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.     

Non‐additive effects of litter mixing are suppressed in a nutrient‐enriched stream

Investigations of how species compositional changes interact with other aspects of global change, such as nutrient mobilization, to affect ecosystem processes are currently lacking. Many studies have shown that mixed species plant litters exhibit non‐additive effects on ecosystem functions in terrestrial and aquatic systems. Using a full‐factorial design of three leaf litter species with distinct initial chemistries (carbon:nitrogen; C:N) and breakdown rates (Liriodendron tulipifera, Acer rubrum and Rhododendron maximum), we tested for additive and non‐additive effects of litter species mixing on breakdown in southeastern US streams with and without added nutrients (N and phosphorus). We found a non‐additive (antagonistic) effect of litter mixing on breakdown rates under reference conditions but not when nutrient levels were elevated. Differential responses among single‐species litters to nutrient enrichment contributed to this result. Antagonistic litter mixing effects on breakdown were consistent with trends in litter C:N, which were higher for mixtures than for single species, suggesting lower microbial colonization on mixtures. Nutrient enrichment lowered C:N and had the greatest effect on the lowest‐ (R. maximum) and the least effect on the highest‐quality litter species (L. tulipifera), resulting in lower interspecific variation in C:N. Detritivore abundance was correlated with litter C:N in the reference stream, potentially contributing to variation in breakdown rates. In the nutrient‐enriched stream, detritivore abundance was higher for all litter and was unrelated to C:N. Thus, non‐additive effects of litter mixing were suppressed by elevated streamwater nutrients, which increased nutrient content of all litter, reduced variation in C:N among litter species and increased detritivore abundance. Nutrients reduced interspecific variation among plant litters, the base of important food web pathways in aquatic ecosystems, affecting predicted mixed‐species breakdown rates. More generally, world‐wide mobilization of nutrients may similarly modify other effects of biodiversity on ecosystem processes.     

Litter quality, stream characteristics and litter diversity influence decomposition rates and macroinvertebrates

1. We examined the relative importance of litter quality and stream characteristics in determining decomposition rate and the macroinvertebrate assemblage living on autumn‐shed leaves. 2. We compared the decomposition rates of five native riparian tree species (Populus fremontii, Alnus oblongifolia, Platanus wrightii, Fraxinus velutina and Quercus gambelii) across three south‐western streams in the Verde River catchment (Arizona, U.S.A.). We also compared the decomposition of three‐ and five‐species mixtures to that of single species to test whether plant species diversity affects rate. 3. Decomposition rate was affected by both litter quality and stream. However, litter quality accounted for most of the variation in decomposition rates. The relative importance of litter quality decreased through time, explaining 97% of the variation in the first week but only 45% by week 8. We also found that leaf mixtures decomposed more quickly than expected, when all the species included were highly labile or when the stream environment led to relatively fast decomposition. 4. In contrast to decomposition rate, differences in the invertebrate assemblage were more pronounced across streams than across leaf litter species within a stream. We also found significant differences between the invertebrate assemblage colonising leaf mixtures compared with that colonising pure species litter, indicating non‐additive properties of litter diversity on stream invertebrates. 5. This study shows that leaf litter diversity has the capacity to affect in‐stream decomposition rates and stream invertebrates, but that these effects depend on both litter quality and stream characteristics.     

Assessing the influence of wildfire on leaf decomposition and macroinvertebrate communities in boreal streams using mixed-species leaf packs

