Climate dependence of the macrofaunal effect on litter decomposition-A global meta-regression analysis.

Litter decomposition by microorganisms and animals is influenced by climate and has been found to be higher in warm and wet than in cold and dry biomes. We, however, hypothesized that the macrofaunal effect on decomposition should increase with temperature and aridity since larger animals are more tolerant to aridity than smaller organisms. This hypothesis was supported by our global analysis of macrofauna exclusion studies. Macrofauna increased litter mass loss on average by 40%, twofold higher than the highest previous estimation of macrofaunal effect on decomposition. The strongest effect was found in subtropical deserts where faunal decomposition had not been considered important. Our results highlight the need to consider animal size when exploring climate dependence of faunal decomposition, and the disproportionately large role of macrofauna in regulating litter decomposition in warm drylands. This new realization is critical for understanding element cycling in the face of global warming and aridification.

Functional traits of predators and decomposer prey determine context dependency in trophic control over ecosystems.

Research Highlight: Piccoli, G. C. d. O., Antiqueira, P. A. P., Srivastava, D. S., & Romero, G. Q. (2024). Trophic cascades within and across ecosystems: The role of anti-predatory defences, predator type and detritus quality. Journal of Animal Ecology, 00, 1-14. https://doi.org/10.1111/1365-2656.14063. Ecosystem functioning is controlled by the interplay between bottom-up supply of limiting nutrients and top-down animal feedback effects. However, the degree of animal versus nutrient control is context-dependent. A key challenge lies in characterizing this context dependency which is hypothesized to depend on differences in animal functional traits. Reporting on an important experiment, Piccoli et al. (2014) evaluate how interactions among functionally different predators and decomposer prey create context dependency in top-down control of a model system-tropical bromeliad tank ecosystems. Bromeliad plants hold water in their tanks supporting microcosm ecosystems containing terrestrial and aquatic insect larvae and arachnids. The ecosystems are supported by nutrients in plant litter that rains down from forest canopies into the tanks. Nutrients are released after litter is decomposed by a functionally diverse community of larval insect decomposers that differ in feeding mode and antipredator defence strategy. This decomposer community is preyed upon by an exclusively narrowly ranging aquatic insect larval predator and widely ranging spider predator that crosses between the aquatic and surrounding terrestrial ecosystems. Experimental manipulation of the animal community to test for the degree of control by predators mediated by the functionally diverse prey community included four treatments: (i) a control with the detritivores composing different function groups but without predators, (ii) the cross-ecosystem spider predator added, (iii) the purely aquatic damselfly larvae predator added and (iv) both predator types added to capture their interacting effect on ecosystem function (decomposition, nutrient release, and plant growth). Notably, the study resolved the causal pathways and strengths of direct and indirect control using structural equation modelling. These findings reveal how context dependency arises due to different capacities of the predators alone and together to overcome prey defences and control their abundances, with attendant cascading effects that diminished as well as enhanced decomposition and nutrient release to support bromeliad plant production. The study reveals that predators have a decided, albeit qualitatively and quantitatively different, hand in shaping the degree of bottom-up control through feedback effect on the release of limiting nutrients. This ground-breaking study provides a way forward in understanding the mechanisms determining context dependency in the control over ecosystem functioning.

Mechanical traits as drivers of trophic interaction between macrodetritivores and leaf litter.

In ecosystems, the rates of resource consumption by animals drive the flows of matter and energy. Consumption rates are known to vary according to consumer energy requirements, resource nutrient content and mechanical properties. The aim of our study is to determine how mechanical constraints, compared to energetic and nutritional constraints, explain the variation in leaf litter consumption rates by macrodetritivores. In particular, we focus on the impact of litter toughness. To this end, we propose a non-linear model describing leaf litter consumption rates of detritivore as a function of litter toughness. We also investigate a possible match between bite force and litter toughness, since consumer-resource co-occurrence is thought to be driven by the match between invertebrate mandibular traits and resource toughness. Our study was designed as follows: leaf litter from oak and hornbeam was exposed to field physical and microbial decomposition in aquatic and terrestrial ecosystems for selected time periods before it was offered to eight macrodetritivore taxa (three forest stream taxa and five forest soil taxa) in no-choice laboratory feeding experiments. Our findings show that, compared to energetic and nutritional constraints, mechanical traits have a greater impact on litter consumption rate by detritivores. After subtracting the contribution of the detritivore body mass, we report that litter consumption rates depend primarily on litter toughness. A sigmoid function is best suited to characterize the relationship between mass-independent consumption rate and litter toughness. We note that the parameters of our sigmoid model are taxon-specific, suggesting biomechanical thresholds and biological differences among taxa. Interestingly, we found no correlation with detritivore bite force, suggesting that food processing by detritivores does not only depend on mandibles strength.

