Climate change is poised to alter mountain stream ecosystem processes via organismal phenological shifts.

Climate change is affecting the phenology of organisms and ecosystem processes across a wide range of environments. However, the links between organismal and ecosystem process change in complex communities remain uncertain. In snow-dominated watersheds, snowmelt in the spring and early summer, followed by a long low-flow period, characterizes the natural flow regime of streams and rivers. Here, we examined how earlier snowmelt will alter the phenology of mountain stream organisms and ecosystem processes via an outdoor mesocosm experiment in stream channels in the Eastern Sierra Nevada, California. The low-flow treatment, simulating a 3- to 6-wk earlier return to summer baseflow conditions projected under climate change scenarios in the region, increased water temperature and reduced biofilm production to respiration ratios by 32%. Additionally, most of the invertebrate species explaining community change (56% and 67% of the benthic and emergent taxa, respectively), changed in phenology as a consequence of the low-flow treatment. Further, emergent flux pulses of the dominant insect group (Chironomidae) almost doubled in magnitude, benefitting a generalist riparian predator. Changes in both invertebrate community structure (composition) and functioning (production) were mostly fine-scale, and response diversity at the community level stabilized seasonally aggregated responses. Our study illustrates how climate change in vulnerable mountain streams at the rain-to-snow transition is poised to alter the dynamics of stream food webs via fine-scale changes in phenology-leading to novel predator-prey "matches" or "mismatches" even when community structure and ecosystem processes appear stable at the annual scale.

Female proportion has a stronger influence on dispersal than body size in nematodes of mountain lakes.

Nematodes disperse passively and are amongst the smallest invertebrates on Earth. Free-living nematodes in mountain lakes are highly tolerant of environmental variations and are thus excellent model organisms in dispersal studies, since species-environment relationships are unlikely to interfere. In this study, we investigated how population or organism traits influence the stochastic physical nature of passive dispersal in a topologically complex environment. Specifically, we analyzed the influence of female proportion and body size on the geographical distribution of nematode species in the mountain lakes of the Pyrenees. We hypothesized that dispersal is facilitated by (i) a smaller body size, which would increase the rate of wind transport, and (ii) a higher female proportion within a population, which could increase colonization success because many nematode species are capable of parthenogenetic reproduction. The results showed that nematode species with a low proportion of females tend to have clustered spatial distributions that are not associated with patchy environmental conditions, suggesting greater barriers to dispersal. When all species were pooled, the overall proportion of females tended to increase at the highest elevations, where dispersal between lakes is arguably more difficult. The influence of body size was barely relevant for nematode distributions. Our study highlights the relevance of female proportion as a mechanism that enhances the dispersal success of parthenogenetic species, and that female sex is a determining factor in metacommunity connectivity.

Isolation controls reestablishment mechanisms and post-drying community structure in an intermittent stream.

Biota in disturbance-prone landscapes have evolved a variety of strategies to persist long term, either locally (resistance) or by regional recolonization (resilience). Habitat fragmentation and isolation can limit the availability of recolonization pathways, and thus the dynamics of post-disturbance community reestablishment. However, empirical studies on how isolation may control the mechanisms that enable community recovery remain scarce. Here, we studied a pristine intermittent stream (Chalone Creek, Pinnacles National Park, California) to understand how isolation (distance from a perennial pool) alters invertebrate community recolonization after drying. We monitored benthic invertebrate reestablishment during the rewetting phase along a ~2-km gradient of isolation, using mesh traps that selected for specific recolonization pathways (i.e., drift, flying, swimming/crawling, and vertical migration from the hyporheic). We collected daily emigration samples, surveyed the reestablished benthic community after 6 weeks, and compared assemblages across trap types and sites. We found that isolation mediated migration dynamics by delaying peak vertical migration from the hyporheic by ca. 1 day on average per 250 m of dry streambed. The relative importance of reestablishment mechanisms varied longitudinally-with more resistance strategists (up to 99.3% of encountered individuals) in the upstream reaches, and increased drift and aerial dispersers in the more fragmented habitats (up to 17.2% and 18%, respectively). Resistance strategists persisting in the hyporheic dominated overall (88.2% of individuals, ranging 52.9%-99.3% across sites), but notably most of these organisms subsequently outmigrated downstream (85.6% on average, ranging 52.1%-96% across sites). Thus, contrary to conventional wisdom, resistance strategists largely contributed to downstream resilience as well as to local community recovery. Finally, increased isolation was associated with a general decrease in benthic invertebrate diversity, and up to a 3-fold increase in the relative abundance of drought-resistant stoneflies. Our results advance the notion that understanding spatial context is key to predicting post-disturbance community dynamics. Considering the interaction between disturbance and fragmentation may help inform conservation in ecosystems that are subject to novel environmental regimes.

Nematode distributions as spatial null models for macroinvertebrate species richness across environmental gradients: A case from mountain lakes.

