Predicting richness effects on ecosystem function in natural communities: insights from high-elevation streams.

. Despite the increased complexity of experimental and theoretical studies on the biodiversity-ecosystem functioning (B-EF) relationship, a major challenge is to demonstrate whether the observed importance of biodiversity in controlled experimental systems also persists in nature. Due to their structural simplicity and their low levels of human impacts, extreme species-poor ecosystems may provide new insights into B-EF relationships in natural systems. We address this issue using shredder invertebrate communities and organic matter decomposition rates in 24 high-altitude (3200-3900 m) Neotropical streams as a study model. We first assessed the effects of stream characteristics and shredder diversity and abundance on organic matter decomposition rates in coarse- and fine-mesh bags. We found the interaction term shredder richness x shredder abundance had the most significant impact on decomposition rates in the field, although water discharge may also play a role locally. We also examined the relative contribution of the three most abundant shredders on decomposition rates by manipulating shredder richness and community composition in a field experiment. Transgressive overyielding was detected among the three shredder species, indicating complementary resource use and/or facilitation. By integrating survey and experimental data in surface response analyses we found that observed B-EF patterns fit those predicted by a linear model that described litter decomposition rates as a function of increasing shredder richness and the relative abundance of the most efficient shredders. Finally, the validity of our approach was tested in a broader context by using two independent but comparable data sets from 49 French and Swedish streams showing more complex shredder community structure. Results revealed that richness and identity effects on decomposition rates were lost with increasing shredder community complexity. Our approach of combining experimental and empirical data with modeling in species-poor ecosystems may serve as an impetus for new B-EF studies. If theory can explain B-EF in low-diversity ecosystems, it may also have credibility in more complex ones.

Global patterns and drivers of ecosystem functioning in rivers and riparian zones.

River ecosystems receive and process vast quantities of terrestrial organic carbon, the fate of which depends strongly on microbial activity. Variation in and controls of processing rates, however, are poorly characterized at the global scale. In response, we used a peer-sourced research network and a highly standardized carbon processing assay to conduct a global-scale field experiment in greater than 1000 river and riparian sites. We found that Earth's biomes have distinct carbon processing signatures. Slow processing is evident across latitudes, whereas rapid rates are restricted to lower latitudes. Both the mean rate and variability decline with latitude, suggesting temperature constraints toward the poles and greater roles for other environmental drivers (e.g., nutrient loading) toward the equator. These results and data set the stage for unprecedented "next-generation biomonitoring" by establishing baselines to help quantify environmental impacts to the functioning of ecosystems at a global scale.

Ecological responses to experimental glacier-runoff reduction in alpine rivers.

Glacier retreat is a worldwide phenomenon with important consequences for the hydrological cycle and downstream ecosystem structure and functioning. To determine the effects of glacier retreat on aquatic communities, we conducted a 4-year flow manipulation in a tropical glacier-fed stream. Compared with an adjacent reference stream, meltwater flow reduction induces significant changes in benthic fauna community composition in less than 2 weeks. Also, both algal and herbivore biomass significantly increase in the manipulated stream as a response to flow reduction. After the flow reduction ceased, the system requires 14-16 months to return to its pre-perturbation state. These results are supported by a multi-stream survey of sites varying in glacial influence, showing an abrupt increase in algal and herbivore biomass below 11% glacier cover in the catchment. This study shows that flow reduction strongly affects glacier-fed stream biota, prefiguring profound ecological effects of ongoing glacier retreat on aquatic systems.

Invertebrate Metacommunity Structure and Dynamics in an Andean Glacial Stream Network Facing Climate Change.

