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.     

Patterns of macroinvertebrate traits along three glacial stream continuums

1. Glacier‐fed streams are characterised by low spatial but high temporal heterogeneity, manifested in seasonal and diurnal discharge and suspended sediment peaks induced by glacial runoff. These streams shelter macroinvertebrate communities adapted to such harsh environmental conditions. Studies relating macroinvertebrate traits to environmental conditions in glacial streams could provide important insights into the structure and function of glacial stream communities. 2. From data collected in three glacial streams from the central Swiss and southern French Alps, we analysed the relationships among six biological traits to define five groups of macroinvertebrate taxa with similar suites of traits. 3. The longitudinal distribution of the five groups and of individual traits was analysed, as well as their variation according to a glaciality index combining water temperature, conductivity, suspended solids and substrate stability. 4. The trait diversity along the three streams showed a strong upstream‐downstream gradient. The upper reaches were dominated by a single group of taxa characterised by small, crawling, deposit feeders. The other trait‐based groups appeared progressively downstream. 5. Changes in the relative frequency of trait‐based groups along the glaciality gradient highlighted the dominance of all‐rounder resistant/resilient traits in the three streams and confirmed that environmental conditions in the glacial streams are too harsh or uniform to allow macroinvertebrate communities to develop alternative suites of traits. The findings are discussed in relation to the question of trait coding in the available literature.     

Genetic isolation among mountains but not between stream types in a tropical high‐altitude mayfly

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Functional structure and diversity of invertebrate communities in a glacierised catchment of the tropical Andes

1. In many mountainous areas, glaciers feed streams characterised by harsh environmental conditions, such as low water temperature, high turbidity, low channel stability, and high temporal variability in flow. Additionally, in many glacierised catchments, the mixture of streams arising from different water sources (glacier melt, groundwater, rainfall) generates high levels of environmental heterogeneity, which enhance species turnover rates and increase regional diversity.
2. Studies from mainly temperate regions have revealed some consistent patterns: a predominance of traits adaptive to harsh environmental conditions and reduced functional diversity with increased glaciality, both strongly related to environmental filtering. Here, we investigated variation in functional structure and diversity between macroinvertebrate communities from 15 stream sites, with different water sources (five glacier-fed, five groundwater-fed, and five mixed source) and level of glacier influence, in a glacierised catchment in the Ecuadorian Andes.
3. Our results revealed functional differences between communities inhabiting the different stream types. As found in temperate regions, high levels of glaciality were associated with an increase of small-sized taxa that do not swim but are temporarily attached to or burrow in the substrate, have a flying-adult stage, and feed by collecting–gathering. Similarly, we found a general decrease in functional diversity at sites with higher glacier influence. A null modelling approach suggested that in some of our glacier-fed sites, environmental filtering may be the main driver of community assembly, whereas other mechanisms, mainly regional (such as dispersal), but also local (such as intraspecific competition), may gain importance as glacier influence decreases.
4. Assemblage composition in streams in tropical glacierised catchments may be driven by both local and regional processes that generate a gradient of decreasing functional diversity with stronger glacier influence. However, lack of knowledge of relevant traits for taxa in tropical glacierised streams currently poses a substantial obstacle to predicting changes likely to arise from global warming and glacier melt in this region.

     

Drivers of beta diversity of macroinvertebrate communities in tropical forest streams

