Marcelo S. Moretti

A global experiment suggests climate warming will not accelerate litter decomposition in streams but might reduce carbon sequestration

The decomposition of plant litter is one of the most important ecosystem processes in the biosphere and is particularly sensitive to climate warming. Aquatic ecosystems are well suited to studying warming effects on decomposition because the otherwise confounding influence of moisture is constant. By using a latitudinal temperature gradient in an unprecedented global experiment in streams, we found that climate warming will likely hasten microbial litter decomposition and produce an equivalent decline in detritivore-mediated decomposition rates. As a result, overall decomposition rates should remain unchanged. Nevertheless, the process would be profoundly altered, because the shift in importance from detritivores to microbes in warm climates would likely increase CO(2) production and decrease the generation and sequestration of recalcitrant organic particles. In view of recent estimates showing that inland waters are a significant component of the global carbon cycle, this implies consequences for global biogeochemistry and a possible positive climate feedback.

Global distribution of a key trophic guild contrasts with common latitudinal diversity patterns

Most hypotheses explaining the general gradient of higher diversity toward the equator are implicit or explicit about greater species packing in the tropics. However, global patterns of diversity within guilds, including trophic guilds (i.e., groups of organisms that use similar food resources), are poorly known. We explored global diversity patterns of a key trophic guild in stream ecosystems, the detritivore shredders. This was motivated by the fundamental ecological role of shredders as decomposers of leaf litter and by some records pointing to low shredder diversity and abundance in the tropics, which contrasts with diversity patterns of most major taxa for which broad-scale latitudinal patterns haven been examined. Given this evidence, we hypothesized that shredders are more abundant and diverse in temperate than in tropical streams, and that this pattern is related to the higher temperatures and lower availability of high-quality leaf litter in the tropics. Our comprehensive global survey (129 stream sites from 14 regions on six continents) corroborated the expected latitudinal pattern and showed that shredder distribution (abundance, diversity and assemblage composition) was explained by a combination of factors, including water temperature (some taxa were restricted to cool waters) and biogeography (some taxa were more diverse in particular biogeographic realms). In contrast to our hypothesis, shredder diversity was unrelated to leaf toughness, but it was inversely related to litter diversity. Our findings markedly contrast with global trends of diversity for most taxa, and with the general rule of higher consumer diversity at higher levels of resource diversity. Moreover, they highlight the emerging role of temperature in understanding global patterns of diversity, which is of great relevance in the face of projected global warming.

Leaf-litter breakdown in tropical streams: is variability the norm?

Many forested headwater streams are heterotrophic ecosystems in which allochthonous inputs of plant litter are a major source of energy. Leaves of riparian vegetation entering the stream are broken down by a combination of biotic and abiotic processes and, in most temperate and boreal streams, provide food and habitat for dense populations of detritivorous invertebrates. However, tropical streams in different parts of the world show substantial variability in the number and diversity of leaf-shredding detritivores (hereafter detritivores). We used data obtained with standardized methods from multiple streams in Africa, the Americas, Asia, and Australia to test the hypothesis that this variability would lead to differences in the relative role of detritivores and microorganisms in the breakdown process. We also tested the hypotheses that variability in litter breakdown rates changes with litter type (native litter mixtures vs nonnative alder [Alnus glutinosa]) and is higher across regions within than outside the tropics. We found that litter breakdown rates were highly variable across sites, with no consistent pattern within geographic areas, although litter consumption by detritivores was negligible at several sites, all in America. Geographic patterns of litter breakdown also varied between litter types, with higher breakdown rates for alder than for native litter in most but not all regions. When litter breakdown rates at the tropical sites were compared to previously reported values from temperate and boreal regions, we found that differences in variability between tropical and temperate sites were inconsistent, with great differences among studies. Further global-scale studies will be needed to assess the extent to which latitudinal changes in the diversity and composition of microbial and detritivore assemblages contribute to variability in litter breakdown rates.