Under the ongoing climate change, understanding the mechanisms structuring the spatial distribution of aquatic species in glacial stream networks is of critical importance to predict the response of aquatic biodiversity in the face of glacier melting. In this study, we propose to use metacommunity theory as a conceptual framework to better understand how river network structure influences the spatial organization of aquatic communities in glacierized catchments. At 51 stream sites in an Andean glacierized catchment (Ecuador), we sampled benthic macroinvertebrates, measured physico-chemical and food resource conditions, and calculated geographical, altitudinal and glaciality distances among all sites. Using partial redundancy analysis, we partitioned community variation to evaluate the relative strength of environmental conditions (e.g., glaciality, food resource) vs. spatial processes (e.g., overland, watercourse, and downstream directional dispersal) in organizing the aquatic metacommunity. Results revealed that both environmental and spatial variables significantly explained community variation among sites. Among all environmental variables, the glacial influence component best explained community variation. Overland spatial variables based on geographical and altitudinal distances significantly affected community variation. Watercourse spatial variables based on glaciality distances had a unique significant effect on community variation. Within alpine catchment, glacial meltwater affects macroinvertebrate metacommunity structure in many ways. Indeed, the harsh environmental conditions characterizing glacial influence not only constitute the primary environmental filter but also, limit water-borne macroinvertebrate dispersal. Therefore, glacier runoff acts as an aquatic dispersal barrier, isolating species in headwater streams, and preventing non-adapted species to colonize throughout the entire stream network. Under a scenario of glacier runoff decrease, we expect a reduction in both environmental filtering and dispersal limitation, inducing a taxonomic homogenization of the aquatic fauna in glacierized catchments as well as the extinction of specialized species in headwater groundwater and glacier-fed streams, and consequently an irreversible reduction in regional diversity.

Diversity patterns of aquatic macroinvertebrates in a tropical high-Andean catchment

Introduction: Glacierised catchments are remote and hostile environments, in which streams from different water sources (e.g., glacier melt, rain/snowmelt, groundwater) converge, creating a complex mosaic of stream sites with varying levels of glacial influence and environmental conditions. This environmental heterogeneity, in turn, influences the assemblage and composition of aquatic communities and produces complex patterns of species diversity at the catchment scale. Objectives: In this contribution, we assessed biodiversity and community composition of aquatic macroinvertebrate communities from 51 stream site types in a glacierised catchment in the tropical Andes. The aim of our study was to: (1) determine diversity, rarity, commonness and spatial distribution patterns of aquatic macroinvertebrate communities from sites with different water sources, and (2) identify which environmental variables influence the density and presence of macroinvertebrate taxa and, in particular, of the subfamilies of the ubiquitous chironomids. Methods: Our study sites were grouped according to their water source and to their percentage of glacier coverage in the catchment (GCC). At each site we sampled aquatic macroinvertebrates, measured environmental variables and assessed community differences and environmental influence with ordination analyses and generalized linear models. Results: Kryal and mixed sites had an important proportion of rare taxa. Mean richness was highest in the mixed sites and lowest in the sites with the highest glacier cover; while sites with an intermediate percentage of glacier cover, had the highest values of α and β diversity. We found that 13 taxa (15.9%) were common to all stream site types. SIMPER analysis showed that Orthocladiinae, Hyalella sp. and Andesiops sp. contributed the most to the dissimilarity between site types (˃ 45% of cumulative contribution). RDA showed that kryal sites were associated with high turbidity and density of Podonomids, and with low temperature, amount of CPOM and densities of both Anomalocosmoecus sp. and Andesiops sp. Orthocladinae was associated with high current velocity and chlorophyll a concentration, whereas Hyalella sp. had a positive relationship with higher pH and streambed stability. Generalized linear models showed that GCC was the main variable explaining all faunal metrics. Current velocity explained macroinvertebrate abundance, water temperature was related to chironomid density and chlorophyll a influenced Orthocladiinae presence-absence. Conclusions: Our results suggest that by favoring the presence of rare taxa and taxa turnover, glacier influence may increase biodiversity in glacierised catchments. In terms of biodiversity conservation, this study confirms an urgent need to increase knowledge of high-Andean stream biodiversity, especially in highly heterogenous glacierised catchments, to better describe regional biodiversity patterns and community composition of these highly vulnerable freshwater ecosystems. Detailed analyses of benthic communities and development of databases are key for conservation strategies. Water management municipalities and/or enterprises should consider water quality and stream types for more sustainable management of these important ecosystems.

