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

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

Context dependency of litter‐mixing effects on decomposition and nutrient release across a long‐term chronosequence

Litter decomposition is an important driver of terrestrial systems, and factors that determine decomposition rate for individual litter species have been widely studied. Fewer studies have explored the factors that regulate how mixing litters of multiple species affects litter decomposition and nutrient dynamics, and only a handful of studies have investigated how litter‐mixing effects may differ among different habitats or ecosystems, or how they respond to environmental gradients. We used a well‐established retrogressive chronosequence involving thirty lake islands in northern Sweden in which time since fire disturbance increases with decreasing island size; smaller islands therefore have reduced rates of aboveground and belowground ecosystem processes. On each of these islands we utilized plots with and without the long‐term experimental removal of shrubs. Litters from the six most common plant species on the islands were prepared in single‐, three‐ and six‐species litterbags, and placed on both the shrub‐removal and non‐removal plots on each island to decompose for one year. We found significant non‐additive effects of litter mixing on litter decomposition rates, on final litter N and P concentrations, and on litter N loss, but these non‐additive effects varied both in direction and magnitude with changed number of species, and even among litter mixtures with the same number of species. Further, the magnitude of non‐additive effects of litter mixing on both litter decomposition and nutrient dynamics was significantly influenced by both island size and the interaction between island size and shrub‐removal treatment. When shrubs were present, there was a U‐shaped relationship between these non‐additive effects and island size, while the relationship was positive when shrubs were removed. Hence, our results support previous findings that litter mixing may produce non‐additive effects on litter decomposition and nutrient dynamics, and that these effects tend to be idiosyncratic due to the importance of effects of individual species in the mixture. Most importantly, our results show that non‐additive litter‐mixing effects change greatly across environmental gradients, meaning that the biotic and abiotic characteristics of an ecosystem can be a powerful driver of the magnitude and even the direction of litter‐mixing effects on ecosystem processes.     

Non‐additive effects of invasive tree litter shift seasonal N release: a potential invasion feedback

Many invasive plant species strongly alter ecosystem processes by producing leaf litter that decomposes faster and releases N more quickly than that of native species. However, while most studies of invasive species litter impacts have only considered the decomposition of species in monoculture, forest litter layers typically contain litter from many species. Many litter mixtures decompose in a non‐additive manner, in which the mixture decomposes more quickly (synergistic effect) or more slowly (antagonistic effect) than would be expected based on decomposition of the component species’ litters in isolation. We investigated the potential for non‐additive effects of invasive species’ litter by conducting a one‐year litter bag experiment in which we mixed the litters of four native tree species with each of four invasive species. Litter mixtures frequently lost mass at non‐additive rates, although not at every loading ratio, and the presence, sign, and strength of effects depended on species composition. Non‐additive effects on N loss occurred in more litter combinations, and were almost always antagonistic at 90 days and synergistic at 365 days. Invasive species litter with lower C:N led to more strongly synergistic N loss with time. During the growing season, non‐additive patterns of N loss almost always resulted in increased N release – up to six times greater than would be expected based on single‐species decomposition. Consequently, we suggest that invasive species may further synchronize N release from the litter layer with plant N demand, enhancing any positive litter feedback to invasion. These results highlight the need to consider non‐additive effects of litter mixing in invaded forest communities, and suggest that estimates of invasive species’ impacts on ecosystem processes would be improved by considering these effects.     