1. There has recently been increasing interest in patterns of beta diversity but we still lack a comprehensive understanding of these patterns in various regions (e.g. the tropics), ecosystems (e.g. streams) and organism groups (e.g. invertebrates). 2. Our aim was to investigate the patterns of beta diversity of stream macroinvertebrates in relation to key environmental (i.e. stream size, pH and habitat degradation) and geographical variables (i.e. latitude, longitude, altitude) in a tropical region. We surveyed a total of 8–10 riffle sites in each of 34 streams (altogether 337 riffle sites were sampled) in Peninsular Malaysia to examine variation in macroinvertebrate community composition at within‐stream and among‐stream scales. 3. Based on test of homogeneity of dispersion, we found that the streams studied differed significantly in within‐stream variation in community composition (i.e. among‐site variation of within stream beta diversity). The patterns were similar based on Bray–Curtis coefficient on abundance data, Sorensen coefficient on presence–absence data and Simpson coefficient on presence–absence data. We also found that within‐stream beta diversity was significantly related to stream size, pH and latitude, with each of these variables individually accounting for around 20% of the variation in beta diversity in simple regressions, while the total variation explained by the three significant variables amounted to around 50% in multiple regressions. By contrast, habitat degradation, longitude and altitude were not significantly related to beta diversity. We also found that the factor drainage basin accounted for much of the variation in beta diversity in general linear models, suppressing the effects of environmental variables. 4. We concluded that within‐stream beta diversity is mainly related to a combination of the identity of a drainage basin and stream environmental factors. Our findings provide important background for stream environmental assessment and conservation planning by emphasising that (i) macroinvertebrate communities within streams are not homogeneous, but show considerable beta diversity, (ii) streams differ in their degree of within‐stream beta diversity, (iii) stream size and water pH should be considered in applied contexts related to within‐stream beta diversity and (iv) historical effects may be different in different drainage basins and may affect present‐day patterns of within‐stream beta diversity.     

Physical and biological changes to a lengthening stream gradient following a decade of rapid glacial recession

In mountains, environmental gradients are steep in both terrestrial and aquatic systems, and climate change is causing upward shifts of physical and biological features of these gradients. Glacial streams are an interesting system to evaluate such shifts both because streams have a linear nature (for simplicity of analysis), and because the stream habitat will at least temporarily lengthen as it follows receding glaciers upward. The Tschierva Glacier, Swiss Alps, receded 482 m upstream from 1997 to 2008. We tested the null hypothesis that the physical and biological stream gradient below this glacier maintained the same structure between these time periods, but simply shifted upward following the glacial source. We compared longitudinal patterns of water temperature and zoobenthic community structure in 1997 and 2007–2008 during three seasons (spring, summer, fall) along the uppermost ca. 5 stream km. Upward shifts were evident, including colonization of the newly exposed stream reaches by cold‐adapted taxa, and the appearance in 2007/2008 of four lower‐altitude species that were previously absent. Overall, however, results rejected the null hypothesis, instead revealing significant changes in gradient structures. These included a more steeply increasing temperature profile downstream of the glacier and increased amplitude of seasonal community turnover in 2007/2008 vs. 1997. Long‐term (1955–2007) flow records revealed increasing short‐term and seasonal hydrologic variability, which might have influenced the increased intra‐annual community variability. The steepening of the temperature gradient was likely caused by a warming lake‐outlet tributary upon which glacial influence was diminished between 1997 and 2007/2008. These results suggest that upward‐shifting gradients in glacial streams can involve complex interactions with other landscape elements and that local‐scale climate response can progress even more rapidly than the rate of glacial recession.     

The influence of channelisation on riparian plant assemblages

1. A comparative study of species diversity and assemblage patterns of herbaceous ground‐flora communities in riparian areas was performed along natural mid‐sized lowland streams and their channelised counterparts. The areas had open vegetation and were positioned along 18 similar‐sized third to fourth order stream reaches. 2. Alpha diversity was significantly higher along natural streams both at the sample plot and reach scale. Sample plot diversity peaked at intermediate distance from the natural stream reaches, whereas it increased with increasing distance from the channelised reaches. Both gamma diversity, measured and estimated from species‐area curves, and beta diversity, which is a measure of the change in diversity between areas, was similar along the two types of streams. 3. Alpha diversity correlated with several of the measured and calibrated environmental variables. The positive correlation between bank slope and alpha diversity indicates that flooding plays a key role in maintaining high levels of diversity along natural streams. 4. Species composition varied significantly between the two stream types. A cluster analysis identified four clusters of which two clusters (one and three) primarily included species associated with sample plots in areas along natural streams. Most cluster one and three species were also identified as indicator species for this stream type. 5. Canonical correspondence analysis revealed that cluster one species were less productive species associated with high total soil carbon and nitrogen contents, whereas cluster three species were highly productive species associated with high soil moisture levels, probably partly resulting from flooding. Our results suggest that distance from the stream channel imposes a probabilistic gradient that sustains co‐occurrence of these two communities in riparian areas along natural mid‐sized lowland streams.     

