Luz Boyero

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.

Are tropical streams really different

Stream shredders have been reported as scarce in several tropical areas. This pattern is in contrast to observations in temperate streams, which support an abundant and diverse shredder fauna. Two possible explanations for this pattern are that most shredders are adapted to cool conditions and that temperate riparian vegetation often produces more palatable and more nutritious leaves than do the more diverse, tropical rainforests. In peninsular Malaysia, most streams flow through lowland dipterocarp forests that are characterized by strikingly high tree diversity and by many species with tough leathery leaves that are high in lignin and toxic secondary compounds and low in protein. In contrast, highland streams flow through montane rain forests and are more similar to temperate streams. We hypothesized that shredder fauna would be distributed along an altitudinal gradient, with more abundant and diverse assemblages in highland streams. We sampled leaf litter in 12 sites at altitudes from 55 to 1560 m above sea level. As expected, highland sites supported higher abundance and diversity of shredders (9–15 species per site) than did lowland sites (3–8 species per site). Shredder densities were similar among lowland sites, but species composition was variable. Large snails (Brotia spp.) were the dominant shredders in nonacidic streams in Taman Negara, but they were absent from acidic streams and from streams elsewhere in peninsular Malaysia. Shredder biomass was generally high because of the large body size of most species (e.g., crabs, snails, semiaquatic cockroaches, calamoceratids, and tipulids). Large mouthparts might allow these species to shred tough leaves efficiently, and large body size might enable them to tolerate a wide variety of toxic secondary compounds. Our results suggest that shredder diversity might depend on elevation, water temperature, characteristics of the riparian vegetation along altitudinal gradients, and variation in water chemistry at local scales.

Shredders in Malaysia: abundance and richness are higher in cool upland tropical streams

Abstract Stream shredders have been reported as scarce in several tropical areas. This pattern is in contrast to observations in temperate streams, which support an abundant and diverse shredder fauna. Two possible explanations for this pattern are that most shredders are adapted to cool conditions and that temperate riparian vegetation often produces more palatable and more nutritious leaves than do the more diverse, tropical rainforests. In peninsular Malaysia, most streams flow through lowland dipterocarp forests that are characterized by strikingly high tree diversity and by many species with tough leathery leaves that are high in lignin and toxic secondary compounds and low in protein. In contrast, highland streams flow through montane rain forests and are more similar to temperate streams. We hypothesized that shredder fauna would be distributed along an altitudinal gradient, with more abundant and diverse assemblages in highland streams. We sampled leaf litter in 12 sites at altitudes from 55 to 1560 m above sea level. As expected, highland sites supported higher abundance and diversity of shredders (9–15 species per site) than did lowland sites (3–8 species per site). Shredder densities were similar among lowland sites, but species composition was variable. Large snails (Brotia spp.) were the dominant shredders in nonacidic streams in Taman Negara, but they were absent from acidic streams and from streams elsewhere in peninsular Malaysia. Shredder biomass was generally high because of the large body size of most species (e.g., crabs, snails, semiaquatic cockroaches, calamoceratids, and tipulids). Large mouthparts might allow these species to shred tough leaves efficiently, and large body size might enable them to tolerate a wide variety of toxic secondary compounds. Our results suggest that shredder diversity might depend on elevation, water temperature, characteristics of the riparian vegetation along altitudinal gradients, and variation in water chemistry at local scales.

Organization of macroinvertebrate communities at a hierarchy of spatial scales in a tropical stream

The variability of benthic macroinvertebrate (bmi) communities at three spatial scales was examined in the Juncal Stream, Coiba Island, Panama. Standard Surber samples were taken at riffle habitats of first, second and third order streams of the Juncal Stream watershed. In a nested design, we had three riffles within each stream order, and three sampling points within each riffle. Bmi total density and richness showed greater variation among stream orders and within riffles, while individual taxa varied mostly among and within riffles, and community evenness varied within riffles. Current velocity was strongly related to the variability of bmi community descriptors. Multidimensional scaling showed that first and third stream orders were more similar in bmi composition than second order. Riffles were more heterogeneous in composition within first order than within second or third orders, probably related to the greater substrate heterogeneity in first order riffles.

Multiscale patterns of spatial variation in stream macroinvertebrate communities

Our ability to detect patterns of variation of communities depends on the spatial scale of observation. I examined the spatial variation of macroinvertebrate community structure: abundance, richness, evenness, percentage of Ephemeroptera, Plecoptera and Trichoptera (EPT), and taxonomic composition across a wide range of spatial scales in two mountain streams. In a nested design, three segments were selected within each basin, three riffles within each segment, three sections within each riffle, and three samples within each section. Significant variation of communities occurred mainly at sample and riffle scales, although different community characteristics may vary at different scales. Environmental factors were strongly related to communities, but these relationships depended on spatial scale in many cases, suggesting that the influence of the environment is ultimately regulated by the grain and extent of organisms. This study highlights the importance of multiscale studies to obtain a complete understanding of the spatial variation of macroinvertebrate communities and their relationship with the environment.