Energetic mismatch induced by warming decreases leaf litter decomposition by aquatic detritivores.

The balance of energetic losses and gains is of paramount importance for understanding and predicting the persistence of populations and ecosystem processes in a rapidly changing world. Previous studies suggested that metabolic rate often increases faster with warming than resource ingestion rate, leading to an energetic mismatch at high temperature. However, little is known about the ecological consequences of this energetic mismatch for population demography and ecosystem functions. Here, we combined laboratory experiments and modelling to investigate the energetic balance of a stream detritivore Gammarus fossarum along a temperature gradient and the consequences for detritivore populations and organic matter decomposition. We experimentally measured the energetic losses (metabolic rate) and supplies (ingestion rate) of Gammarus and we modelled the impact of rising temperatures and changes in Gammarus body size induced by warming on population dynamics and benthic organic matter dynamics in freshwater systems. Our experimental results indicated an energetic mismatch in a Gammarus population where losses via metabolic rate increase faster than supplies via food ingestion with warming, which translated in a decrease in energetic efficiency with temperature rising from 5 to 20°C. Moreover, our consumer-resource model predicts a decrease in the biomass of Gammarus population with warming, associated with lower maximum abundances and steeper abundance decreases after biomass annual peaks. These changes resulted in a decrease in leaf litter decomposition rate and thus longer persistence of leaf litter standing stock over years in the simulations. In addition, Gammarus body size reductions led to shorter persistence for both leaf litter and Gammarus biomasses at low temperature and the opposite trend at high temperature, revealing that body size reduction was weakening the effect of temperature on resource and consumer persistence. Our model contributes to identifying the mechanisms that explain how thermal effects at the level of individuals may cascade through trophic interactions and influence important ecosystem processes. Considering the balance of physiological processes is crucial to improve our ability to predict the impact of climate change on carbon stocks and ecosystem functions.

Invertebrate functional traits and terrestrial nutrient cycling: Insights from a global meta-analysis.

Functional traits are useful for characterizing variation in community and ecosystem dynamics. Most advances in trait-based ecology to date centre on plant functional traits, although there is an increasing recognition that animal traits are also key contributors to processes operating at the community or ecosystem scale. Terrestrial invertebrates are incredibly diverse and ubiquitous animals with important roles in nutrient cycling. Despite their widespread influence on ecosystem processes, we currently lack a synthetic understanding of how invertebrate functional traits affect terrestrial nutrient cycling. We present a meta-analysis of 511 paired observations from 122 papers that examined how invertebrate functional traits affected litter decomposition rates, nitrogen pools and litter C:N ratios. Based on the available data, we specifically assessed the effects of feeding mode (bioturbation, detritus shredding, detritus grazing, leaf chewing, leaf piercing, ambush predators, active hunting predators) and body size (macro- and micro-invertebrates) on nutrient cycling. The effects of invertebrates on terrestrial nutrient cycling varied according to functional trait. The inclusion of both macro- (≥2 mm) and micro-invertebrates (<2 mm) increased litter decomposition by 20% and 19%, respectively. All detritivorous feeding modes enhanced litter decomposition rates, with bioturbators, detritus shredders and detritus grazers increasing decomposition by 28%, 22% and 15%, respectively. Neither herbivore feeding mode (e.g. leaf chewers and leaf piercers) nor predator hunting mode (ambush and active hunting) affected decomposition. We also revealed that bioturbators and detritus grazers increased soil nitrogen availability by 99% and 70%, respectively, and that leaf-chewing herbivores had a weak effect on litterfall stoichiometry via reducing C:N ratios by 11%. Although functional traits might be useful predictors of ecosystem processes, our findings suggest context-dependent effects of invertebrate traits on terrestrial nutrient cycling. Detritivore functional traits (i.e. bioturbators, detritus shredders and detritus grazers) are more consistent with increased rates of nutrient cycling, whereas our currently characterized predator and herbivore traits are less predictive. Future research is needed to identify, standardize and deliberately study the impacts of invertebrate functional traits on nutrient cycling in hopes of revealing the key functional traits governing ecosystem functioning worldwide.

