processes by shrubs at a patch scale also work at a larger scale in terms of the maintenance of biodiversity. Because observed larger-scale facilitation processes are enhanced at some intermediate levels of shrub cover, this study offers practical insight into the need for management practices that allow some intermediate levels of grazing by livestock for optimizing the role of larger-scale facilitation processes in the maintenance of biodiversity and ecosystem functioning in arid and semi-arid rangelands.     

Viral Richness is Positively Related to Group Size,but Not Mating System,in Bats

Characterizing host traits that influence viral richness and diversification is important for understanding wildlife pathogens affecting conservation and/or human health. Behaviors that affect contact rates among hosts could be important for viral diversification because more frequent intra- and inter-specific contacts among hosts should increase the potential for viral diversification within host populations. We used published data on bats to test the contact-rate hypothesis. We predicted that species forming large conspecific groups, that share their range with more heterospecifics (i.e., sympatry), and with mating systems characterized by high contact rates (polygynandry: multi-male/multi-female), would host higher viral richness than species with small group sizes, lower sympatry, or low contact-rate mating systems (polygyny: single male/multi-female). Consistent with our hypothesis and previous research, viral richness was positively correlated with conspecific group size although the relationship plateaued at group sizes of approximately several hundred thousand bats. This pattern supports epidemiological theory that, up to a point, larger groups have higher contact rates, greater likelihood of acquiring and transmitting viruses, and ultimately greater potential for viral diversification. However, contrary to our hypothesis, there was no effect of sympatry on viral richness and no difference in viral richness between mating systems. We also found no residual effect of host phylogeny on viral richness, suggesting that closely related species do not necessarily host similar numbers of viruses. Our results support the contact-rate hypothesis that intra-specific viral transmission can enhance viral diversification within species and highlight the influence of host group size on the potential of viruses to propagate within host populations.     

Chaco forest fragmentation effects on leaf litter decomposition are not explained by changes in litter fauna

Forest fragmentation is a component of global change, with substantial impact on biodiversity and ecosystem functioning. Despite extensive evidence of forest fragmentation effects on above‐ground ecological processes, little is understood about its below‐ground effects. Abundance and richness of leaf litter fauna can be affected by forest fragmentation, and this can have cascading effects on the decomposition process. Here, we examine how fragmentation of a subtropical dry forest affects aspects of ecosystem structure and functioning, by unravel area and edge effects on leaf litter fauna and decomposition rates and testing whether changes in abundance or richness of litter fauna mediated fragment area and edge effects on litter decomposition. We incubated litterbags filled with a common substrate, at the edge and interior of 12 fragments of Chaco Serrano forest in Central Argentina, for 180 days. We found that invertebrate abundance was higher at the forest edge but independent of fragment area, whereas decomposition declined with fragment size independently of edge or interior location. According to our results, the effect of forest size on decomposition was not mediated by changes in abundance or richness of leaf litter fauna, suggesting independent changes in ecosystem structure and functioning.     

Species diversity and decomposition in laboratory aquatic systems: the role of species interactions

1. Interest in the effects of biodiversity on ecosystem processes is increasing, stimulated by the global species decline. Different hypotheses about the biodiversity‐ecosystem functioning (BEF) relationship have been put forward and various underlying mechanisms proposed for different ecosystems. 2. We investigated BEF relationships and the role of species interactions in laboratory experiments focussing on aquatic decomposition. Species richness at three different trophic levels (leaf detritus, detritus‐colonising fungi and invertebrate detritivores) was manipulated, and its effects on leaf mass loss and fungal growth were assessed in two experiments. In the first, monocultures and mixtures of reed (Phragmites australis), alder (Alnus glutinosa) and oak (Quercus cerris) leaf disks were incubated with zero, one or eight fungal species. Leaf mixtures were also incubated with combinations of three and five fungal species. In the second experiment, reed leaf disks were incubated with all eight fungal species and offered to combinations of one, two, three, four or five macroinvertebrate detritivores with different feeding modes. 3. Results from the first experiment showed that leaf mass loss was directly related to fungal mass and varied unimodally with the number of fungi, with a maximum rate attained at intermediate diversity in oak and reed and at maximum diversity in alder (the fastest decomposing leaf). 4. Mixing litter species stimulated fungal growth but interactions between species of fungi slowed down decomposition. In contrast, mixtures of macroinvertebrate detritivores reduced fungal mass and accelerated leaf decomposition. Possible explanations of the positive relationship between detritivore diversity and decomposition are a reduction in fungal dominance and a differentiation in the use of different resource patches promoted by higher fungal diversity. 5. In conclusion, the results show a general increase in decomposition rate with increasing biodiversity that is controlled by within‐ and between‐trophic level interactions, and support the hypothesis of both bottom‐up and top‐down effects of diversity on this process.