Leaf litter identity alters the timing of lotic nutrient dynamics

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Contrasting effects of functionally distinct tadpole species on nutrient cycling and litter breakdown in a tropical rainforest stream

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Leaf‐Litter Mixtures Affect Breakdown and Macroinvertebrate Colonization Rates in a Stream Ecosystem

Previous work in terrestrial and aquatic ecosystems has suggested that the relationship between breakdown rates of leaf litter and plant species richness may change unpredictability due to non‐additive effects mediated by the presence of key‐species. By using single‐ and mixed‐species leaf bags (7 possible combinations of three litter species differing in toughness; common alder [Alnus glutinosa ], sweet chestnut [Castanea sativa ], and Spanish oak [Quercus ilex ilex ]), I tested whether leaf species diversity, measured as richness and composition, affects breakdown dynamics and macroinvertebrate colonization (abundance, richness and composition) during 90 days incubation in a stream. Decomposition rates were additive, i.e., observed decomposition rates were not different from expected ones. However, decomposition rates of individual leaf species were affected by the mixture, i.e., there were species‐specific responses to mixing litter. The invertebrate communities colonizing the mixtures were not richer and more diverse in mixtures than in single‐species leaf bags. On the opposite, mixing leaf species had a negative, non‐additive effect on rates of shredder and taxa colonization and on macroinvertebrate diversity. (© 2009 WILEY‐VCH Verlag GmbH & Co. KGaA, Weinheim)     

Effects of nutrient enrichment on boreal streams: invertebrates, fungi and leaf-litter breakdown

1. The effect of nutrient enrichment on structural (invertebrate indices) and functional (leaf‐litter breakdown rates) characteristics of stream integrity was studied in nine boreal streams. 2. The results showed predicted changes in biotic indices and leaf‐litter breakdown along a complex (principal component) nutrient gradient. Biotic indices were better correlated with nutrient effects than leaf‐litter breakdown. 3. Fungal biomass and invertebrate densities in the litter bags were positively correlated with leaf‐litter breakdown, and both were also positively related to the nutrient gradient. 4. Invertebrate community composition influenced breakdown rate. High breakdown rates at one site were associated with the high abundance of the detritivore Asellus aquaticus. 5. This study lends support to the importance of invertebrate and fungi as mediators of leaf‐litter decomposition. However, our study also shows that study design (length of incubation) can confound the interpretation of nutrient‐induced effects on decomposition.     

Macro‐detritivore identity drives leaf litter diversity effects

The importance of leaf litter diversity for decomposition, an important process in terrestrial ecosystems, is much debated. Previous leaf litter‐mixing studies have shown that non‐additive leaf litter diversity effects can occur, but it is not clear why they occurred in only half of the studies and which underlying mechanisms can explain these conflicting results. We hypothesized that incorporating the role of macro‐detritivores could be important. Although often ignored, macro‐detritivores are known to strongly influence decomposition. To better understand the importance of macro‐detritivores for leaf litter mixing effects during decomposition, four common leaf litter species were added separately and in two and four species combinations to monocultures of three different macro‐detritivores and a control without fauna. Our results clearly show that leaf litter‐mixing effects occurred only in the presence of two macro‐detritivores (earthworms and woodlice). Application of the additive partitioning method revealed that in the specific combination of woodlice and the presence of a slow‐decomposing leaf litter species in the mixture, leaf litter mixing effects were strongly driven by a selection effect. This was caused by food preference of the isopod: the animals avoided the slow decomposing species when given the choice. However, most leaf litter mixing effects were caused by complementarity effects. The potential mechanisms underlying the complementarity effects are discussed. Our results clearly show that that both leaf litter and macro‐detritivore identity can affect litter diversity. This may help to explain the conflicting results obtained in previous experiments.     

Tree species richness affects litter production and decomposition rates in a tropical biodiversity experiment