Introducción: Las cuencas glaciares son entornos remotos y hostiles, en los que los arroyos de diferentes fuentes de agua (p.ej., deshielo de glaciares, lluvia/deshielo, agua subterránea) convergen, creando un mosaico complejo de tramos con diferentes niveles de influencia glacial y condiciones ambientales. Esta heterogeneidad ambiental influye, a su vez, en el ensamblaje y composición de las comunidades acuáticas y produce complejos patrones de diversidad a la escala de la cuenca. Objetivos: En esta contribución, evaluamos la biodiversidad y composición de comunidades de macroinvertebrados acuáticos en 51 sitios de una cuenca glaciar en los Andes tropicales. Los objetivos de nuestro estudio fueron: (1) determinar la diversidad, la contribución de taxones raros y comunes y los patrones de distribución espacial de las comunidades de macroinvertebrados acuáticos en sitios con diferentes fuentes de agua, y (2) identificar qué variables ambientales influyen en la densidad y presencia de taxones de macroinvertebrados y, en particular, de las subfamilias de los omnipresentes quironómidos. Métodos: Agrupamos a nuestros sitios de estudio según su fuente de agua y su porcentaje de cobertura de glaciar en la cuenca (GCC). En cada sitio donde muestreamos macroinvertebrados acuáticos, medimos variables ambientales y evaluamos las diferencias entre comunidades y la influencia ambiental con análisis de ordenación y modelos lineales generalizados. Resultados: Los sitios kryal y mixtos tuvieron una proporción importante de taxones raros. La riqueza media fue más alta en los sitios mixtos y más baja en los sitios con mayor cobertura glaciar; mientras que los sitios con un porcentaje intermedio de cobertura glaciar tuvieron los valores más altos de diversidad α y β. Encontramos que 13 taxones (15,9%) fueron comunes a todos los tipos de sitios de estudio. El análisis SIMPER mostró que Orthocladiinae, Hyalella sp. y Andesiops sp. contribuyeron más a la disimilitud entre tipos de sitios (˃ 45% de la contribución acumulada). El RDA mostró que los sitios kryal estaban asociados con alta turbidez y densidad de podonómidos, y con baja temperatura, cantidad de CPOM y densidad de Anomalocosmoecus sp. y Andesiops sp. Orthocladinae se asoció con una alta velocidad de corriente y concentración de clorofila a, mientras que Hyalella sp. tuvo una relación positiva con pH más alto y estabilidad del lecho del río. Los modelos lineales generalizados mostraron que GCC fue la variable principal para explicar todas las métricas de fauna. La velocidad de corriente explicó la abundancia de macroinvertebrados, la temperatura del agua estuvo relacionada con la densidad de los quironómidos y la clorofila influenció la presencia-ausencia de Orthocladiinae. Conclusiones: Nuestros resultados sugieren que, al favorecer la presencia de taxones raros y la rotación de taxones, la influencia de los glaciares puede aumentar la biodiversidad en cuencas con influencia glaciar. En términos de conservación de la biodiversidad, este estudio confirma la necesidad urgente de incrementar el conocimiento de la biodiversidad en arroyos de la región altoandina, especialmente en cuencas glaciares altamente heterogéneas, para describir mejor los patrones de biodiversidad regional y la composición de las comunidades en estos ecosistemas altamente vulnerables. Análisis detallados de las comunidades bentónicas y el desarrollo de bases de datos son claves para diseñar estrategias de conservación. Los municipios y/o empresas administradoras de agua deben considerar la calidad del agua y los tipos de arroyos para una gestión más sostenible de estos importantes ecosistemas.