Microbial biodiversity in glacier-fed streams

While glaciers become increasingly recognised as a habitat for diverse and active microbial communities, effects of their climate change-induced retreat on the microbial ecology of glacier-fed streams remain elusive. Understanding the effect of climate change on microorganisms in these ecosystems is crucial given that microbial biofilms control numerous stream ecosystem processes with potential implications for downstream biodiversity and biogeochemistry. Here, using a space-for-time substitution approach across 26 Alpine glaciers, we show how microbial community composition and diversity, based on 454-pyrosequencing of the 16S rRNA gene, in biofilms of glacier-fed streams may change as glaciers recede. Variations in streamwater geochemistry correlated with biofilm community composition, even at the phylum level. The most dominant phyla detected in glacial habitats were Proteobacteria, Bacteroidetes, Actinobacteria and Cyanobacteria/chloroplasts. Microorganisms from ice had the lowest α diversity and contributed marginally to biofilm and streamwater community composition. Rather, streamwater apparently collected microorganisms from various glacial and non-glacial sources forming the upstream metacommunity, thereby achieving the highest α diversity. Biofilms in the glacier-fed streams had intermediate α diversity and species sorting by local environmental conditions likely shaped their community composition. α diversity of streamwater and biofilm communities decreased with elevation, possibly reflecting less diverse sources of microorganisms upstream in the catchment. In contrast, β diversity of biofilms decreased with increasing streamwater temperature, suggesting that glacier retreat may contribute to the homogenisation of microbial communities among glacier-fed streams.     

Homage to the Virgin of Ecology,or why an aquatic insect unadapted to desiccation may maintain populations in very small,temporary Mediterranean streams

In temporary streams, the annual constriction of drying is associated with high local extinction risk. To survive in such habitats, organisms with no specific biological traits for coping with dry periods should experience high colonisation rates from permanent reaches of the same basin or from other basins. Hydropsyche siltalai is a widespread caddisfly common in permanent and temporary headwaters reaches in the Mediterranean climate region of the Iberian Peninsula. In this study, we used genetic analyses to test if populations of H. siltalai in temporary streams are resettled from populations of the same basin or from other basins. The geographical distribution of H. siltalai was surveyed in 97 temporary and permanent reaches across four basins; larvae were found in 22 reaches (12 temporary and 10 permanent). Population genetic analyses of 11 selected reaches (6 temporary and 5 permanent) revealed low genetic diversity and no genetic population structure among and within basins. Overall, H. siltalai appeared to disperse well among basins independent of stream temporality. Permanent reaches from different basins act as a source of the individuals that recolonise temporary reaches after local extinctions, indicating a metapopulation structure at regional scale. Moreover, our results support other studies that showed that dispersal among basins is a recurrent pattern in aquatic insects.     

Beta diversity of stream diatoms at two hierarchical spatial scales: implications for biomonitoring

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Longitudinal distribution of macroinvertebrate assemblages in a glacially influenced stream system in the Italian Alps

1. The longitudinal distribution of macroinvertebrates was investigated in June, August and September 1996 and 1997 in the Conca glacial stream and its tributary (Italian Alps; 46°N, 10°E). The principal aim was to test the 22 model that predicts the succession of faunal groups downstream of the glacial snout in relation to water temperature and channel stability. The effect of a non‐glacial tributary on the taxonomic richness and density patterns occurring in the glacial stream was also considered. 2. Channel stability showed an atypical longitudinal trend in the Conca glacial stream, being high in the upper part with Pfankuch Index values between 30 and 33. Water temperature exceeded 6 °C at all stations, with average values below 2 °C occurring only within 700 m from the glacial snout. 3. Taxonomic richness and diversity increased downstream. Taxonomic richness in the glacial stream (at about 1.5 km from the glacier) was comparable with the tributary and the reach after the confluence. Abundance also increased downstream in the glacial stream, but not as greatly as the number of taxa. 4. At higher taxonomic levels, the community structure in the tributary stations appeared to be similar to the two stations in the glacial stream just upstream of the confluence. The effect of the tributary was evident mainly at the genus or species level of the Chironomidae community. Some taxa found in the non‐glacial stream (e.g. Cricotopus fuscus, Eukiefferiella coerulescens, Metriocnemus sp., Paratrichocladius rufiventris, P. skirwitensis, Rheocricotopus effusus and Smittia sp.) were found also in the Conca stream but only after the confluence. 5. The upper glacial reach (within 700 m from the glacier snout) was dominated by the chironomid Diamesa spp. Less than 400 m from the glacier other Diamesinae (Pseudokiefferiella parva) and a few Orthocladiinae, especially Orthocladius (Euorthocladius) rivicola gr., colonized the stream. Some Diamesinae maintained relatively dense populations at mean water temperature around 5 °C, while some Orthocladiinae colonized reaches with mean water temperature <3 °C. 6. Contrary to the 22 model, Dipteran families such as Empididae and Limoniidae were more abundant in the upper stations than Simuliidae; non‐insects such as Nematoda and Oligochaeta were also numerous at some sites. Leuctridae, Taeniopterygidae and Nemouridae were the first Plecoptera to appear upstream, while Chloroperlidae were restricted to the lower reaches. Among Ephemeroptera, Heptageniidae were more abundant than Baetidae in the glacial sites. 7. In this glacial system channel stability and maximum temperature did not show the expected longitudinal trend and thus a typical kryal community was confined within 700 m from the glacier snout where summer mean water temperature was below 4 °C.     