We report data on leaf litter production and decomposition from a manipulative biodiversity experiment with trees in tropical Panama, which has been designed to explore the relationship between tree diversity and ecosystem functioning. A total of 24 plots (2025 m²) were established in 2001 using six native tree species, with 1‐, 3‐, and 6‐species mixtures. We estimated litter production during the dry season 2005 with litter traps; decomposition was assessed with a litter bag approach during the following wet season. Litter production during the course of the dry season was highly variable among the tree species. Tree diversity significantly affected litter production, and the majority of the intermediate diverse mixtures had higher litter yields than expected based on yields in monoculture. In contrast, high diverse mixtures did not show such overyielding in litter production. Litter decomposition rates were also highly species‐specific, and were related to various measures of litter quality (C/N, lignin/N, fibre content). We found no overall effect of litter diversity if the entire litter mixtures were analyzed, i.e. mixing species resulted in pure additive effects and observed decomposition rates were not different from expected rates. However, the individual species changed their decomposition pattern depending on the diversity of the litter mixture, i.e. there were species‐specific responses to mixing litter. The analysis of temporal C and N dynamics within litter mixtures gave only limited evidence for nutrient transfer among litters of different quality. At this early stage of our tree diversity experiment, there are no coherent and general effects of tree species richness on both litter production and decomposition. Within the scope of the biodiversity‐ecosystem functioning relationship, our results therefore highlight the process‐specific effects diversity may have. Additionally, species‐specific effects on ecosystem processes and their temporal dynamics are important, but such effects may change along the gradient of tree diversity.     

General empirical models for predicting the release of nutrients by fish,with a comparison between detritivores and non‐detritivores

1. We derived models of nutrient release [nitrogen (N) and phosphorus (P)] by fish based on studies that directly measured the release rates from 56 species across a broad range of fish mass, feeding histories and temperature. 2. We developed four separate models of nutrient release from multiple regression analysis: detritivore release rates of N and P, and non‐detritivore release rates of N and P. 3. Fish mass explained most of the variance (78–92%) in release rates. 4. Our predicted rates of release of P by fish (g ha^?1 day^?1) were similar to observed rates in the literature from other lakes. 5. The influence of a shift in diet (planktivory to detritivory) by a single species (gizzard shad, Dorosoma cepedianum, a facultative detritivore) on nutrient release rates was estimated. During periods of detritivory, gizzard shad accounted for on average 39% (<1–96%) of all nutrients released by the fish assemblage, and increased total fish assemblage release rates on average by 59% (<1–331%) compared to when gizzard shad were modelled as planktivores. 6. These models provide a rapid means for predicting the release of nutrients by fish assemblages and may facilitate more comprehensive comparisons of nutrient cycling by fish with other internal pathways.     

Insect biological control accelerates leaf litter decomposition and alters short‐term nutrient dynamics in a Tamarix‐invaded riparian ecosystem

Insect herbivory can strongly influence ecosystem nutrient dynamics, yet the indirect effects of herbivore‐altered litter quality on subsequent decomposition remain poorly understood. The northern tamarisk beetle Diorhabda carinulata was released across several western states as a biological control agent to reduce the extent of the invasive tree Tamarix spp. in highly‐valued riparian ecosystems; however, very little is currently known about the effects of this biocontrol effort on ecosystem nutrient cycling. In this study, we examined alterations to nutrient dynamics resulting from beetle herbivory in a Tamarix‐invaded riparian ecosystem in the Great Basin Desert in northern Nevada, USA, by measuring changes in litter quality and decomposition, as well as changes in litter quantity. Generally, herbivory resulted in improved leaf litter chemical quality, including significantly increased nitrogen (N) and phosphorus (P) concentrations and decreased carbon (C) to nitrogen (C:N), C:P, N:P, and lignin:N ratios. Beetle‐affected litter decomposed 23% faster than control litter, and released 16% more N and 60% more P during six months of decomposition, as compared to control litter. Both litter types showed a net release of N and P during decomposition. In addition, herbivory resulted in significant increases in annual rates of total aboveground litter and leaf litter production of 82% and 71%, respectively, under the Tamarix canopy. Our finding that increased rates of N and P release linked with an increased rate of mass loss during decomposition resulting from herbivore‐induced increases in litter quality provides new support to the nutrient acceleration hypothesis. Moreover, results of this study demonstrate that the introduction of the northern tamarisk beetle as biological control to a Tamarix‐invaded riparian ecosystem has lead to short‐term stimulation of nutrient cycling. Alterations to nutrient dynamics could have implications for future plant community composition, and thus the potential for restoration of Tamarix‐invaded ecosystems.     