High sensitivity of invertebrate detritivores from tropical streams to different pesticides.

Freshwater organisms are often sensitive to pesticides, but their sensitivity varies across different taxa and with pesticide type and action mode, as shown by multiple acute toxicity tests. Such variability hampers predictions about how freshwater ecosystems may be altered by pesticide toxicity, which is especially critical for understudied areas of the world such as the tropics. Furthermore, there is little information about the sensitivity of some organisms that are key components of stream food webs; this is the case of litter-feeding detritivorous invertebrates, which contribute to the fundamental process of litter decomposition. Here, we examined the sensitivity of three common detritivores [Anchytarsus sp. (Coleoptera: Ptilodactylidae), Hyalella sp. (Amphipoda: Hyalellidae) and Lepidostoma sp. (Trichoptera: Lepidostomatidae)] to three pesticides commonly used (the insecticides bifenthrin and chlorpyrifos and the fungicide chlorothalonil) using acute (48 or 96 h) toxicity tests. Our study demonstrates that common-use pesticides provoke the mortality of half their populations at concentrations of 0.04-2.7 µg L-1. We found that all species were sensitive to the three pesticides, with the highest sensitivity found for chlorpyrifos. Additionally, we used the approach of species sensitivity distributions (SSD) to compare our study species with Daphnia magna and other temperate and tropical invertebrates. We found that the study species were among the most sensitive species to chlorpyrifos and chlorothalonil. Our results suggest that tropical detritivores merit special attention in ecological risk assessment of pesticides and highlight the need for accurate ecotoxicological information from ecologically relevant species in the tropics.

Habitat patchiness, ecological connectivity and the uneven recovery of boreal stream ecosystems from an experimental drought.

Ongoing climate change is increasing the occurrence and intensity of drought episodes worldwide, including in boreal regions not previously regarded as drought prone, and where the impacts of drought remain poorly understood. Ecological connectivity is one factor that might influence community structure and ecosystem functioning post-drought, by facilitating the recovery of sensitive species via dispersal at both local (e.g. a nearby habitat patch) and regional (from other systems within the same region) scales. In an outdoor mesocosm experiment, we investigated how impacts of drought on boreal stream ecosystems are altered by the spatial arrangement of local habitat patches within stream channels, and variation in ecological connectivity with a regional species pool. We measured basal ecosystem processes underlying carbon and nutrient cycling: (a) algal biomass accrual; (b) microbial respiration; and (c) decomposition of organic matter, and sampled communities of aquatic fungi and benthic invertebrates. An 8-day drought event had strong impacts on both community structure and ecosystem functioning, including algal accrual, leaf decomposition and microbial respiration, with many of these impacts persisting even after water levels had been restored for 3.5 weeks. Enhanced connectivity with the regional species pool and increased aggregation of habitat patches also affected multiple response variables, especially those associated with microbes, and in some cases reduced the effects of drought to a small extent. This indicates that spatial processes might play a role in the resilience of communities and ecosystem functioning, given enough time. These effects were however insufficient to facilitate significant recovery in algal growth before seasonal dieback began in autumn. The limited resilience of ecosystem functioning in our experiment suggests that even short-term droughts can have extended consequences for stream ecosystems in the world's vast boreal region, and especially on the ecosystem processes and services mediated by algal biofilms.

A common fungicide impairs stream ecosystem functioning through effects on aquatic hyphomycetes and detritivorous caddisflies.

