Biodiversity and litter decomposition: a case study in a Mediterranean stream

The importance of riparian diversity for the functioning of headwater streams has been demonstrated repeatedly. For example, mixing litter from different riparian tree species can influence their decomposition rates, an effect that is contingent on both the litter assemblage and the detritivore community. However, the effects of mixing litter species have been studied mostly in temperate streams, and very few studies have been done in non- temperate streams. Mediterranean streams are often subjected to recurrent flow intermittency, and their riparian tree and stream detritivore communities have unique sets of species and traits, which probably influence the effects of litter mixtures on decomposition. We hypothesized that high dissimilarity in litter traits could promote effects of litter mixtures on decomposition that would be counteracted by the low abundance and small body size of detritivores in Mediterranean streams. We manipulated litter diversity and the size-class and presence/absence of detritivores in a 2nd-order Mediterranean stream in a 46-d experiment and found substantial but contingent effects on litter mixtures. Mixture effects were not significant on average, but both negative and positive effects of litter mixtures occurred. For instance, mixing soft and nutrient-rich litter species led to up to 9.6% increase in leaf mass loss. Microbial activity accounted for 85% of total leaf mass loss, and no effect of litter mixture was observed when detritivores were excluded. In contrast, the presence of detritivores, despite their relatively low abundance and diversity, was a key factor for litter decomposition and promoted effects of litter mixture. These results suggest that the extinction of a few key taxa (riparian tree species or large detritivores) could impair nutrient and C cycling in Mediterranean streams with potential consequences for stream food webs

Fungi are involved in the effects of litter mixtures on consumption by shredders

1. Decomposition of litter mixtures in both terrestrial and aquatic ecosystems often shows non- additive diversity effects on decomposition rate, generally interpreted in streams as a result of the feeding activity of macroinvertebrates. The extent to which fungal assemblages on mixed litter may influence consumption by macroinvertebrates remains unknown. 2. We assessed the effect of litter mixing on all possible three-species combinations drawn from four tree species (Alnus glutinosa, Betula pendula, Juglans regia and Quercus robur) on both fungal assemblages and the rate of litter consumption by a common shredder, Gammarus fossarum. After a 9-week inoculation in a stream, batches of leaf discs were taken from all leaf species within litter mixture combinations. Ergosterol, an indicator of fungal biomass, and the composition of fungal assemblages, assessed from the conidia released, were determined, and incubated litter offered to G. fossarum in a laboratory-feeding experiment. 3. Mixing leaf litter species enhanced both the Simpson’s index of the fungal assemblage and the consumption of litter by G. fossarum, but had no clear effect on mycelial biomass. Specifically, consumption rates of J. regia were consistently higher for mixed-species litter packs than for single-species litter. In contrast, the consumption rates of B. pendula were not affected by litter mixing, because of the occurrence of both positive and negative litter-mixing effects in different litter species combinations that counteracted each other. 4. In some litter combinations, the greater development of some fungal species (e.g. Clavariopsis aquatica) as shown by higher sporulation rates coincided with increased leaf consumption, which may have resulted from feeding preferences by G. fossarum for these fungi. 5. Where litter mixture effects on decomposition rate are mediated via shredder feeding, this could be due to indirect effects of the fungal assemblage

Litter identity mediates predator impacts on the functioning of an aquatic detritus-based food web