Nematode species are widely tolerant of environmental conditions and disperse passively. Therefore, the species richness distribution in this group might largely depend on the topological distribution of the habitats and main aerial and aquatic dispersal pathways connecting them. If so, the nematode species richness distributions may serve as null models for evaluating that of other groups more affected by environmental gradients. We investigated this hypothesis in lakes across an altitudinal gradient in the Pyrenees. We compared the altitudinal distribution, environmental tolerance, and species richness, of nematodes with that of three other invertebrate groups collected during the same sampling: oligochaetes, chironomids, and nonchironomid insects. We tested the altitudinal bias in distributions with t-tests and the significance of narrow-ranging altitudinal distributions with randomizations. We compared results between groups with Fisher's exact tests. We then explored the influence of environmental factors on species assemblages in all groups with redundancy analysis (RDA), using 28 environmental variables. And, finally, we analyzed species richness patterns across altitude with simple linear and quadratic regressions. Nematode species were rarely biased from random distributions (5% of species) in contrast with other groups (35%, 47%, and 50%, respectively). The altitudinal bias most often shifted toward low altitudes (85% of biased species). Nematodes showed a lower portion of narrow-ranging species than any other group, and differed significantly from nonchironomid insects (10% and 43%, respectively). Environmental variables barely explained nematode assemblages (RDA adjusted R2 = 0.02), in contrast with other groups (0.13, 0.19 and 0.24). Despite these substantial differences in the response to environmental factors, species richness across altitude was unimodal, peaking at mid elevations, in all groups. This similarity indicates that the spatial distribution of lakes across altitude is a primary driver of invertebrate richness. Provided that nematodes are ubiquitous, their distribution offers potential null models to investigate species richness across environmental gradients in other ecosystem types and biogeographic regions.

Environmental factors prevail over dispersal constraints in determining the distribution and assembly of Trichoptera species in mountain lakes.

Aiming to elucidate whether large-scale dispersal factors or environmental species sorting prevail in determining patterns of Trichoptera species composition in mountain lakes, we analyzed the distribution and assembly of the most common Trichoptera (Plectrocnemia laetabilis, Polycentropus flavomaculatus, Drusus rectus, Annitella pyrenaea, and Mystacides azurea) in the mountain lakes of the Pyrenees (Spain, France, Andorra) based on a survey of 82 lakes covering the geographical and environmental extremes of the lake district. Spatial autocorrelation in species composition was determined using Moran's eigenvector maps (MEM). Redundancy analysis (RDA) was applied to explore the influence of MEM variables and in-lake, and catchment environmental variables on Trichoptera assemblages. Variance partitioning analysis (partial RDA) revealed the fraction of species composition variation that could be attributed uniquely to either environmental variability or MEM variables. Finally, the distribution of individual species was analyzed in relation to specific environmental factors using binomial generalized linear models (GLM). Trichoptera assemblages showed spatial structure. However, the most relevant environmental variables in the RDA (i.e., temperature and woody vegetation in-lake catchments) were also related with spatial variables (i.e., altitude and longitude). Partial RDA revealed that the fraction of variation in species composition that was uniquely explained by environmental variability was larger than that uniquely explained by MEM variables. GLM results showed that the distribution of species with longitudinal bias is related to specific environmental factors with geographical trend. The environmental dependence found agrees with the particular traits of each species. We conclude that Trichoptera species distribution and composition in the lakes of the Pyrenees are governed predominantly by local environmental factors, rather than by dispersal constraints. For boreal lakes, with similar environmental conditions, a strong role of dispersal capacity has been suggested. Further investigation should address the role of spatial scaling, namely absolute geographical distances constraining dispersal and steepness of environmental gradients at short distances.

Pollutant dehalogenation capability may depend on the trophic evolutionary history of the organism: PBDEs in freshwater food webs.

Organohalogen compounds are some of the most notorious persistent pollutants disturbing the Earth biosphere. Although human-made, these chemicals are not completely alien to living systems. A large number of natural organohalogens, part of the secondary metabolism, are involved in chemical trophic interactions. Surprisingly, the relationship between organisms' trophic position and synthetic organohalogen biotransformation capability has not been investigated. We studied the case for polybromodiphenyl ethers (PBDE), a group of flame-retardants of widespread use in the recent years, in aquatic food webs from remote mountain lakes. These relatively simple ecosystems only receive pollution by atmospheric transport. A large predominance of the PBDE congener currently in use in Europe, BDE-209, largely dominated the PBDE composition of the basal resources of the food web. In contrast, primary consumers (herbivores and detritivores) showed a low proportion of BDE-209, and dominance of several less brominated congeners (e.g. BDE-100, BDE47). Secondary consumers (predators) showed large biomagnification of BDE-209 compare to other congeners. Finally, top predator fish characterized by low total PBDE concentrations. Examination of the bromine stable isotopic composition indicates that primary consumers showed higher PBDE biotransformation capability than secondary consumers. We suggest that the evolutionary response of primary consumers to feeding deterrents would have pre-adapted them for PBDE biotransformation. The observed few exceptions, some insect taxa, can be interpreted in the light of the trophic history of the evolutionary lineage of the organisms. Bromine isotopic composition in fish indicates that low PBDE values are due to not only biotransformation but also to some other process likely related to transport. Our finding illustrates that organohalogen compounds may strongly disturb ecosystems even at low concentrations, since the species lacking or having scarce biotransformation capability may be selectively more exposed to these halogenated hydrophobic semi-volatile organic pollutants due to their high bioaccumulation potential.