Microbial assemblages reflect environmental heterogeneity in alpine streams

Alpine streams are dynamic habitats harboring substantial biodiversity across small spatial extents. The diversity of alpine stream biota is largely reflective of environmental heterogeneity stemming from varying hydrological sources. Globally, alpine stream diversity is under threat as meltwater sources recede and stream conditions become increasingly homogeneous. Much attention has been devoted to macroinvertebrate diversity in alpine headwaters, yet to fully understand the breadth of climate change threats, a more thorough accounting of microbial diversity is needed. We characterized microbial diversity (specifically Bacteria and Archaea) of 13 streams in two disjunct Rocky Mountain subranges through 16S rRNA gene sequencing. Our study encompassed the spectrum of alpine stream sources (glaciers, snowfields, subterranean ice, and groundwater) and three microhabitats (ice, biofilms, and streamwater). We observed no difference in regional (γ) diversity between subranges but substantial differences in diversity among (β) stream types and microhabitats. Within‐stream (α) diversity was highest in groundwater‐fed springs, lowest in glacier‐fed streams, and positively correlated with water temperature for both streamwater and biofilm assemblages. We identified an underappreciated alpine stream type—the icy seep—that are fed by subterranean ice, exhibit cold temperatures (summer mean <2°C), moderate bed stability, and relatively high conductivity. Icy seeps will likely be important for combatting biodiversity losses as they contain similar microbial assemblages to streams fed by surface ice yet may be buffered against climate change by insulating debris cover. Our results show that the patterns of microbial diversity support an ominous trend for alpine stream biodiversity; as meltwater sources decline, stream communities will become more diverse locally, but regional diversity will be lost. Icy seeps, however, represent a source of optimism for the future of biodiversity in these imperiled ecosystems.     

滇西北高山微水体与溪流生境底栖动物多样性和环境特征

高山微水体由于面积微小且通过地表径流形成串联结构常常被认为与高山溪流具有类似的生境, 然而由于这两类生境中环境因子与底栖动物多样性存在差异, 它们在生态系统中的作用可能完全不同。滇西北地区是全球生物多样性热点区域之一, 境内高山微水体和高山溪流分布密集, 在区域底栖生物多样性维持方面具有重要的功能, 然而目前对这两类高山淡水生态系统的研究较少。为了比较这两类生境环境因子的异同及其对底栖动物多样性的维持作用, 2015年6月, 作者在云南省怒江州贡山县的高山峡谷内, 对27个高山微水体和同区域分布的1条高山溪流(海拔高差500 m范围)的底栖动物多样性和水环境因子进行了实地调查。结果表明: (1)高山微水体和高山溪流底栖动物群落中优势分类单元种群数量均比较庞大, 而稀有分类单元数量较多且种群较小; (2)两种生境在环境因子、物种多样性、功能多样性和群落结构方面的差异明显, 高山溪流有较高的物种丰富度、物种多样性和功能多样性; (3)高山微水体底栖动物多样性的分布与水环境因子无关, 而高山溪流底栖动物多样性与群落结构的形成受到与流速关联的水环境因子和海拔的影响。因此, 高山微水体与高山溪流不能简单地视为类似的生境类型, 它们对区域底栖动物多样性和生态功能维持可能具有不同的作用。     

Glacial loss and its effect on riparian vegetation of alpine streams

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Vulnerability of stream community composition and function to projected thermal warming and hydrologic change across ecoregions in the western United States

Shifts in biodiversity and ecological processes in stream ecosystems in response to rapid climate change will depend on how numerically and functionally dominant aquatic insect species respond to changes in stream temperature and hydrology. Across 253 minimally perturbed streams in eight ecoregions in the western USA, we modeled the distribution of 88 individual insect taxa in relation to existing combinations of maximum summer temperature, mean annual streamflow, and their interaction. We used a heat map approach along with downscaled general circulation model (GCM) projections of warming and streamflow change to estimate site‐specific extirpation likelihood for each taxon, allowing estimation of whole‐community change in streams across these ecoregions. Conservative climate change projections indicate a 30–40% loss of taxa in warmer, drier ecoregions and 10–20% loss in cooler, wetter ecoregions where taxa are relatively buffered from projected warming and hydrologic change. Differential vulnerability of taxa with key functional foraging roles in processing basal resources suggests that climate change has the potential to modify stream trophic structure and function (e.g., alter rates of detrital decomposition and algal consumption), particularly in warmer and drier ecoregions. We show that streamflow change is equally as important as warming in projected risk to stream community composition and that the relative threat posed by these two fundamental drivers varies across ecoregions according to projected gradients of temperature and hydrologic change. Results also suggest that direct human modification of streams through actions such as water abstraction is likely to further exacerbate loss of taxa and ecosystem alteration, especially in drying climates. Management actions to mitigate climate change impacts on stream ecosystems or to proactively adapt to them will require regional calibration, due to geographic variation in insect sensitivity and in exposure to projected thermal warming and hydrologic change.     