1. We investigated how compositional differences in riparian leaf litter derived from burned and undisturbed forests influenced leaf breakdown and macroinvertebrate communities using experimental mixed-species leaf packs in boreal headwater streams. Leaf pack mixtures simulating leaf litter from dominant riparian woody-stem species in burned and undisturbed riparian zones were incubated in two references and two fire-disturbed streams for 5 weeks prior to measuring temperature-corrected breakdown rates and macroinvertebrate community composition, richness, and functional metrics associated with decomposers such as shredder abundance and % shredders.
2. Leaf litter breakdown rates were higher and had greater variability in streams bordered by reference riparian forests than in streams where riparian forests had been burned during a wildfire. Streams bordered by fire disturbance showed significant effects of litter mixture on decomposition rates, observed as significantly higher decomposition rates of a fire-simulated leaf mixture compared to all other mixtures.
3. Variation among sites was higher than variation among litter mixtures, especially for macroinvertebrate community composition. In general, fire-simulated leaf mixtures had greater shredder abundances and proportions, but lower overall macroinvertebrate abundance; however, the shredder abundance trend was not consistent across all leaf mixtures at each stream.
4. These results show that disturbance-driven riparian forest condition and resulting composition of leaf subsidies to streams can influence aquatic invertebrate community composition and their function as decomposers. Therefore, if one of the primary goals of modern forest management is to emulate natural disturbance patterns, boreal forest managers should adapt silvicultural practices to promote leaf litter input that would arise post-fire, thereby supporting stream invertebrate communities and their function.

     

Do physical plant litter traits explain non‐additivity in litter mixtures? A test of the improved microenvironmental conditions theory

The decomposition rates of plant litter mixtures often deviate from the averaged rates of monocultures of their component litter species. The mechanisms behind these non‐additive effects in decomposition of litter mixtures are lively debated. One plausible explanation for non‐additive effects is given by the improved microenvironmental condition (IMC) theory. According to this theory, plant litter species, whose physical characteristics improve the microclimatic conditions for decomposers, will promote the decomposition of their co‐occurring litter species. We tested the IMC theory in relation to leaf litter and soil moisture in two contrasting moisture conditions in a dry subarctic mountain birch forest with vascular plant leaf litters of poor and high quality. The non‐additive effects in mass loss of litter mixtures increased when moisture conditions in litter and soil became more favourable for plant litter decomposition. The sign of this increase (antagonistic or synergistic) in non‐additive effects was more predictable for litter mixtures of poor litter quality. Although the specific mechanisms underlying the IMC theory depended on the litter quality of the litter mixtures, a standardized water holding capacity (WHC) was the litter trait most closely related to the non‐additive effects in mixtures of both poor and high quality litter types. Furthermore, we found that higher dissimilarity in WHC traits between the component litter species in a mixture increased synergistic effects in litter mixtures under limiting moisture conditions. However, under improved moisture conditions, increased antagonistic effects were observed. Thus, we found clear support for the IMC theory and showed that climatic conditions and leaf litter physical traits determine whether the non‐additive effects in litter mixtures are antagonistic or synergistic. Our study emphasizes the need to include litter physical traits into predictive models of mixing effects on plant litter decomposition and in general suggests climate specificity into these models.     

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.     

The Effect of Macroinvertebrate Exclusion on Leaf Breakdown Rates in a Tropical Headwater Stream1

The role of macroinvertebrates in the process of leaf breakdown is well studied in temperate streams, but less is known about their role in the tropics. We investigated the effect of reducing macroinvertebrate access to leaf material on leaf breakdown rates in a forested headwater stream in the Luquillo Experimental Forest, Puerto Rico. We measured leaf mass loss using fine and coarse mesh bags over 12 weeks for two common riparian species: Cecropia schreberiana (Moraceae) and Dacryodes excelsa (Burseraceae). Coarse mesh allowed freshwater shrimp and other macroinvertebrates to access leaf material, while fine mesh did not. Leaf breakdown rates did not differ between C. schreberiana and D. excelsa in coarse mesh bags (?0.0375/day vs. ?0.0395/day, respectively), but C. schreberiana breakdown was significantly slower than D. excelsa in fine mesh bags (?0.0159/day vs. ?0.0266/day). Additionally, breakdown in fine mesh bags was significantly slower compared to coarse mesh bags for C. schreberiana, but less so for D. excelsa. Breakdown rates for all treatments were fast relative to those in temperate‐zone streams indicating that both macroinvertebrates and macroinvertebrate‐independent processing can strongly influence leaf decomposition in tropical streams. The difference between C. schreberiana and D. excelsa indicates that the effect of macroinvertebrate exclusion can change with leaf type.     