祁连山区特有物种黑虎耳草的居群遗传多样性研究

利用叶绿体基因(rbcL和trnS-G)及核糖体DNA内转录间隔区(ITS),对祁连山区狭域分布的特有物种黑虎耳草8个居群(115个个体)进行遗传多样性研究,旨在揭示黑虎耳草的居群遗传结构及其历史进化过程。结果表明:(1)所有取样个体共检测到4个cpDNA单倍型和9个ITS单倍型,其中祁连山东南部的居群固定较多的单倍型和特有单倍型,而西北部居群只固定少数几个广泛分布的单倍型,且遗传多样性普遍较低。(2)基于cpDNA数据和ITS数据的分子变异分析(AMOVA)显示,遗传变异主要来源于居群内。(3)基于cpDNA数据的中性检验表明,Tajima’s D(-1.012 30,P 0.05)和FuLi’s D*(-2.066 77,P 0.05)均为负值,均不显著;歧点分布分析结果显示,黑虎耳草居群经历过近期的扩张事件。根据物种现有遗传分布格局推测,黑虎耳草在第四纪冰期时退缩到祁连山东南部的边缘避难所,间冰期或冰期后回迁到祁连山西北部地区,在回迁过程中由于奠基者效应导致祁连山西北部的居群仅固定少数广泛分布的单倍型,并呈现出较低的遗传多样性;由于居群较小且相互隔离,该物种经历了严重的瓶颈效应和遗传漂变,导致该物种总遗传多样性(H_T)和居群平均遗传多样性(H_S)远低于虎耳草属其他青藏高原-喜马拉雅广域分布的物种。     

Thermal patterns in the surface waters of a glacial river corridor (Val Roseg, Switzerland)

SUMMARY 1. We examined the thermal patterns of the surface waters in the catchment of the Roseg River, which is fed by the meltwaters of two valley glaciers. One of the glaciers has a lake at its terminus. The river corridor comprised a proglacial stream reach below one glacier, the glacier lake outlet stream, a 2.5‐km long complex floodplain and a constrained reach extending to the end of the catchment. 2. Temperatures were continuously measured with temperature loggers at 27 sites between 1997 and 1998. Moreover, from 1997 to 1999, spot measurements were taken at 33–165 floodplain sites (depending on water level) at monthly intervals. 3. The temperature regime of glacial streams, including the glacier lake outlet, was characterised by rapidly increasing temperatures in April and May, a moderate decline from June to September (period of glacial melt) and a subsequent fast decline in autumn. During summer, the lake increased temperatures in the outlet stream by 2–4 °C, compared with the adjacent proglacial stream reach. 4. In the main channel (thalweg) of the Roseg River, annual degree‐days (DD) ranged from 176 DD in the upper proglacial reach to 1227 DD at the end of the catchment. 5. Thermal variation among different channels within the floodplain was higher than the variation along the entire main channel. Floodplain channels lacking surface connection to the main channel accumulated up to 1661 annual DDs. 6. Thermal heterogeneity within the floodplain was linked to the glacial flow pulse. With the onset of ice melt, temperatures in the main channel and in channels surface‐connected to the main channel began to decline, whereas in surface‐disconnected channels temperatures continued to increase; as a consequence, thermal heterogeneity at the floodplain scale rose slightly until September. 7. High thermal heterogeneity was not anticipated in the harsh environment of a largely glacierised alpine catchment. The relatively wide range of thermal environments may contribute to the highly diverse zoobenthic community.     