Gradients in regional diversity of freshwater taxa

Abstract Opinion about latitudinal diversity gradients in freshwater fauna has varied over past decades. Global data have been compiled for diversity of Ephemeroptera, Plecoptera, and Trichoptera at the site scale, but no assessment of global variability in regional diversity has been done for any taxon except fish. Global variation cannot be inferred from site-scale data because the relationship between site and regional diversity is not necessarily linear. We have assembled global data on regional diversity for 7 freshwater taxa. Here, we plot diversity against regional area, develop diversity–area regressions, and then plot the residuals against latitude to investigate latitudinal diversity gradients. The existence and directions of gradients vary substantially among taxa. For Ephemeroptera and Plecoptera, significant latitudinal gradients exist, and diversity is greater at higher latitudes. For Trichoptera and Caudata, latitudinal gradients are not apparent, although taxa within the Caudata show distinct patterns. For Odonata, Osteichthyes, and Anura, latitudinal gradients are highly significant, and diversity is greatest at low latitudes. A clear distinction between the life cycles of tropically diverse taxa and of other taxa (excluding fish) is a long terrestrial phase in the tropically diverse taxa (e.g., Odonata). We discuss reasons for the gradients and this contrast in ecological and evolutionary contexts (e.g., habitat complexity and cross-habitat adaptation).

The quantification of local substrate heterogeneity in streams and its significance for macroinvertebrate assemblages

The effect of substrate heterogeneity on the structure of stream macroinvertebrate assemblages (total abundance, taxon richness, and evenness) is still not clear, but this could be due to the lack of standard methods for quantifying substrate heterogeneity. An accurate quantification of substrate heterogeneity was obtained from photographs of sampled areas (each 225 cm2), which were used to create maps that were subsequently digitized and analyzed using image analysis software. These maps allowed the calculation of multiple metrics quantifying two aspects of substrate heterogeneity: composition and spatial configuration of substrate patches. The diversity of substrate types (calculated as the Shannon diversity index), and the heterogeneity of patch compactness (calculated as the coefficient of variation of the relationship between patch dimensions) were the metrics explaining more biotic variance at the sample scale, but at higher scales there were no relationships between assemblage structure and substrate heterogeneity. Most variation in substrate heterogeneity occurred at the sample scale, while some metrics varied significantly at riffle or segment scales; these patterns of variation match those of macroinvertebrate assemblages, which had been previously studied. The importance of quantifying substrate heterogeneity and considering the spatial scales of its study are discussed.

Leaf litter diversity and shredder preferences in an Australian tropical rain-forest stream

Consumption of terrestrial leaf litter by stream invertebrates is an important process, but little attention has been paid to the influence of leaf diversity on the process. Tropical forests are known to have much greater diversity of plant species than their temperate counterparts, but studies of litter processing in tropical streams have not explicitly addressed this issue. This paper documents the streambed leaf litter composition and diversity of an Australian tropical rain-forest stream and the leaf preferences of consumers in the stream. We hypothesized that there would be high diversity of litter in the stream and that the shredders would have broad preferences, given that litterfall of any one species would occur over a restricted period. Leaf litter was characterized by high species diversity (81 species from one stream reach sampled on two occasions). Leaf consumers (‘shredders’) were associated with a relatively broad suite of leaf species (38 species) and did not indicate clear leaf preferences. However, in a laboratory feeding experiment, using the three most common shredder species and some of the most abundant leaf species in the stream, all shredder species exhibited clear preference for a single leaf species (Endiandra bessaphila). Preference for this and other species was affected by the conditioning age of leaves (i.e. the length of time leaves were exposed to leaching and microbial colonization), with conditioned leaves usually being preferred, and previously non-selected leaves becoming more palatable with conditioning. Thus, different successional stages were more important than the identity of leaf species in determining the distribution of shredders among the leaves.

How biological diversity influences ecosystem function: a test with a tropical stream detritivore guild

We investigated the relationship between diversity and ecosystem function, which is controversial and has rarely been examined for consumer assemblages, for the process of leaf breakdown by the shredder guild in a tropical stream. We manipulated species richness, evenness and identity of four macroinvertebrate shredder species (three caddisflies and one mayfly) in microcosms and tested their effect on leaf breakdown rates measured as leaf mass loss per capita and per milligram of animal. Species richness, evenness and species identity all affected leaf breakdown rates. Breakdown rates tended to increase with higher richness, but only for the three caddisflies, probably through a release of intraspecific interference, although other mechanisms such as niche complementarity or facilitation cannot be discarded. Leaf breakdown by the caddisflies was reduced in the presence of the mayfly, possibly because of its mode of movement by swimming instead of crawling and its similarity to some predators that are common in leaf litter. Species identity was more important than species richness in determining leaf breakdown rates, indicating that some species within the shredder guild are not redundant, and suggesting important consequences of particular species loss for the functioning of the ecosystem.