Limited evidence that mixing leaf litter accelerates decomposition or increases diversity of decomposers in streams of eastern Canada

Most studies of terrestrial litter decomposition in streams and rivers have used leaves from a single tree species, but leaf packs in streams in eastern North America are usually mixtures of two or more species. Litter mixtures may decay more quickly than either of the component species. If so, estimates of stream energy and nutrient budgets may be inaccurate. In northern Nova Scotia, Canada, we measured mass loss from binary mixtures (1:1 mass ratio) of leaf litter in mesh bags, using freshly fallen or air-dried litter from five species of canopy trees. We repeated the experiment eight times, in summer and fall, in two streams and a small river, over 3 years. In some trials we enumerated benthic invertebrate and fungal colonization of decaying litter. Although there were marked differences in mass loss rates among litter types, decomposition was accelerated in mixtures relative to the mean of the component species in only three of eight trials, and only in mixtures containing N-rich speckled alder leaves. Mixing yellow birch and red maple leaves inhibited decomposition. Diversity (Shannon–Weaver Index), species richness, and abundance of aquatic hyphomycete fungi, as indexed by conidial production, were never greater (and sometimes less) on litter mixtures than on the component species. Total numbers, taxonomic richness and diversity of benthic invertebrates generally, and litter-feeding species in particular, were not augmented by mixing litter types. Litter mixtures appear to dilute a preferred substrate with patches of a less preferred substrate. Our results provide only weak support for the contention that combining two litter types leads to acceleration of decomposition rates. Handling editor: K. Martens     

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.     

Composition of riparian litter input regulates organic matter decomposition: Implications for headwater stream functioning in a managed forest landscape

Although the importance of stream condition for leaf litter decomposition has been extensively studied, little is known about how processing rates change in response to altered riparian vegetation community composition. We investigated patterns of plant litter input and decomposition across 20 boreal headwater streams that varied in proportions of riparian deciduous and coniferous trees. We measured a suite of in‐stream physical and chemical characteristics, as well as the amount and type of litter inputs from riparian vegetation, and related these to decomposition rates of native (alder, birch, and spruce) and introduced (lodgepole pine) litter species incubated in coarse‐ and fine‐mesh bags. Total litter inputs ranged more than fivefold among sites and increased with the proportion of deciduous vegetation in the riparian zone. In line with differences in initial litter quality, mean decomposition rate was highest for alder, followed by birch, spruce, and lodgepole pine (12, 55, and 68% lower rates, respectively). Further, these rates were greater in coarse‐mesh bags that allow colonization by macroinvertebrates. Variance in decomposition rate among sites for different species was best explained by different sets of environmental conditions, but litter‐input composition (i.e., quality) was overall highly important. On average, native litter decomposed faster in sites with higher‐quality litter input and (with the exception of spruce) higher concentrations of dissolved nutrients and open canopies. By contrast, lodgepole pine decomposed more rapidly in sites receiving lower‐quality litter inputs. Birch litter decomposition rate in coarse‐mesh bags was best predicted by the same environmental variables as in fine‐mesh bags, with additional positive influences of macroinvertebrate species richness. Hence, to facilitate energy turnover in boreal headwaters, forest management with focus on conifer production should aim at increasing the presence of native deciduous trees along streams, as they promote conditions that favor higher decomposition rates of terrestrial plant litter.     

Comparison of leaf decomposition and macroinvertebrate colonization between exotic and native trees in a freshwater ecosystem