Fungicides can reach streams through runoff or adhered to leaf litter, and have the potential to adversely affect processes such as litter decomposition and associated communities. This study investigated the effects of chlorothalonil, a widely used fungicide, on litter decomposition, detritivorous invertebrates (larvae of the insect Sericostoma pyrenaicum) and aquatic hyphomycetes (AHs), using stream microcosms. We considered the single and combined effects of two exposure modes: waterborne fungicide (at two concentrations: 0.125 µg L-1 and 1.25 µg L-1) and litter previously sprayed with the fungicide (i.e., pre-treated litter, using the application dose concentration of 1250 µg L-1). We also assessed whether fungicide effects on invertebrates, AHs and decomposition varied among litter types (i.e., different plant species), and whether plant diversity mitigated any of those effects. Invertebrate survival and AH sporulation rate and taxon richness were strongly reduced by most combinations of fungicide exposure modes; however, invertebrates were not affected by the low waterborne concentration, whereas AHs suffered the highest reduction at this concentration. Total decomposition was slowed down by both exposure modes, and microbial decomposition was reduced by litter pre-treatment, while the waterborne fungicide had different effects depending on plant species. In general, with the exception of microbial decomposition, responses varied little among litter types. Moreover, and contrary to our expectation, plant diversity did not modulate the fungicide effects. Our results highlight the severity of fungicide inputs to streams through effects on invertebrate and microbial communities and ecosystem functioning, even in streams with well-preserved, diverse riparian vegetation.

Sea urchins mediate the availability of kelp detritus to benthic consumers.

Detritus can fundamentally shape and sustain food webs, and shredders can facilitate its availability. Most of the biomass of the highly productive giant kelp, Macrocystis pyrifera, becomes detritus that is exported or falls to the seafloor as litter. We hypothesized that sea urchins process kelp litter through shredding, sloppy feeding and egestion, making kelp litter more available to benthic consumers. To test this, we conducted a mesocosm experiment in which an array of kelp forest benthic consumers were exposed to 13C- and 15N-labelled Macrocystis with or without the presence of sea urchins, Strongylocentrotus purpuratus. Our results showed that several detritivore species consumed significant amounts of kelp, but only when urchins were present. Although they are typically portrayed as antagonistic grazers in kelp forests, sea urchins can have a positive trophic role, capturing kelp litter before it is exported and making it available to a suite of benthic detritivores.

Burrowing detritivores regulate nutrient cycling in a desert ecosystem.

Nutrient cycling in most terrestrial ecosystems is controlled by moisture-dependent decomposer activity. In arid ecosystems, plant litter cycling exceeds rates predicted based on precipitation amounts, suggesting that additional factors are involved. Attempts to reveal these factors have focused on abiotic degradation, soil-litter mixing and alternative moisture sources. Our aim was to explore an additional hypothesis that macro-detritivores control litter cycling in deserts. We quantified the role different organisms play in clearing plant detritus from the desert surface, using litter baskets with different mesh sizes that allow selective entry of micro-, meso- or macrofauna. We also measured soil nutrient concentrations in increasing distances from the burrows of a highly abundant macro-detritivore, the desert isopod Hemilepistus reaumuri. Macro-detritivores controlled the clearing of plant litter in our field site. The highest rates of litter removal were measured during the hot and dry summer when isopod activity peaks and microbial activity is minimal. We also found substantial enrichment of inorganic nitrogen and phosphorous near isopod burrows. We conclude that burrowing macro-detritivores are important regulators of litter cycling in this arid ecosystem, providing a plausible general mechanism that explains the unexpectedly high rates of plant litter cycling in deserts.

When things don't add up: quantifying impacts of multiple stressors from individual metabolism to ecosystem processing.

Ecosystems are exposed to multiple stressors which can compromise functioning and service delivery. These stressors often co-occur and interact in different ways which are not yet fully understood. Here, we applied a population model representing a freshwater amphipod feeding on leaf litter in forested streams. We simulated impacts of hypothetical stressors, individually and in pairwise combinations that target the individuals' feeding, maintenance, growth and reproduction. Impacts were quantified by examining responses at three levels of biological organisation: individual-level body sizes and cumulative reproduction, population-level abundance and biomass and ecosystem-level leaf litter decomposition. Interactive effects of multiple stressors at the individual level were mostly antagonistic, that is, less negative than expected. Most population- and ecosystem-level responses to multiple stressors were stronger than expected from an additive model, that is, synergistic. Our results suggest that across levels of biological organisation responses to multiple stressors are rarely only additive. We suggest methods for efficiently quantifying impacts of multiple stressors at different levels of biological organisation.

Interactive effects of climate change and biodiversity loss on ecosystem functioning.