Length–dry mass relationships for a typical shredder in Brazilian streams (Trichoptera: Calamoceratidae)

The aims of this study were to determine which linear body dimensions are best suitable and which mathematical functions can be used to describe length–dry mass relationships for a population of Phylloicus sp. (Trichoptera: Calamoceratidae) larvae. We measured three linear body dimensions (body length, head capsule width and interocular distance) of 54 larvae to use as dry mass predictors. For the description of length–dry mass relationships we used linear, exponential and power function models. Body length provided the best fitted equations to estimate biomass, followed by head capsule width and interocular distance. The highest coefficients of determination were found in power function and exponential models. These relationships can be useful to determine the growth rate and/or secondary production of Phylloicus larvae in future laboratory experiments, as well as to understand the importance of these shredders in the energy flux of shaded tropical streams.

Biotic and abiotic variables influencing plant litter breakdown in streams: A global study

Plant litter breakdown is a key ecological process in terrestrial and freshwater ecosystems. Streams and rivers, in particular, contribute substantially to global carbon fluxes. However, there is little information available on the relative roles of different drivers of plant litter breakdown in fresh waters, particularly at large scales. We present a global-scale study of litter breakdown in streams to compare the roles of biotic, climatic and other environmental factors on breakdown rates. We conducted an experiment in 24 streams encompassing latitudes from 47.8° N to 42.8° S, using litter mixtures of local species differing in quality and phylogenetic diversity (PD), and alder (Alnus glutinosa) to control for variation in litter traits. Our models revealed that breakdown of alder was driven by climate, with some influence of pH, whereas variation in breakdown of litter mixtures was explained mainly by litter quality and PD. Effects of litter quality and PD and stream pH were more positive at higher temperatures, indicating that different mechanisms may operate at different latitudes. These results reflect global variability caused by multiple factors, but unexplained variance points to the need for expanded global-scale comparisons.

No evidence for leaf-trait dissimilarity effects on litter decomposition, fungal decomposers, and nutrient dynamics

Biodiversity and ecosystem-functioning theory suggest that litter mixtures composed of dissimilar leaf species can enhance decomposition due to species trait complementarity. Here we created a continuous gradient of litter chemistry trait variability within species mixtures to assess effects of litter dissimilarity on three related processes in a natural stream: litter decomposition, fungal biomass accrual in the litter, and nitrogen and phosphorus immobilization. Litter from a pool of eight leaf species was analyzed for chemistry traits affecting decomposition (lignin, nitrogen, and phosphorus) and assembled in all of the 28 possible two-species combinations. Litter dissimilarity was characterized in terms of a range of trait-diversity measures, using Euclidean and Gower distances and dendrogram-based indices. We found large differences in decomposition rates among leaf species, but no significant relationships between decomposition rate of individual leaf species and litter trait dissimilarity, irrespective of whether decomposition was mediated by microbes alone or by both microbes and litter-consuming invertebrates. Likewise, no effects of trait dissimilarity emerged on either fungal biomass accrual or changes during decomposition of nitrogen or phosphorus concentrations in individual leaf species. In line with recent meta-analyses, these results provide support for the contention that litter diversity effects on decomposition, at least in streams, are less pronounced than effects on terrestrial primary productivity.

Leaf traits induce the feeding preference of a shredder of the genus Triplectides Kolenati, 1859 (Trichoptera) in an Atlantic Forest stream, Brazil: a test with native and exotic leaves

This study assessed the feeding preference of larvae of Triplectides sp. (Trichoptera, Leptoceridae) exposed to leaves of native (Hoffmannia dusenii Standley, 1931) and exotic (Eucalyptus globulus Labillardiere, 1799) trees. We hypothesized that, regardless of the origin of the leaves, larval preference is determined mainly by leaf anatomy and quality. Leaves from both species were conditioned with and without nutrient enrichment (NPK), and the four food items were offered in paired combinations to 162 larvae. Larval preference varied according to leaf combinations. In treatments containing both species, larvae preferred to feed on H. dusenii because of softer tissues and anatomical structure. The only exception was the treatment containing discs of enriched E. globulus and non-enriched H. dusenii where enhanced microbial activity on enriched leaves provided a softer resource to shredders. Our results corroborate the initial hypothesis and suggest that introduction of exotic leaves and changes in nutrient availability may affect shredder activity in streams and, consequently, organic-matter processing and ecosystem functioning.