One of the most important sources of energy in aquatic ecosystems is the allochthonous input of detritus. Replacement of native tree species by exotic ones affects the quality of detritus entering freshwater ecosystems. This replacement can alter nutrient cycles and community structure in aquatic ecosystems. The aims of our study were (1) to compare leaf litter decomposition of two widely distributed exotic species (Ailanthus altissima and Robinia pseudoacacia) with the native species they coexist with (Ulmus minor and Fraxinus angustifolia), and (2) to compare macroinvertebrate colonization among litters of the invasive and native species. Litter bags of the four tree species were placed in the water and collected every 2, 25, 39, 71, and 95 days in a lentic ecosystem. Additionally, the macroinvertebrate community on litter bags was monitored after 25, 39, and 95 days. Several leaf chemistry traits were measured at the beginning (% lignin; lignin:N, C:N, LMA) and during the study (leaf total nitrogen). We detected variable rates of decomposition among species (k values of 0.009, 0.008, 0.008, and 0.005 for F. angustifolia, U. minor, A. altissima and R. pseudoacacia, respectively), but we did not detect an effect of litter source (from native/exotic). In spite of its low decay, the highest leaf nitrogen was found in R. pseudoacacia litter. Macroinvertebrate communities colonizing litter bags were similar across species. Most of them were collectors (i.e., they feed on fine particulate organic matter), suggesting that leaf material of either invasive or native trees was used as substrate both for the animals and for the organic matter they feed on. Our results suggest that the replacement of the native Fraxinus by Robinia would imply a reduction in the rate of leaf processing and also a slower release of leaf nitrogen to water.     

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.     

Effects of leaf litter species on macroinvertebrate community properties and mosquito yield in Neotropical tree hole microcosms

Detritus quality and quantity affect macroinvertebrate productivity and distribution in many freshwater ecosystems. This study experimentally investigated the effects of leaf litter from Ceiba pentandra, Dipteryx panamensis, Ficus yoponensis, and Platypodium elegans on macroinvertebrate species composition, richness, and abundance in artificial water-filled tree holes in a lowland moist forest of Panama. Species composition was similar among treatments, but species richness and longevity differed among litter types and were consistently highest with Platypodium litter. Similar patterns were observed in natural tree holes of the focal tree species. The mosquito Culex mollis was the most abundant species in the field experiment. Average conductivity and dissolved oxygen concentration differed among leaf species, but pH did not. Leaf toughness was positively correlated with mean macroinvertebrate abundance and cumulative species richness. A laboratory experiment measured C. mollis yield and pupation time in tree hole microcosms containing the four litter species. Cumulative mosquito mass and time to pupation differed among leaf litter species, with Platypodium litter supporting the greatest yield. Pupation was slowest on Ceiba litter. Grazing by mosquito larvae facilitated leaf decomposition in all treatments. Results suggest that differences in macroinvertebrate species richness and mosquito yield can be attributed to differences in nutritional quality among litter species. Received: 14 October 1998 / Accepted: 21 February 1999     

Non-additive, identity-dependent effects of detrital species mixing on soft-sediment communities

Accelerating rates of species extinction and invasion have sparked recent interest in how changes in plant community composition can be propagated through food webs. Research in this area has, however, been largely restricted to considerations of how detrital species mixing affects litter decay processes. The consequences of changing detrital resources for whole assemblages of sediment‐dwelling invertebrates remain largely unknown. We manipulated the availability of three detrital sources, Avicennia marina leaves, Posidonia australis blades and Sargassum sp. thalli, on an Australian mudflat to test hypotheses about how changes in the type and number of macrophytes contributing to detrital resources might impact benthic invertebrate assemblages of estuarine soft‐sediments. By controlling for changes in total detrital biomass and ensuring that each detrital source was present in two‐ and three‐species mixes as well as monocultures, our experimental design was able to distinguish among effects of mixing, identity and biomass. Three months after detrital manipulation, macroinvertebrate abundance and species richness differed among treatments according to the biomass of detritus added and non‐additive effects of detrital species mixing. Whereas the mixing of two detrital species generally had an antagonistic effect on macroinvertebrate abundance and richness, faunal assemblages did not appreciably differ between three‐species mixes and monocultures. Generally negative effects of two‐species mixes on macroinvertebrates were opposed by positive effects on microphytobenthos, an important food‐source for many of the animals. Non‐additive effects on sediment communities were particularly apparent when Sargassum sp., the most labile of the three detrital sources considered, was included in two‐species mixes. This demonstration of non‐additive and identity‐dependent effects of detrital species mixing on soft‐sediment communities suggests that predicted compositional changes to aquatic macrophyte communities, resulting from coastal development and climate change, will flow on to effect other components of the estuarine food‐web.     

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.     