During past decades, several mechanisms such as resource quality and habitat complexity have been proposed to explain variations in the strength of trophic cascades across ecosystems. In detritus-based headwater streams, litter accumulations constitute both a habitat and a resource for detritivorous macroinvertebrates. Because litter edibility (which promotes trophic cascades) is usually inversely correlated with its structural complexity (which weakens trophic cascades), there is a great scope for stronger trophic cascades in litter accumulations that are dominated by easily degradable litter species. However, it remains unclear how mixing contrasting litter species (conferring both habitat complexity and high quality resource) may influence top–down controls on communities and processes. In enclosures exposed in a second-order stream, we manipulated litter species composition by using two contrasting litter (alder and oak), and the presence–absence of a macroinvertebrate predator (Cordulegaster boltonii larvae), enabling it to effectively exert predation pressure, or not, on detritivores (consumptive versus non-consumptive predation effects). Leaf mass loss, detritivore biomass and community structure were mostly controlled independently by litter identity and mixing and by predator consumption. However, the strength of predator control was mediated by litter quality (stronger on alder), and to a lesser extent by litter mixing (weaker on mixed litter). Refractory litter such as oak leaves may contribute to the structural complexity of the habitat for stream macroinvertebrates, allowing the maintenance of detritivore communities even when strong predation pressure occurs. We suggest that considering the interaction between top–down and bottom–up factors is important when investigating their influence on natural communities and ecosystem processes in detritus-based ecosystems

Seasonal variations overwhelm temperature effects on microbial processes in headwater streams: insights from a temperate thermal spring

Carbon cycling in headwater streams is mostly driven by the decomposition of allochthonous organic matter, and to a lesser extent by primary production. Quantifying the influence of temperature on these processes is therefore essential to better anticipate the consequences of global warming for stream ecological functioning. In this study, we measured alder litter microbial decomposition and associated fungal biomass and diversity, using leaf discs enclosed in fine-mesh bags along a natural geothermal temperature gradient, in both spring and winter. We monitored the chlorophyll-a accrual in biofilms growing on ceramic tiles. The temperature gradient, from upstream to downstream, ranged from 15.3 to 14.2 °C in spring and 18.2 to 13.2 °C in winter. Autotrophs and heterotrophs exhibited contrasting responses to temperature. The expected positive effect of temperature was actually observed for chlorophyll-a accrual only, while an apparent temperature-independence of litter decomposition rate was found. Moreover, temperature effects on heterotrophic and autotrophic organisms depended on the season, with higher litter decomposition rates, sporulation rates, fungal biomass and chlorophyll-a in spring, despite a lower mean water temperature than in winter. Together, these results suggest that the influence of temperature remained largely overrode by seasonal effects. This result is likely due to annual variations in light availability, and may involve indirect positive interactions between microbial primary producers and decomposers

Litter Quality Modulates Effects of Dissolved Nitrogen on Leaf Decomposition by Stream Microbial Communities

Rates of leaf litter decomposition in streams are strongly influenced both by inorganic nutrients dissolved in stream water and by litter traits such as lignin, nitrogen (N) and phosphorus (P) concentrations. As a result, decomposition rates of different leaf species can show contrasting responses to stream nutrient enrichment resulting from human activities. It is unclear, however, whether the root cause of such discrepancies in field observations is the interspecific variation in either litter nutrient or litter lignin concentrations. To address this question, we conducted a controlled laboratory experiment with a known fungal community to determine decomposition rates of 38 leaf species exhibiting contrasting litter traits (N, P and lignin concentrations), which were exposed to 8 levels of dissolved N concentrations representative of field conditions across European streams (0.07 to 8.96 mg N L−1). The effect of N enrichment on decomposition rate was modelled using Monod kinetics to quantify N effects across litter species. Lignin concentration was the most important litter trait determining decomposition rates and their response to N enrichment. In particular, increasing dissolved N supply from 0.1 to 3.0 mg N L−1 accelerated the decomposition of lignin-poor litter (e.g.  15% of lignin, 1.4× increase ± 0.2 SD, n = 9). Litter nutrient concentrations were less important, with a slight positive effect of P on decomposition rates and no effect of litter N. These results indicate that shifts in riparian vegetation towards species characterized by high litter lignin concentrations could alleviate the stimulation of C turnover by stream nutrient enrichment

Diversity patterns of leaf-associated aquatic hyphomycetes along a broad latitudinal gradient