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.     

Local environment rather than past climate determines community composition of mountain stream macroinvertebrates across Europe

Community assembly is determined by a combination of historical events and contemporary processes that are difficult to disentangle, but eco‐evolutionary mechanisms may be uncovered by the joint analysis of species and genetic diversity across multiple sites. Mountain streams across Europe harbour highly diverse macroinvertebrate communities whose composition and turnover (replacement of taxa) among sites and regions remain poorly known. We studied whole‐community biodiversity within and among six mountain regions along a latitudinal transect from Morocco to Scandinavia at three levels of taxonomic hierarchy: genus, species and haplotypes. Using DNA barcoding of four insect families (>3100 individuals, 118 species) across 62 streams, we found that measures of local and regional diversity and intraregional turnover generally declined slightly towards northern latitudes. However, at all hierarchical levels we found complete (haplotype) or high (species, genus) turnover among regions (and even among sites within regions), which counters the expectations of Pleistocene postglacial northward expansion from southern refugia. Species distributions were mostly correlated with environmental conditions, suggesting a strong role of lineage‐ or species‐specific traits in determining local and latitudinal community composition, lineage diversification and phylogenetic community structure (e.g., loss of Coleoptera, but not Ephemeroptera, at northern sites). High intraspecific genetic structure within regions, even in northernmost sites, reflects species‐specific dispersal and demographic histories and indicates postglacial migration from geographically scattered refugia, rather than from only southern areas. Overall, patterns were not strongly concordant across hierarchical levels, but consistent with the overriding influence of environmental factors determining community composition at the species and genus levels.     

Aquatic insect β‐diversity is not dependent on elevation in Southern Rocky Mountain streams

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Spatial variability in macroinvertebrate assemblages along and among neighbouring equatorial glacier‐fed streams

1. During the past two decades, understanding of the structure and function of glacier‐fed stream ecosystems at temperate latitudes has increased substantially. In contrast, information on their tropical counterparts is very limited. We studied three neighbouring glacier‐fed streams in the tropical Andes of Ecuador. Our main goals were (i) to determine overall longitudinal patterns in density, taxon richness and the composition of macroinvertebrate assemblages and driving factors in equatorial glacial streams and (ii) to examine variability among replicate streams in faunal metrics and assemblages, and stream‐specific effects of supposed environmental key factors. 2. We measured four geographical and 17 environmental factors and collected five Surber samples (500 cm²) of macroinvertebrates at each of nine sites, three sites along three streams. The streams were located 1–5 km apart. In each stream, the three sites were placed at comparable distances from the glacier and were grouped as ‘upper’ (50–200 m), ‘middle’ (1.5 km) and ‘lower’ sites (3.5–5.6 km). 3. In total, 2200 individuals (64% chironomids) were collected and 47 taxa (30 dipterans, 18 of these Chironomidae) identified. Density ranged from 176 to 372 ind. m^?2, and the number of taxa ranged from 2 to 6 at the upper sites and 868–3044 ind. m^?2 and 21–27 taxa at the lower sites. Density, number of taxa, rarefied richness and axis‐1 coordinates from a MDS ordination increased logarithmically with distance from the glacier. These faunal metrics were equally related to altitude and glacier per cent of catchment and correlated with maximum conductivity, mean temperature, mean daily maximum temperature and a channel stability index. As expected, the mean difference in distance decay in similarity was higher at the upper (47% km^?1) than at the lower reaches (20% km^?1) of the streams. 4. The number of taxa varied among sites within the upper and middle groups, but not among the lower sites. In contrast, but in accordance with our expectation, assemblage composition did not differ among upper sites but did so at middle and lower sites, following a supposed decrease in environmental harshness along the streams. Relationships between faunal metrics and the four environmental variables mean temperature, the stability index, chlorophyll a and coarse particulate organic matter also varied among the three streams. Generalised linear model analyses revealed that temperature interacted with stream on macroinvertebrate density, while chlorophyll a had a significant effect on the number of taxa in interaction with stream and stability. 5. The basic predictions of the Milner et al. (2001a) , model regarding longitudinal faunal patterns and temperature and stability as main driving factors were met by our three replicate equatorial glacial streams. Qualitative departures from the model were mainly because of zoogeographical differences. We demonstrated that variability in assemblages between comparable sites in closely situated streams was considerable, and the effect of key environmental factors varied among streams and interacted with other factors. Quantifying spatial variation in benthic assemblages may help us foresee possible consequences for biodiversity as a result of glacial retreat.     