Decomposition dynamics in mixed-species leaf litter

Literature on plant leaf litter decomposition is substantial, but only in recent years have potential interactions among leaves of different species during decomposition been examined. We review emerging research on patterns of mass loss, changes in nutrient concentration, and decomposer abundance and activity when leaves of different species are decaying in mixtures. Approximately 30 papers have been published that directly examine decomposition in leaf mixtures as well as in all component species decaying alone. From these litter‐mix experiments, it is clear that decomposition patterns are not always predictable from single‐species dynamics. (Characteristics of decomposition in litter‐mixes that deviate from responses predicted from decomposition of single‐species litters alone are designated "non‐additive"; "additive" responses in mixes are predictable from component species decaying alone.) Non‐additive patterns of mass loss were observed in 67% of tested mixtures; mass loss is often (though not always) increased when litters of different species are mixed. Observed mass loss in some mixtures is as much as 65% more extensive than expected from decomposition of single‐species litter, but more often mass loss in mixtures exceeds expected decay by 20% or less. Nutrient transfer among leaves of different species is striking, with 76% of the mixtures showing non‐additive dynamics of nutrient concentrations. Non‐additive patterns in the abundance and activity of decomposers were observed in 55% and 65% of leaf mixes, respectively. We discuss some methodological details that likely contribute to conflicting results among mixed‐litter studies to date. Enough information is available to begin formulating mechanistic hypotheses to explain patterns in litter‐mix experiments. Emerging patterns in the mixed‐litter decomposition literature have implications for relationships between biodiversity and ecosystem function (in this case, the function being decomposition), and for potential mechanisms through which invasive plant species could alter carbon and nutrient dynamics in ecosystems.     

C : N : P stoichiometry of dominant riparian trees and arthropods along the Middle Rio Grande

1. We examined the role of flooding on the leaf nutrient content of riparian trees by comparing the carbon : nitrogen : phosphorus (C : N : P) ratio of leaves and litter of Rio Grande cottonwood (Populus deltoides ssp. wislizenii) in flood and non‐flood sites along the Middle Rio Grande, NM, U.S.A. The leaf C : N : P ratio was also examined for two non‐native trees, saltcedar (Tamarix chinensis) and Russian olive (Elaeagnus angustifolia), and six species of dominant riparian arthropods. 2. Living leaves and leaf litter of cottonwoods at flood sites had a significantly lower leaf N : P ratio and higher %P compared with leaves and litter at non‐flood sites. A non‐flood site downstream from wastewater effluent had a significantly lower litter C : N ratio than all other sites, suggesting N fertilisation through ground water. The non‐native trees, saltcedar and Russian olive, had higher mean leaf N content, N : P ratio, and lower C : N ratio compared with cottonwoods across study sites. 3. Riparian arthropods ranged from 5.2 to 7.1 for C : N ratio, 56–216 for C : P ratio, and 8.9–34 for N : P ratio. C content ranged from 25 to 52% of dry mass, N content from 4.7 to 10.8%, and P content from 0.59 to 1.2%. Differences in stoichiometry between high C : nutrient leaf litter and low C : nutrient invertebrates suggests possible food‐quality constraints for detritivores. 4. These results suggest that spatial and temporal variation in the C : N : P ratio of cottonwood leaves and leaf litter is influenced by surface and subsurface hydrologic connection within the floodplain. Reach‐scale variation in the elemental composition of riparian organic matter inputs may have important implications for decomposition, nutrient cycling, and food webs in river floodplain systems.     