Information about the global distribution of aquatic hyphomycetes is scarce, despite the primary importance of these fungi in stream ecosystem functioning. In particular, the relationship between their diversity and latitude remains unclear, due to a lack of coor- dinated surveys across broad latitudinal ranges. This study is a first report on latitudinal patterns of aquatic hyphomycete diversity associated with native leaf-litter species in five streams located along a gradient extending from the subarctic to the tropics. Exposure of leaf litter in mesh bags of three different mesh sizes facilitated assessing the effects of including or excluding different size-classes of litter-consuming invertebrates. Aquatic hyphomycete evenness was notably constant across all sites, whereas species richness and diversity, expressed as the Hill number, reached a maximum at mid-latitudes (Medi- terranean and temperate streams). These latitudinal patterns were consistent across litter species, despite a notable influence of litter identity on fungal communities at the local scale. As a result, the bell-shaped distribution of species richness and Hill diversity devi- ated markedly from the latitudinal patterns of most other groups of organisms. Differences in the body-size distribution of invertebrate communities colonizing the leaves had no effect on aquatic hyphomycete species richness, Hill diversity or evenness, but inverte- brates could still influence fungal communities by depleting litter, an effect that was not captured by the design of our experiment

Variable temperature effects between heterotrophic stream processes and organisms

Temperature is known to stimulate metabolism with cascading effects on multiple biological processes. These effects may, however, vary across processes, types of organisms or levels of biological organisation. They can also vary with nutrient availability, with potentially stronger temperature effects when nutrients are not limiting. This context dependence of temperature effects on processes challenges our ability to anticipate their consequences on ecosystems in a changing world. In headwater streams, the decomposition of allochthonous leaf litter, driven by both microbial decomposers and invertebrates, is known to respond to both temperature and nutrient availability. These food webs are highly tractable and a useful model system to investigate the variations of temperature effects on processes across types of organisms (microbes versus invertebrates), resource availability levels (nutrient concentration), and levels of biological organisation (from individual to ecosystem). In a microcosm experiment, we measured the effects of temperature and nitrogen availability (four levels each) on respiration rates of litter-consuming microbes and invertebrates and their decomposition activity in different contexts of food web complexity. The latter included one treatment without invertebrate detritivore (microbial decomposers only), three single invertebrate taxa (Gammarus, Potamophylax, and Sericostoma) treatments, and one mixed invertebrate taxa treatment (three‐species altogether). Microbial processes increased nearly exponentially with temperature (Arrhenius model, activation energy (± 95% confidence interval) = 0.56 ± 0.53 and 1.00 ± 0.23 eV for litter decomposition and respiration), while invertebrate‐driven processes increased (activation energy from 0.47–1.15 eV) up to a maximal value at an intermediate temperature (c. 11–15°C depending on species and process), above which process rates decreased. By contrast, litter consumption in mixed invertebrate species treatments was not significantly influenced by temperature, because of a negative effect of species mixing occurring above 12°C. Nitrogen had a weaker influence, only slightly stimulating litter consumption by mixed‐species invertebrates, which limited the scope for synergies with temperature effects. Our results raise issues about how aquatic litter consumers meet their energy requirements at high temperature and suggest that a general consequence of warming could be loss of carbon through mineralisation in headwater stream food webs. In several aspects, our results deviate from expectations based on universal relationships between temperature and individual metabolism (e.g. metabolic theory of ecology), suggesting that we may need to develop less simplistic assumptions to predict the consequence of warming on ecosystem processes

Biodiversity and litter decomposition in streams (the influence of trophic interactions)

TOULOUSE3-BU Sciences (315552104) / SudocTOULOUSE-Observ. Midi Pyréné (315552299) / SudocSudocFranceF

Data from: Phenotypic determinants of inter-individual variability of litter consumption rate in a detritivore population

No description availabl

Rota_et_al_2018_Oikos_Data_sets

Rota_et_al_2018_Oikos_Data_set