Grasping the heterogeneity of kettle hole water quality in Northeast Germany

Riparian vegetation typically provides substantial allochthonous material to aquatic ecosystems where micro-organisms can play an important role in organic matter degradation which can support consumer biomass. We examined the effects of leaf litter quality (e.g., leaf nutrients, lignin and cellulose content), leaf species mixing, and microbial community diversity on in-stream breakdown rates of litter from dominant riparian trees (Melaleuca argentea, M. leucadendra, and Nauclea orientalis) in both a perennial and intermittent river in Australia’s wet-dry tropics. Leaf mass remaining after 82 days of in-stream incubation was negatively correlated (P < 0.05) with initial leaf N and P content while initial lignin and cellulose content had no statistically significant effect. Breakdown rates of incubated leaves of both Melaleuca and Nauclea were significantly higher in mixed litter bags compared with single species litter bags. Although it was expected that leaf N content would decrease from initial levels during decomposition, we found either similar or slightly higher N content following in-stream incubation suggesting microbial colonisation increased overall N content. Stable isotopes of δ¹³C and δ¹⁵N for the major sources and consumers in both rivers provide evidence that leaf litter was an important macroinvertebrate food source in the perennial river where heavy shading may limit algal production. However, in the intermittent river where riparian cover was low, benthic algae were the major organic carbon source for consumers. Our findings suggest that riparian tree species influence rates of in-stream organic matter processing, microbial community composition, and aquatic food web dynamics in tropical wet-dry streams.     

Effects of organic matter and season on leaf litter colonisation and breakdown in cave streams

1. Low organic matter availability is thought to be a primary factor influencing evolutionary and ecological processes in cave ecosystems. We examined links among organic matter abundance, macroinvertebrate community structure and breakdown rates of red maple (Acer rubrum) and corn litter (Zea mays) in coarse‐ (10 × 8 mm) and fine‐mesh (500‐μm) litter bags over two seasonal periods in four cave streams in the south‐eastern U.S.A. 2. Organic matter abundance differed among cave streams, averaging from near zero to 850 g ash‐free dry mass m^?2. Each cave system harboured a different macroinvertebrate community. However, trophic structure was similar among caves, with low shredder biomass (2–17% of total biomass). 3. Corn litter breakdown rates (mean k = 0.005 day^?1) were faster than red maple (mean k = 0.003 day^?1). Breakdown rates in coarse‐mesh bags (k = 0.001–0.012 day^?1) were up to three times faster than in fine‐mesh bags (k = 0.001–0.004 day^?1). Neither invertebrate biomass in litter bags nor breakdown rates were correlated with the ambient abundance of organic matter. Litter breakdown rates showed no significant temporal variation. Epigean (surface‐adapted) invertebrates dominated biomass in litter bags, suggesting that their effects on cave ecosystem processes may be greater than hypogean (cave‐adapted) taxa, the traditional focus of cave studies. 4. The functional diversity of our cave communities and litter breakdown rates are comparable to those found in previous litter breakdown studies in cave streams, suggesting that the factors that control organic matter processing (e.g. trophic structure of communities) may be broadly similar across geographically diverse areas.     

Mixed leaf litter effects on decomposition rates and soil microarthropod communities in an oak–pine stand in Japan

Rates of decomposition, and soil faunal abundance and diversity associated with single-species and mixed-species litters were studied in a litter bag experiment in an oak–pine forest. We used two canopy species of leaf litter, pine and oak, and one shrub species, Sasa, and compared decomposition rates, and soil microarthropod abundance and community structure of oribatid mites in the litter bags. Mass loss of single species decreased in the order: oak > pine > Sasa. While the total mass loss rates of mixed litter were intermediate between those of the constituent species, enhancement of mass loss from the three-species mixture and from mixed slow-decomposing litters (pine and Sasa) was observed. Faunal abundance in litter bags was higher in mixed-species litter than in those with single-species litter, and species richness of oribatid mites was also higher in the three-species mixed litter. Faunal abundance in single-species litter bags was not correlated with mass loss, although enhancement of mass loss in mixed litter bags corresponded with higher microarthropod abundance. Habitat heterogeneity in mixed litter bags seemed to be responsible for the more abundant soil microarthropod community.     

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.     

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.