Decomposition and nutrient release patterns of mistletoe litters in a semi‐arid savanna,southwest Zimbabwe

Nutrient loss from litter plays an essential role in carbon and nutrient cycling in nutrient‐constrained environments. However, the decomposition and nutrient dynamics of nutrient‐rich mistletoe litter remains unknown in semi‐arid savanna where productivity is nutrient limited. We studied the decomposition and nutrient dynamics (nitrogen: N, phosphorous; P, carbon: C) of litter of three mistletoe species, Erianthemum ngamicum, Plicosepalus kalachariensis, and Viscum verrucosum and N‐fixing Acacia karroo using the litter‐bag method in a semi‐arid savanna, southwest Zimbabwe. The temporal dynamics of the soil moisture content, microbial populations, and termite activity during decomposition were also assessed. Decay rates were slower for A. karroo litter (k = 0.63), but faster for the high quality mistletoe litters (mean k‐value = 0.79), which supports the premise that mistletoes can substantially influence nutrient availability to other plants. Nitrogen loss was between 1.3 and 3 times greater in E. ngamicum litter than in the other species. The litter of the mistletoes also lost C and P faster than A. karroo litter. However, soil moisture content and bacterial and fungal colony numbers changed in an opposite direction to changes in the decomposition rate. Additionally, there was little evidence of termite activity during the decay of all the species litters. This suggests that other factors such as photodegradation could be important in litter decomposition in semi‐arid savanna. In conclusion, the higher rate of decay and nutrient release of mistletoe than A. karroo litter indicate that mistletoes play an important role in carbon and nutrient fluxes in semi‐arid savanna.     

Grazing history effects on above- and below-ground litter decomposition and nutrient cycling in two co-occurring grasses

Large herbivores may alter carbon and nutrient cycling in soil by changing above- and below-ground litter decomposition dynamics. Grazing effects may reflect changes in plant allocation patterns, and thus litter quality, or the site conditions for decomposition, but the relative roles of these broad mechanisms have rarely been tested. We examined plant and soil mediated effects of grazing history on litter mass loss and nutrient release in two grazing-tolerant grasses, Lolium multiflorum and Paspalum dilatatum, in a humid pampa grassland, Argentina. Shoot and root litters produced in a common garden by conspecific plants collected from grazed and ungrazed sites were incubated under both grazing conditions. We found that grazing history effects on litter decomposition were stronger for shoot than for root material. Root mass loss was neither affected by litter origin nor incubation site, although roots from the grazed origin immobilised more nutrients. Plants from the grazed site produced shoots with higher cell soluble contents and lower lignin:N ratios. Grazing effects mediated by shoot litter origin depended on the species, and were less apparent than incubation site effects. Lolium shoots from the grazed site decomposed and released nutrients faster, whereas Paspalum shoots from the grazed site retained more nutrient than their respective counterparts from the ungrazed site. Such divergent, species-specific dynamics did not translate into consistent differences in soil mineral N beneath decomposing litters. Indeed, shoot mass loss and nutrient release were generally faster in the grazed grassland, where soil N availability was higher. Our results show that grazing influenced nutrient cycling by modifying litter breakdown within species as well as the soil environment for decomposition. They also indicate that grazing effects on decomposition are likely to involve aerial litter pools rather than the more recalcitrant root compartment.     

Shifts in leaf litter breakdown along a forest–pasture–urban gradient in Andean streams