Does Barypenthus concolor Burmeister (Trichoptera: Odontoceridae) select particles of different sizes for case building?

Many species of Trichoptera larvae use organic or inorganic particles to build cases that protect them from predators. The study of the wide variety of structures produced by trichopterans is important to understand the evolution of case building behavior. The objective of this work was to investigate if larvae of Barypenthus concolor (Burmeister) are capable of selecting particles of specific sizes to build their cases. The study site was an altitudinal stream in Serra do Cipo National Park, MG, Brazil). Cases and substrate samples were collected at three sites of the stream and submitted to granulometric analyses. The average mass of each particle size found in each site was used to determine the particle size proportion present in the substrate. The proportions of particle size of each case (site I: n = 10, site II: n = 19, site III: n = 13) were also analyzed. Possible differences between the proportions of the particle sizes of the substrate and of those used by B. concolor larvae were assessed by the Chi-square test. Medium and fine gravel predominated at the three sites (>70%). There were no significant differences between the expected and observed particle size proportion in the cases. Therefore, B. concolor does not select specific particle sizes for case building.

Exotic Eucalyptus leaves are preferred over tougher native species but affect the growth and survival of shredders in an Atlantic Forest stream (Brazil)

We evaluated the effect of leaves of native and exotic tree species on the feeding activity and performance of the larvae of Triplectides gracilis, a typical caddisfly shredder in Atlantic Forest streams. Leaves of four native species that differ in chemistry and toughness (Hoffmannia dusenii, Miconia chartacea, Myrcia lineata and Styrax pohlii) and the exotic Eucalyptus globulus were used to determine food preferences and rates of consumption, production of fine particulate organic matter (FPOM), growth and survival of shredders. We hypothesized that the consumption rates of leaves of Eucalyptus and their effects on the growth and survival of shredders could be predicted by leaf chemistry and toughness. The larvae preferred to feed on soft leaves (H. dusenii and M. chartacea) independently of the content of nutrients (N and P) and secondary compounds (total phenolics). When such leaves were absent, they preferred E. globulus and did not consume the tough leaves (M. lineata and S. pohlii). In monodietary experiments, leaf consumption and FPOM production differed among the studied leaves, and the values observed for the E. globulus treatments were intermediate between the soft and tough leaves. The larvae that fed on H. dusenii and M. chartacea grew constantly over five weeks, while those that fed on E. globulus lost biomass. Larval survival was higher on leaves of H. dusenii, M. chartacea and S. pohlii than on E. globulus and M. lineata leaves. Although E. globulus was preferred over tougher leaves, long-term consumption of leaves of the exotic species may affect the abundance of T. gracilis in the studied stream. Additionally, our results suggest that leaf toughness can be a determining factor for the behavior of shredders where low-quality leaves are abundant, as in several tropical streams.

Structural and functional composition of invertebrate communities associated with leaf patches in forest streams: a comparison between mesohabitats and catchments

We evaluated the influence of mesohabitats and catchments on the structural and functional composition of invertebrate communities associated with leaf patches in Atlantic Forest streams. We hypothesized that invertebrate communities would be more influenced by inter-habitat than inter-catchment variation, i.e., invertebrate taxonomic and trophic composition would differ more between pools and riffles than among catchments. Invertebrate richness differed among catchments only, while values of total abundance and biomass were higher in pools. The influence of mesohabitats on the structure of invertebrate communities was high, and most taxa had specificities with one mesohabitat. Among insect shredders, Grypopterigidae (Plecoptera) and Blattodea occurred more in riffles; Phylloicus, Nectopsyche, and Triplectides (Trichoptera) occurred more in pools. The biomass of omnivorous macroconsumers (Trichodactylus fluviatilis and Macrobrachium potiuna) was higher in pools and also differed among catchments. Except gathering-collectors, the taxonomic composition of functional feeding groups differed between pools and riffles. The obtained results corroborated the hypothesis that the structural and functional composition of invertebrate communities is more influenced by inter-habitat than by inter-catchment variation, and reinforced the importance of maintaining the biological and geomorphological characteristics of streams to allow the accumulation of leaves in different mesohabitats and preserve the aquatic biodiversity.