Parallels and contrasts between intermittently freezing and drying streams: From individual adaptations to biodiversity variation

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Vulnerability of alpine stream biodiversity to shrinking glaciers and snowpacks

Climate change poses a considerable threat to the biodiversity of high latitude and altitude ecosystems, with alpine regions across the world already showing responses to warming. However, despite probable hydrological change as alpine glaciers and snowpacks shrink, links between alpine stream biota and reduced meltwater input are virtually unknown. Using data from the French Pyrénées, we demonstrate that taxonomic richness and total abundance of stream macroinvertebrates increase significantly as meltwater (snow melt and glacier melt) contributions to river flow decrease. Macroinvertebrate species showed a gradation of optimum meltwater conditions at which they persist. For example: Habroleptoides berthelemyi (Ephemeroptera), Perla grandis (Plecoptera) and Rhithrogena spp. (Ephemeroptera) increased in abundance when meltwater contributions to streamflow decrease, whereas in contrast, Rhyacophila angelieri (Trichoptera) and Diamesa latitarsis spp. (Diptera) decreased in abundance. Changes in alpine stream macroinvertebrate community composition as meltwater contributions decline were associated with lower suspended sediment concentration, and higher water temperature, electrical conductivity and pH. Our results suggest α diversity (at a site) of streams presently fed by meltwaters will increase with future meltwater reductions. However, β diversity (between‐sites) will be reduced as snow melt and glacier melt decrease because the habitat heterogeneity associated with spatiotemporal variability of water source contributions will become lower as meltwater contributions decline. Extinction of some endemic alpine aquatic species (such as the Pyrenean caddis fly R. angelieri) is predicted with reduced meltwater inputs, leading to decreases in γ diversity (region). Our identification of significant links between meltwater production and stream macroinvertebrate biodiversity has wider implications for the conservation of alpine river ecosystems under scenarios of climate change induced glacier and snowpack loss.     

Altitudinal patterns of diversity and functional traits of metabolically active microorganisms in stream biofilms

Resources structure ecological communities and potentially link biodiversity to energy flow. It is commonly believed that functional traits (generalists versus specialists) involved in the exploitation of resources depend on resource availability and environmental fluctuations. The longitudinal nature of stream ecosystems provides changing resources to stream biota with yet unknown effects on microbial functional traits and community structure. We investigated the impact of autochthonous (algal extract) and allochthonous (spruce extract) resources, as they change along alpine streams from above to below the treeline, on microbial diversity, community composition and functions of benthic biofilms. Combining bromodeoxyuridine labelling and 454 pyrosequencing, we showed that diversity was lower upstream than downstream of the treeline and that community composition changed along the altitudinal gradient. We also found that, especially for allochthonous resources, specialisation by biofilm bacteria increased along that same gradient. Our results suggest that in streams below the treeline biofilm diversity, specialisation and functioning are associated with increasing niche differentiation as potentially modulated by divers allochthonous and autochthonous constituents contributing to resources. These findings expand our current understanding on biofilm structure and function in alpine streams.     

Meta‐Analysis of Lost Ecosystem Attributes in Urban Streams and the Effectiveness of Out‐of‐Channel Management Practices

Urban development is a leading cause of stream impairment that reduces biodiversity and negatively affects ecosystem processes and habitat. Out‐of‐stream restoration practices, such as stormwater ponds, created wetlands, and restored riparian vegetation, are increasingly implemented as management strategies to mitigate impacts. However, uncertainty exists regarding how effectively they improve downstream ecosystems because monitoring is uncommon and results are typically reported on a case‐by‐case basis. We conducted a meta‐analysis of literature and used response ratios to quantify how downstream ecosystems change in response to watershed development and to out‐of‐stream restoration. Biodiversity in unrestored urban streams was 47% less than that in reference streams, and ecological communities, habitat, and rates of nutrient cycling were negatively affected as well. Mean measures of ecosystem attributes in restored streams were significantly greater than, and 156% of, those in unrestored urban streams. Measures of biodiversity in restored streams were 132% of those in unrestored urban streams, and indices of biotic condition, community structure, and nutrient cycling significantly improved. However, ecosystem attributes and biodiversity at restored sites were significantly less than, and only 60% and 45% of, those in reference streams, respectively. Out‐of‐stream management practices improved ecological conditions in urban streams but still failed to restore reference stream conditions. Despite statistically significant improvements, assessing restoration success remains difficult due to few comparisons to reference sites or to clearly defined targets. These findings can inform future monitoring, management, and development strategies and highlight the need for preventative actions in a watershed context.     