Tropical montane ecosystems of the Andes are critically threatened by a rapid land‐use change which can potentially affect stream variables, aquatic communities, and ecosystem processes such as leaf litter breakdown. However, these effects have not been sufficiently investigated in the Andean region and at high altitude locations in general. Here, we studied the influence of land use (forest–pasture–urban) on stream physico‐chemical variables (e.g., water temperature, nutrient concentration, and pH), aquatic communities (macroinvertebrates and aquatic fungi) and leaf litter breakdown rates in Andean streams (southern Ecuador), and how variation in those stream physico‐chemical variables affect macroinvertebrates and fungi related to leaf litter breakdown. We found that pH, water temperature, and nutrient concentration increased along the land‐use gradient. Macroinvertebrate communities were significantly different between land uses. Shredder richness and abundance were lower in pasture than forest sites and totally absent in urban sites, and fungal richness and biomass were higher in forest sites than in pasture and urban sites. Leaf litter breakdown rates became slower as riparian land use changed from natural to anthropogenically disturbed conditions and were largely determined by pH, water temperature, phosphate concentration, fungal activity, and single species of leaf‐shredding invertebrates. Our findings provide evidence that leaf litter breakdown in Andean streams is sensitive to riparian land‐use change, with urban streams being the most affected. In addition, this study highlights the role of fungal biomass and shredder species (Phylloicus; Trichoptera and Anchytarsus; Coleoptera) on leaf litter breakdown in Andean streams and the contribution of aquatic fungi in supporting this ecosystem process when shredders are absent or present low abundance in streams affected by urbanization. Finally, we summarize important implications in terms of managing of native vegetation and riparian buffers to promote ecological integrity and functioning of tropical Andean stream ecosystems.     

Trophic‐level dependent effects on CO2 emissions from experimental stream ecosystems

Concern over accelerating rates of species invasions and losses have initiated investigations into how local and global changes to predator abundance mediate trophic cascades that influence CO₂ fluxes of aquatic ecosystems. However, to date, no studies have investigated how species additions or losses at other consumer trophic levels influence the CO₂ flux of aquatic ecosystems. In this study, we added a large predatory stonefly, detritivorous stonefly, or grazer tadpole to experimental stream food webs and over a 70‐day period quantified their effects on community composition, leaf litter decomposition, chlorophyll‐a concentrations, and stream CO₂ emissions. In general, streams where the large grazer or large detritivore were added showed no change in total invertebrate biomass, leaf litter loss, chlorophyll‐a concentrations, or stream CO₂ emissions compared with controls; although we did observe a spike in CO₂ emissions in the large grazer treatment following a substantial reduction in chlorophyll‐a concentrations on day 28. However, the large grazer and large detritivore altered the community composition of streams by reducing the densities of other grazer and detritivore taxa, respectively, compared with controls. Conversely, the addition of the large predator created trophic cascades that reduced total invertebrate biomass and increased primary producer biomass. The cascading effects of the predator additions on the food web ultimately led to decreased CO₂ emissions from stream channels by up to 95%. Our results suggest that stream ecosystem processes were more influenced by changes in large predator abundance than large grazer or detritivore abundance, because of a lack of functionally similar large predators. Our study demonstrates that the presence/absence of species with unique functional roles may have consequences for the exchange of CO₂ between the ecosystem and the atmosphere.     

Resource quality and stoichiometric constraints on stream ecosystem functioning

1. Resource quality and stoichiometric imbalances in carbon : nutrient ratios between consumers and resources can influence key ecosystem processes. In many streams, this has important implications for food webs that are based largely upon the utilization of terrestrial leaf‐litter, which varies widely among litter types in its value as a food source for detritivores and as a substrate for microbial decomposers. 2. We measured breakdown rates and macroinvertebrate colonization of leaf‐litter from a range of native and exotic plants of differing resource quality and palatability to consumers [e.g. carbon : nitrogen : phosphorus (C : N : P) ratios, lignin and cellulose content], in a field experiment. We also measured C : N : P ratios of the principal leaf‐shredding invertebrates, which revealed strong stoichiometric imbalances across trophic levels: C : N and C : P ratios typically differed by at least one order of magnitude between consumers and resources, whereas N : P imbalances were less marked. Application of the threshold elemental ratio approach, which integrates animal bioenergetics and body elemental composition in examining nutrient deficiency between consumers and resources, revealed less marked C : P imbalances than those based on the simpler arithmetic differences described above. 3. Litter breakdown rates declined as nutrient imbalances widened and resource quality fell, but they were independent of whether resources were exotic or native. The principal drivers of total, microbial and invertebrate‐mediated breakdown rates were lignin : N, lignin : P and fungal biomass, respectively. However, multiple regression using orthogonal predictors yielded even more efficient models of litter breakdown, as consumers responded to more than one aspect of resource quality. For example, fungal biomass and litter C : N both influenced invertebrate‐mediated breakdown. 4. Large stoichiometric imbalances and changes in resource quality are likely to have serious consequences for stream ecosystem functioning, especially when riparian zones have been invaded by exotic plant species whose chemical composition differs markedly from that of the native flora. Consequently, the magnitude and direction of change in breakdown rates and, thus, resource depletion, will be driven to a large extent by the biochemical traits (rather than taxonomic identity per se) of the resident and invading flora.     