Direct and indirect effects of glaciers on aquatic biodiversity in high Andean peatlands

The rapid melting of glacier cover is one of the most obvious impacts of climate change on alpine ecosystems and biodiversity. Our understanding of the impact of a decrease in glacier runoff on aquatic biodiversity is currently based on the ‘glacier‐heterogeneity‐diversity’ paradigm, according to which there is high α‐diversity at intermediate levels of glacial influence due to the high degree of environmental heterogeneity caused by glacier water. This α‐diversity pattern generates high levels of between‐site aquatic community variation (high β diversity) and increases regional diversity (γ‐diversity). There is a rich conceptual background in favor of this paradigm, but empirical data supporting it are scarce. We investigated this paradigm by analyzing the different diversity patterns (α, β and γ‐diversity) of four aquatic groups (zooplankton, macroinvertebrates, algae and macrophytes) living in high‐elevation peatlands (>4500 m above sea level). We sampled 200 pools from 20 peatlands along a glacier gradient in the Cordillera Real of Bolivia. We performed structural equation modeling (SEM) to analyze the potential mechanisms underlying the observed diversity patterns. Intermediate levels of glacial influence (15–20% cover) resulted in high heterogeneity, but α‐diversity responded to glacial influence only for the zooplankton group (Cladocera). Our SEM analysis did not identify environmental heterogeneity as a significant variable explaining the relationship between glacier and α‐diversity. Peatland area had a strong positive effect on heterogeneity and diversity. β‐diversity was significantly associated with glacier gradient, and 12.9% of the total regional diversity (γ‐diversity) was restricted to peatlands with a high degree of glacial influence. These species might be lost in a context of glacial retreat. These findings provide new insight into the potential effects of glacial retreat on the aquatic environment and biodiversity in the peatlands of the tropical Andes.     

BIODIVERSITY RESEARCH: Conserving macroinvertebrate diversity in headwater streams: the importance of knowing the relative contributions of α and β diversity

Aim We investigated partitioning of aquatic macroinvertebrate diversity in eight headwater streams to determine the relative contributions of α and β diversity to γ diversity, and the scale dependence of α and β components. Location Great Dividing Range, Victoria, Australia. Methods We used the method of Jost (Ecology, 2007, 88, 2427–2439) to partition γ diversity into its α and β components. We undertook the analyses at both reach and catchment scales to explore whether inferences depended on scale of observation. Results We hypothesized that β diversity would make a large contribution to the γ diversity of macroinvertebrates in our dendritic riverine landscape, particularly at the larger spatial scale (among catchments) because of limited dispersal among sites and especially among catchments. However, reaches each had relatively high taxon richness and high α diversity, while β diversity made only a small contribution to γ diversity at both the reach and catchment scales. Main conclusions Dendritic riverine landscapes have been thought to generate high β diversity as a consequence of limited dispersal and high heterogeneity among individual streams, but this may not hold for all headwater stream systems. Here, α diversity was high and β diversity low, with individual headwater stream reaches each containing a large portion of γ diversity. Thus, each stream could be considered to have low irreplaceability since losing the option to use one of these sites in a representative reserve network does not greatly diminish the options available for completing the reserve network. Where limited information on individual taxonomic distributions is available, or time and money for modelling approaches are limited, diversity partitioning may provide a useful ‘first‐cut’ for obtaining information about the irreplaceability of individual streams or subcatchments when establishing representative freshwater reserves.     

Higher β‐ and γ‐diversity at species and genetic levels in headwaters than in mid‐order streams in Hydropsyche (Trichoptera)

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