Climate change and biodiversity loss are expected to simultaneously affect ecosystems, however research on how each driver mediates the effect of the other has been limited in scope. The multiple stressor framework emphasizes non-additive effects, but biodiversity may also buffer the effects of climate change, and climate change may alter which mechanisms underlie biodiversity-function relationships. Here, we performed an experiment using tank bromeliad ecosystems to test the various ways that rainfall changes and litter diversity may jointly determine ecological processes. Litter diversity and rainfall changes interactively affected multiple functions, but how depends on the process measured. High litter diversity buffered the effects of altered rainfall on detritivore communities, evidence of insurance against impacts of climate change. Altered rainfall affected the mechanisms by which litter diversity influenced decomposition, reducing the importance of complementary attributes of species (complementarity effects), and resulting in an increasing dependence on the maintenance of specific species (dominance effects). Finally, altered rainfall conditions prevented litter diversity from fueling methanogenesis, because such changes in rainfall reduced microbial activity by 58%. Together, these results demonstrate that the effects of climate change and biodiversity loss on ecosystems cannot be understood in isolation and interactions between these stressors can be multifaceted.

Decadal-scale response of detritivorous surgeonfishes (family Acanthuridae) to no-take marine reserve protection and changes in benthic habitat.

No-take marine reserves (NTMR) are increasingly being implemented to mitigate the effects of fishing on coral reefs, yet determining the efficacy of NTMRs depends largely on partitioning the effects of fishing from the effect of benthic habitat. Species of coral-reef fishes typically decline in density when subjected to fishing or benthic disturbances, but this is not always the case. This study documents the long-term (8-31 years) response of six species of detritivorous surgeonfishes (family Acanthuridae) to NTMR protection and benthic habitat change at four islands (Apo, Sumilon, Mantigue, Selinog) in the central Philippines, each island with a NTMR and a monitored fished site. Despite being subject to moderate fishing pressure, these species did not increase in density with NTMR protection. However, density of these surgeonfishes had a strong negative relationship with cover of live hard coral and a strong positive relationship with cover of dead substratum (sand, rubble, hard dead substratum). These surgeonfishes typically feed over dead substrata and thus probably increase in density following large environmental disturbances that substantially reduce live hard coral cover. Here, we describe effects of environmental disturbance events (e.g., use of explosives, typhoons) that reduced live hard-coral cover and subsequent large increases (up to 25 fold) in surgeonfish densities, which then slowly (over 5-15 years) decreased in density as live hard coral recovered. Density of these functionally important surgeonfish species was influenced more by changes to benthic cover than by NTMR protection. Thus, we highlight the greater importance of bottom-up controls (i.e., benthic changes to food availability) than top-down control (i.e., fishing) on a functionally important group of coral-reef fishes.

Litter chemistry and chemical diversity drive ecosystem processes in forest ponds.

Research suggests that a positive relationship exists between diversity and ecological function, yet the multi-trophic effects of biodiversity remain poorly understood. The resource complementarity hypothesis suggests that increasing the trait diversity of resources provides a more complete diet for consumers, elevating consumer feeding rates. Whereas previous tests of this mechanism have measured trait diversity as the variation of single traits or the richness of functional groups, we employed a multivariate trait index to manipulate the chemical diversity of temperate tree litter species in outdoor pond mesocosms. We inoculated outdoor mesocosms with diverse and multi-trophic communities of microbial and macro-consumer species that rely on leaf litter for energy and nutrients. Litter was provided at three levels of chemical trait diversity, a constant level of species richness, and an equal representation of all litter species. Over three months, we measured more than 65 responses, and assessed the effects of litter chemical diversity and chemical trait means (i.e., community-weighted means). We found that litter chemical diversity positively correlated with decomposition rate of leaf litter, but had no effect on biomass or density of producers and consumers. However, the pond communities often responded to chemical trait means, particularly those related to nutrients, structure, and defense. Our results suggest that resource complementarity does have some effect on the release of energy and nutrients from decomposing substrates in forest ponds, but does not have multi-trophic effects. Our results further suggest that loss of tree biodiversity could affect forest ecosystem functionality, and particularly the processes occurring in and around ponds and wetlands.

Plant herbivory responses through changes in leaf quality have no effect on subsequent leaf-litter decomposition in a neotropical rain forest tree community.