Environmental DNA metabarcoding: Transforming how we survey animal and plant communities

The genomic revolution has fundamentally changed how we survey biodiversity on earth. High‐throughput sequencing (“HTS”) platforms now enable the rapid sequencing of DNA from diverse kinds of environmental samples (termed “environmental DNA” or “eDNA”). Coupling HTS with our ability to associate sequences from eDNA with a taxonomic name is called “eDNA metabarcoding” and offers a powerful molecular tool capable of noninvasively surveying species richness from many ecosystems. Here, we review the use of eDNA metabarcoding for surveying animal and plant richness, and the challenges in using eDNA approaches to estimate relative abundance. We highlight eDNA applications in freshwater, marine and terrestrial environments, and in this broad context, we distill what is known about the ability of different eDNA sample types to approximate richness in space and across time. We provide guiding questions for study design and discuss the eDNA metabarcoding workflow with a focus on primers and library preparation methods. We additionally discuss important criteria for consideration of bioinformatic filtering of data sets, with recommendations for increasing transparency. Finally, looking to the future, we discuss emerging applications of eDNA metabarcoding in ecology, conservation, invasion biology, biomonitoring, and how eDNA metabarcoding can empower citizen science and biodiversity education.     

Effects of pasture conversion to sugarcane for biofuel production on stream fish assemblages in tropical agroecosystems

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Aquifer and surface-water ostracodes in Quaternary paleowetland deposits of southern Nevada,USA

Despite the importance of small tropical streams for maintaining freshwater biodiversity and providing essential ecosystem services to humans, relatively few studies have investigated multiple-stressor effects of climate and land-use change on these ecosystems, and how these effects may interact. To illustrate these knowledge gaps, we reviewed the current state of knowledge regarding the ecological impacts of climate change and catchment land use on small tropical streams. We consider the effects of predicted changes in streamflow dynamics and water temperatures on water chemistry, habitat structure, aquatic biota, and ecosystem processes. We highlight the pervasive individual effects of climate and land-use change on algal, macroinvertebrate, and fish communities, and in stream metabolism and decomposition processes. We also discuss potential responses of tropical streams in a multiple-stressor scenario, considering higher temperatures and shifts in hydrological dynamics. Finally, we identify six key knowledge gaps in the ecology of low-order tropical streams and indicate future research directions that may improve catchment management in the tropics to help alleviate climate-change impacts and biodiversity losses.     

Climate‐induced glacier and snow loss imperils alpine stream insects

Climate warming is causing rapid loss of glaciers and snowpack in mountainous regions worldwide. These changes are predicted to negatively impact the habitats of many range‐restricted species, particularly endemic, mountaintop species dependent on the unique thermal and hydrologic conditions found only in glacier‐fed and snow melt‐driven alpine streams. Although progress has been made, existing understanding of the status, distribution, and ecology of alpine aquatic species, particularly in North America, is lacking, thereby hindering conservation and management programs. Two aquatic insects – the meltwater stonefly (Lednia tumana) and the glacier stonefly (Zapada glacier) – were recently proposed for listing under the U.S. Endangered Species Act due to climate‐change‐induced habitat loss. Using a large dataset (272 streams, 482 total sites) with high‐resolution climate and habitat information, we describe the distribution, status, and key environmental features that limit L. tumana and Z. glacier across the northern Rocky Mountains. Lednia tumana was detected in 113 streams (175 sites) within Glacier National Park (GNP) and surrounding areas. The probability of L. tumana occurrence increased with cold stream temperatures and close proximity to glaciers and permanent snowfields. Similarly, densities of L. tumana declined with increasing distance from stream source. Zapada glacier was only detected in 10 streams (24 sites), six in GNP and four in mountain ranges up to ~600 km southwest. Our results show that both L. tumana and Z. glacier inhabit an extremely narrow distribution, restricted to short sections of cold, alpine streams often below glaciers predicted to disappear over the next two decades. Climate warming‐induced glacier and snow loss clearly imperils the persistence of L. tumana and Z. glacier throughout their ranges, highlighting the role of mountaintop aquatic invertebrates as sentinels of climate change in mid‐latitude regions.     