陆地生态系统混合凋落物分解研究进展

凋落物分解在陆地生态系统养分循环与能量流动中具有重要作用,是碳、氮及其他重要矿质养分在生态系统生命组分间循环与平衡的核心生态过程。自然生态系统中,植物群落大多具有较高的物种丰富度和多样性,其混合凋落物在分解过程中也更有可能发生养分传递、化学抑制等种间互作,形成多样化的分解生境,多样性较高的分解者类群以及复杂的级联效应分解,这些因素和过程均对研究混合凋落物分解过程、揭示其内在机制形成了极大的挑战。从构成混合凋落物物种丰富度和多样性对分解生境、分解者多样性及其营养级联效应的影响等方面,综合阐述混合凋落物对陆地生态系统凋落物分解的影响,探讨生物多样性在凋落物分解中的作用。通过综述近些年的研究发现,有超过60%的混合凋落物对其分解速率的影响存在正向或负向的效应。养分含量有差异的凋落物混合分解过程中,分解者优先利用高质量凋落物,使低质量的凋落物反而具有了较高的养分有效性,引起低质量凋落物分解加快并最终使混合凋落物整体分解速率加快;而凋落物物种丰富度对土壤动物群落总多度有轻微的影响或几乎没有影响,但是对线虫和大型土壤动物的群落组成和多样性有显著影响,并随着分解阶段呈现一定动态变化;混合凋落物改变土壤微生物生存的理化环境,为微生物提供更多丰富的分解底物和养分,优化微生物种群数量和群落结构及其分泌酶的活性,并进一步促进了混合凋落物的分解。这些基于植物-土壤-分解者系统的动态分解过程的研究,表明混合凋落物分解作用不只是经由凋落物自身质量的改变,更会通过逐级影响分解者多样性水平而进一步改变分解速率和养分释放动态,说明生物多样性确实在一定程度上调控凋落物分解及其养分释放过程。     

Inter‐annual changes in detritus‐based food chains can enhance plant growth response to elevated atmospheric CO2

Elevated atmospheric CO₂ generally enhances plant growth, but the magnitude of the effects depend, in part, on nutrient availability and plant photosynthetic pathway. Due to their pivotal role in nutrient cycling, changes in abundance of detritivores could influence the effects of elevated atmospheric CO₂ on essential ecosystem processes, such as decomposition and primary production. We conducted a field survey and a microcosm experiment to test the influence of changes in detritus‐based food chains on litter mass loss and plant growth response to elevated atmospheric CO₂ using two wetland plants: a C₃ sedge (Scirpus olneyi) and a C₄ grass (Spartina patens). Our field study revealed that organism's sensitivity to climate increased with trophic level resulting in strong inter‐annual variation in detritus‐based food chain length. Our microcosm experiment demonstrated that increased detritivore abundance could not only enhance decomposition rates, but also enhance plant growth of S. olneyi in elevated atmospheric CO₂ conditions. In contrast, we found no evidence that changes in the detritus‐based food chains influenced the growth of S. patens. Considered together, these results emphasize the importance of approaches that unite traditionally subdivided food web compartments and plant physiological processes to understand inter‐annual variation in plant production response to elevated atmospheric CO_2.