It is commonly accepted that plant responses to foliar herbivory (e.g. plant defenses) can influence subsequent leaf-litter decomposability in soil. While several studies have assessed the herbivory-decomposability relationship among different plant species, experimental tests at the intra-specific level are rare, although critical for a mechanistic understanding of how herbivores affect decomposition and its consequences at the ecosystem scale. Using 17 tree species from the Yasuní National Park, Ecuadorian Amazonia, and applying three different herbivore damage treatments, we experimentally tested whether the plant intra-specific responses to herbivory, through changes in leaf quality, affect subsequent leaf-litter decomposition in soil. We found no effects of herbivore damage on the subsequent decomposition of leaf litter within any of the species tested. Our results suggest that leaf traits affecting herbivory are different from those influencing decomposition. Herbivore damage showed much higher intra-specific than inter-specific variability, while we observed the opposite for decomposition. Our findings support the idea that interactions between consumers and their resources are controlled by different factors for the green and the brown food-webs in tropical forests, where herbivory may not necessarily generate any direct positive or negative feedbacks for nutrient cycling.

Effects of invasive plants on arthropods.

Non-native plants have invaded nearly all ecosystems and represent a major component of global ecological change. Plant invasions frequently change the composition and structure of vegetation communities, which can alter animal communities and ecosystem processes. We reviewed 87 articles published in the peer-reviewed literature to evaluate responses of arthropod communities and functional groups to non-native invasive plants. Total abundance of arthropods decreased in 62% of studies and increased in 15%. Taxonomic richness decreased in 48% of studies and increased in 13%. Herbivorous arthropods decreased in response to plant invasions in 48% of studies and increased in 17%, likely due to direct effects of decreased plant diversity. Predaceous arthropods decreased in response to invasive plants in 44% of studies, which may reflect indirect effects due to reductions in prey. Twenty-two percent of studies documented increases in predators, which may reflect changes in vegetation structure that improved mobility, survival, or web-building for these species. Detritivores increased in 67% of studies, likely in response to increased litter and decaying vegetation; no studies documented decreased abundance in this functional group. Although many researchers have examined effects of plant invasions on arthropods, sizeable information gaps remain, specifically regarding how invasive plants influence habitat and dietary requirements. Beyond this, the ability to predict changes in arthropod populations and communities associated with plant invasions could be improved by adopting a more functional and mechanistic approach. Understanding responses of arthropods to invasive plants will critically inform conservation of virtually all biodiversity and ecological processes because so many organisms depend on arthropods as prey or for their functional roles, including pollination, seed dispersal, and decomposition. Given their short generation times and ability to respond rapidly to ecological change, arthropods may be ideal targets for restoration and conservation activities.

Effects of nanoplastic exposure routes on leaf decomposition in streams.

Polystyrene nanoparticles (PS NPs) released from plastic products have been demonstrated to pose a threat to leaf litter decomposition in streams. Given the multitrophic systems of species interactions, the effects of PS NPs through different exposure routes on ecosystem functioning remain unclear. Especially dietary exposure, a frequently overlooked pathway leading to toxicity, deserves more attention. A microcosm experiment was conducted in this study to assess the effects of waterborne and dietary exposure to PS NPs on the litter-based food chain involving leaves, microbial decomposers, and detritivores (river snails). Compared to waterborne contamination, dietary contamination resulted in lower microbial enzyme activities and a significantly higher decrease in the lipid content of leaves. For river snails, their antioxidant activity was significantly increased by 20.21%-69.93%, and their leaf consumption rate was significantly reduced by 16.60% through the dietary route due to the lower lipid content of leaves. Besides, the significantly decreased nutritional quality of river snails would negatively influence their palatability to predators. The findings of this study indicate that dietary exposure to PS NPs significantly impacts microbial and detritivore activities, thus affecting their functions in the detritus food chain as well as nutrient cycling.

The synergistic effects of a leaf mixture on decomposition change with a period of terrestrial exposure prior to immersion in a stream.