Spatial patterns of plant diversity and communities in Alpine ecosystems of the Hengduan Mountains,Northwest Yunnan,China

Aims Conduct a quantitative, but rapid, regional-level assessment of the alpine flora across northwest Yunnan (NWY) to provide a broad-based understanding of local and regional patterns of the composition, diversity and health of alpine ecosystems across NWY.Methods A stratified random sampling design was employed to select sites across the different mountain ranges of NWY. Vegetation was sampled by stratifying each site by the three major alpine vegetation community types: meadow, dwarf shrub and scree. Two 50-m transects were randomly located within each community type at each sampling site with 10 1-m 2 subplots systematically placed along each transect. Environmental variables were recorded at each transect. Multivariate analyses were used to classify the major plant community assemblages and link community patterns to environmental and habitat variables.Important findings Forb species richness varied from 19 to 105 species per site (21 sites total) with an average of 59 species per site (60 m 2 sampled per site). Most species were patchily distributed with narrow distributions and/or small population sizes; over half the species occurred at only one or two sites. Distinct species assemblages were identified in the meadow vegetation that was strongly aggregated by geographic location suggesting the presence of distinct phytogeographic zones of the meadow alpine flora. Elevation and geographic location were the dominant environmental gradients underlying the variations in species composition. Jaccard's coefficient of similarity averaged only 10% among sites indicating there was little similarity in the alpine flora across the region. The alpine vegetation is highly heterogeneous across the complex landscape of the Hengduan Mountains of NWY. Conservation strategies need to take into account the large geographic differences in the flora to maximize protection of biodiversity.     

Surveying biodiversity in protected and managed areas: Algae, macrophytes and macroinvertebrates in boreal forest streams

Extensive surveys of biodiversity in protected and managed areas have not been conducted for a majority of taxonomic groups and ecosystem types, which makes it difficult to assess how large a portion of biodiversity is at least potentially under protection. The situation is the same in boreal regions, and only preliminary analyses of the biodiversity patterns of less well-known organism groups, including many freshwater taxa, within the protected area network have been conducted. We studied patterns of species richness and community composition of algae, macrophytes (bryophytes and vascular plants), and macroinvertebrates of headwater streams draining protected areas and managed forests in a boreal drainage basin in Finland. We found no significant differences in the species richness and community composition of these organism groups between the protected and managed streams. Gamma- and beta-diversity varied strongly among the protected and managed stream groups, yet this variation was contingent on the organism group and the beta-diversity measure used. In general, there was much species turnover within both protected and managed stream groups, masking any between-group differences. However, we found a number of redlisted and rare species in our surveys. Of these species, several macrophyte species occurred more frequently in the protected streams. By contrast, rare species of algae and macrophytes did not generally show such inclinations to the protected streams. We found no strong congruence in species richness or community dissimilarity between algae, macrophytes, and macroinvertebrates, suggesting that the main anthropogenic gradient in terms of forestry is not strong enough to modify stream environmental conditions and thereby shape biodiversity in the focal drainage basin. This finding also suggests that surveys of aquatic biodiversity across protected and managed landscapes should not rely too heavily on the surrogate taxon approach, but instead should consider patterns shown by multiple taxonomic groups that represent biologically and ecologically disparate organisms. Our results indeed suggest that the levels of alpha-, beta-, and gamma-diversity show differing among-taxon responses to forest management and naturalness of headwater streams.     

Decline in lizard species diversity,abundance and ectoparasite load across an elevational gradient in the Australian alps

The rapid changes in altitude, and associated habitat, of mountain ecosystems make them ideal natural laboratories for testing the effect of environmental heterogeneity on species assemblage. Our understanding of the sensitivity of Australian reptiles to elevational clines is limited. We examined lizard distribution across three elevation zones (montane, subalpine and alpine), spanning from 900 to 1840 m above sea level, in the Australian alps. We aimed to examine how elevation influences species diversity and abundance, and ectoparasite load, and whether species alter their habitat use amongst different elevational zones. Active searches were conducted across the elevation zones to identify lizard community structure (at least 16 species) across elevational zones, along with skink habitat preferences and the ectoparasite load. Skink diversity and abundance were negatively correlated with increased elevation. The alpine zone had significantly lower diversity and abundance of skinks. Habitat use differed amongst both elevations and species. Ectoparasite prevalence was also significantly diminished in the alpine zone. Ectoparasites only infected a subset of the skink community, with ectoparasite load increasing as the active season progressed. This study provides evidence of the complex interplay between elevation and species diversity, as well as the differences in ectoparasite pressure along elevational gradients in the Australian alps.     

Ecological gradients drive insect wing loss and speciation: The role of the alpine treeline

Alpine ecosystems are frequently characterized by an abundance of wing‐reduced insect species, but the drivers of this biodiversity remain poorly understood. Insect wing reduction in these environments has variously been attributed to altitude, temperature, isolation, habitat stability or decreased habitat size. We used fine‐scale ecotypic and genomic analyses, along with broad‐scale distributional analyses of ecotypes, to unravel the ecological drivers of wing reduction in the wing‐dimorphic stonefly Zelandoperla fenestrata complex. Altitudinal transects within populations revealed dramatic wing reduction over very fine spatial scales, tightly linked to the alpine treeline. Broad biogeographical analyses confirm that the treeline has a much stronger effect on these ecotype distributions than altitude per se. Molecular analyses revealed parallel genomic divergence between vestigial‐winged (high altitude) and full‐winged (low altitude) ecotypes across distinct streams. These data thus highlight the role of the alpine treeline as a key driver of rapid speciation, providing a new model for ecological diversification along exposure gradients.     

Litter decomposition in a temperate and a tropical stream: the effects of species mixing,litter quality and shredders

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Loss of small glaciers will diminish beta diversity in Pyrenean streams at two levels of biological organization

Aim Small (< 1 km²) alpine glaciers are likely to disappear in this century, resulting in decreased regional habitat heterogeneity in associated streams. Both heterogeneity within and spatial isolation among glacier‐influenced streams can enhance beta diversity of stream‐dwelling organisms. We measured beta at both community and population‐genetic levels within and among streams currently influenced by small Pyrenean glaciers. We aimed to evaluate whether patterns are analogous between the two levels, to apply various approaches for characterizing beta, and to infer the outcome of future glacier loss on regional biodiversity. Location Four glacier‐fed basins in the Parc National des Pyrénées, France. Methods We classified each of 18 stream reaches across the basins into either high‐, mid‐ or low‐‘glaciality’ (glacial influence) groups according to four physicochemical characteristics. At each reach, we collected macroinvertebrate communities and evaluated mitochondrial DNA haplotypes for 11–13 individuals of Baetis alpinus Pictet. Using taxa/haplotypes as basic units, we evaluated community and population‐genetic beta diversity simultaneously. We measured beta diversity in three major ways: as multivariate (Sørensen's dissimilarity, Jost D) and ‘classical’ (gamma/alpha) variation to compare among glaciality groups, and as turnover along the glaciality gradient within each basin. Results For most approaches at both organizational levels, beta was greatest among high‐glaciality reaches, absolute values of variation of beta in high‐glaciality streams were strikingly similar between levels, and the steepest turnover within basins occurred between high‐ and mid‐glaciality reaches. Therefore, high‐glaciality reaches contained assemblages and populations that were unique both within that stream type (among basins) and compared with other stream types within basins. Main conclusions Parallel beta diversity patterns at population‐genetic and community levels suggested that environmental drivers influence these levels analogously. Extreme conditions (e.g. low temperature, high instability, isolation) in high‐glaciality streams probably enhance beta at both levels. Stream beta diversity is likely to decrease substantially with continued glacial reduction in this system.     

Spatial sampling bias and model complexity in stream‐based species distribution models: A case study of Paddlefish (Polyodon spathula) in the Arkansas River basin,USA

Leveraging existing presence records and geospatial datasets, species distribution modeling has been widely applied to informing species conservation and restoration efforts. Maxent is one of the most popular modeling algorithms, yet recent research has demonstrated Maxent models are vulnerable to prediction errors related to spatial sampling bias and model complexity. Despite elevated rates of biodiversity imperilment in stream ecosystems, the application of Maxent models to stream networks has lagged, as has the availability of tools to address potential sources of error and calculate model evaluation metrics when modeling in nonraster environments (such as stream networks). Herein, we use Maxent and customized R code to estimate the potential distribution of paddlefish (Polyodon spathula) at a stream‐segment level within the Arkansas River basin, USA, while accounting for potential spatial sampling bias and model complexity. Filtering the presence data appeared to adequately remove an eastward, large‐river sampling bias that was evident within the unfiltered presence dataset. In particular, our novel riverscape filter provided a repeatable means of obtaining a relatively even coverage of presence data among watersheds and streams of varying sizes. The greatest differences in estimated distributions were observed among models constructed with default versus AIC_C‐selected parameterization. Although all models had similarly high performance and evaluation metrics, the AIC_C‐selected models were more inclusive of westward‐situated and smaller, headwater streams. Overall, our results solidified the importance of accounting for model complexity and spatial sampling bias in SDMs constructed within stream networks and provided a roadmap for future paddlefish restoration efforts in the study area.