The effect of mixing litter on decomposition has received considerable attention in terrestrial and aquatic (but rarely in both) ecosystems, with a striking lack of consensus in the obtained results. We studied the decomposition of a mixture of poplar and alder in three terrestrial: aquatic exposures to determine (1) if the effect of mixing litter on mass loss, associated decomposers (fungal biomass, sporulation rates, and richness), and detritivores (abundance, biomass, and richness of invertebrate shredders) differs between the stream (fully aquatic exposure) and when litter is exposed to a period of terrestrial exposure prior to immersion and (2) the effect of the mixture across exposure scenarios. The effect of the mixture was additive on mass loss and synergistic on decomposers and detritivores across exposure scenarios. Within scenarios, mass loss and decomposers showed synergistic effects only in the fully aquatic exposure, detritivores showed synergistic effects only when the period of terrestrial was shorter than the period of aquatic exposure, and when the period of terrestrial was equal to the period of aquatic exposure the effect of the mixture was additive on mass loss, decomposers, and detritivores. The species-specific effects also differed among exposure scenarios. Alder affected poplar only when there was a period of terrestrial exposure, with increased sporulation rates and fungal richness in exposure 25:75, and increased mass loss in exposure 50:50. Poplar affected alder only under fully aquatic exposure, with increased mass loss. In conclusion, the synergistic effects of the mixture changed with a period of terrestrial exposure prior to immersion. These results provide a cross-boundary perspective on the effect of mixing litter, showing a legacy effect of exposure to terrestrial decomposition on the fate of plant litter in aquatic ecosystems and highlighting the importance of also assessing the effect of mixing litter on the associated biota and not only on mass loss.

Trophic complexity enhances ecosystem functioning in an aquatic detritus-based model system.

1. Understanding the functional significance of species interactions in ecosystems has become a major challenge as biodiversity declines rapidly worldwide. Ecosystem consequences arising from the loss of diversity either within trophic levels (horizontal diversity) or across trophic levels (vertical diversity) are well documented. However, simultaneous losses of species at different trophic levels may also result in interactive effects, with potentially complex outcomes for ecosystem functioning. 2. Because of logistical constraints, the outcomes of such interactions have been difficult to assess in experiments involving large metazoan species. Here, we take advantage of a detritus-based model system to experimentally assess the consequences of biodiversity change within both horizontal and vertical food-web components on leaf-litter decomposition, a fundamental process in a wide range of ecosystems. 3. Our concurrent manipulation of fungal decomposer diversity (0, 1 or 5 species), detritivore diversity (0, 1 or 3 species), and the presence of predatory fish scent showed that trophic complexity is key to eliciting diversity effects on ecosystem functioning. Specifically, although fungi and detritivores tended to promote decomposition individually, rates were highest in the most complete community where all trophic levels were represented at the highest possible species richness. In part, the effects were trait-mediated, reflected in the contrasting foraging responses of the detritivore species to predator scent. 4. Our results thus highlight the importance of interactive effects of simultaneous species loss within multiple trophic levels on ecosystem functioning. If a common phenomenon, this outcome suggests that functional ecosystem impairment resulting from widespread biodiversity loss could be more severe than inferred from previous experiments confined to varying diversity within single trophic levels.

Warming overrides eutrophication effects on leaf litter decomposition in stream microcosms.

Several human activities often result in increased nitrogen (N) and phosphorus (P) inputs to running waters through runoff. Although headwater streams are less frequently affected by these inputs than downstream reaches, the joint effects of moderate eutrophication and global warming can affect the functioning of these ecosystems, which represent two thirds of total river length and thus are of major global relevance. In a microcosm study representing streams from a temperate area (northern Spain), we assessed the combined effects of increased water temperature (10.0, 12.5, and 15.0 °C) and nutrient enrichment (control, high N, high P, and high N + P concentrations) on the key process of leaf litter decomposition (mediated by microorganisms and detritivores) and associated changes in different biological compartments (leaf litter, aquatic hyphomycetes and detritivores). While warming consistently enhanced decomposition rates and associated variables (leaf litter microbial conditioning, aquatic hyphomycete sporulation rate and taxon richness, and detritivore growth and nutrient contents), effects of eutrophication were weaker and more variable: P addition inhibited decomposition, addition of N + P promoted leaf litter conditioning, and detritivore stoichiometry was affected by the addition of both nutrients separately or together. In only a few cases (variables related to detritivore performance, but not microbial performance or leaf litter decomposition) we found interactions between warming and eutrophication, which contrasts with other experiments reporting synergistic effects. Our results suggest that both stressors can importantly alter the functioning of stream ecosystems even when occurring in isolation, although non-additive effects should not be neglected and might require exploring an array of ecosystem processes (not just leaf litter decomposition) in order to be detected.