Cryptic diversity within the rotifer Polyarthra dolichoptera along an altitudinal gradient

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Influence of environmental variables on macroinvertebrate community structure in Lianhuan Lake

The structural characteristics of the macroinvertebrate community can effectively reflect the health status of lake ecosystems and the quality of the lake ecological environment. It is therefore important to identify the limiting factors of macroinvertebrate community structure for the maintenance of lake ecosystem health. In this study, the community composition of macroinvertebrate assemblages and their relationships with environmental variables were investigated in 13 small lakes within Lianhuan Lake in northern China. A self‐organizing map and K‐means clustering analysis grouped the macroinvertebrate communities into five groups, and the indicator species reflected the environmental characteristics of each group. Principal component analysis indicated that the classification of the macroinvertebrate communities was affected by environmental variables. The Kruskal–Wallis test results showed that environmental variables (pH, total phosphorus, nitrate, water temperature, dissolved oxygen, conductivity, permanganate index, and ammonium) had a significant effect on the classification of the macroinvertebrate communities. Redundancy analysis showed that mollusks were significantly negatively correlated with pH and chlorophyll a, while annelids and aquatic insects were significantly positively correlated with chlorophyll a and dissolved oxygen. Spearman correlation analysis showed that the species richness and Shannon''s diversity of macroinvertebrates were significantly negatively correlated with total phosphorus, while the biomass of macroinvertebrates was significantly negatively correlated with pH. High alkalinity and lake eutrophication have a serious impact on the macroinvertebrate community. Human disturbances, such as industrial and agricultural runoff, negatively impact the ecological environment and affect macroinvertebrate community structure. Thus, macroinvertebrate community structure should be improved by enhancing the ecological environment and controlling environmental pollution at a watershed scale.     

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.     

Stable isotopes of lakes and precipitation along an altitudinal gradient in the Eastern Alps

Altitude encompasses broad environmental gradients that influence the isotopic composition of lake water. We selected 55 lakes in the Eastern Alps along an altitudinal gradient [214–2,532 m above sea level (a.s.l.)] to model the isotopic signal of surface water dependent on intrinsic (lake geomorphometry) and extrinsic (air temperature, precipitation) factors. Ordinary and generalised least squared regression were used for statistical analysis. The isotope signal of lake water was lower in spring than in summer and decreased with altitude (?0.21 δ¹⁸O ‰/100 m; ?1.5 δ²H ‰/100 m). This pattern largely depended on temperature and a pseudo-latitude effect. The isotopic signal in monthly precipitation (12 stations; altitudinal gradient 90–2,730 m a.s.l.) generally showed the expected pattern of less enriched values with altitude; however, unusual values were related to weather anomalies. The local meteoric water line was similar to the global meteoric water line as shown by overlapping confidence intervals. By discriminating different elevational bands, we could show that high elevation lakes (>1,500 m a.s.l.) experience different patterns of evaporation with respect to low elevation lakes (<1,500 m a.s.l.). Our study showed that lakes have a unique isotopic fingerprint along an altitudinal gradient, potentially useful for tracing ecological processes and for paleoclimatic studies.     

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.     

Rotifer species richness along an altitudinal gradient in the Alps

Aim Biodiversity patterns along altitudinal gradients are less studied in aquatic than terrestrial systems, even though aquatic sites provide a more homogeneous environment independent of moisture constraints. We studied the altitudinal species richness pattern for planktonic rotifers in freshwater lakes and identified the environmental predictors for which altitude is a proxy. Location Two hundred and eighteen lakes of Trentino–South Tyrol (Italy) in the eastern Alps; lakes covered 98% (range 65–2960 m above sea level) of the altitudinal gradient in the Alps. Methods We performed: (1) linear regression between species richness and altitude to evaluate the general pattern, (2) multiple linear regression between species richness and environmental predictors excluding altitude to identify the most important predictors, and (3) linear regression between the residuals of the best model of step (2) and altitude to investigate any additional explanatory power of altitude. Selection of environmental predictors was based on limnological importance and non‐parametric Spearman correlations. We applied ordinary least squares regression, generalized linear, and generalized least squares modelling to select the most statistically appropriate model. Results Rotifer species richness showed a monotonic decrease with altitude independent of scale effects. Species richness could be explained (R²= 51%) by lake area as a proxy for habitat diversity, reactive silica and total phosphorus as proxies for productivity, water temperature as a proxy for energy, nitrate as a proxy for human influence and north–south and east–west directions as covariates. These predictors completely accounted for the species richness–altitude pattern, and altitude had no additional effect on species richness. Main conclusions The linear decrease of species richness along the altitudinal gradient was related to the interplay of habitat diversity, productivity, heat content and human influence. These factors are the same in terrestrial and aquatic habitats, but the greater environmental stability of aquatic systems seems to favour a linear pattern.     

Diversity and abundance of littoral cladocerans and copepods in nine Ecuadorian highland lakes

The diversity and abundance of littoral cladocerans and copepods were studied in nine lakes at Reserva Ecológica Cayambe-Coca (Páramo de Guamaní), Ecuador. Six samples were taken in the littoral zone of each lake using a 500 microm mesh plankton conic net. One species of cladocerans (Ephemeroporus acanthoides) is reported for the first time in Ecuador. The diversity (H') and evenness (E) of the lakes were determined and correlated with PCA axes based on their environmental variables. The principal parameters that distinguished these lakes were altitude and pH, an unexpected finding considering that the altitudinal range was very small. Lake size is of secondary importance for this group of lakes. None of the environmental axes correlated with H' or E; nevertheless, a larger than expected species richness was found in a small oligotrophic lake with a high level of DO. Based on our results, we hypothesize that altitude and pH are important factors determining the zooplankton diversity (directly or indirectly) in highland lakes.     

Structure of chironomid assemblages along environmental and geographical gradients in the Bohemian Forest lakes (Central Europe): An exploratory analysis

Eight glacial lakes of the Bohemian Forest (Czech Republic and Germany) were characterised by the distribution of chironomids collected as pupal exuviae. Twenty-eight taxa were identified, including some faunistically interesting species of the region. Two-way indicator species analysis (TWINSPAN) was used to classify lakes according to their taxonomic composition. Canonical correspondence analysis (CCA) and multiple regression were used to relate the chironomid assemblages to two sets of explanatory variables: (i) local environmental variables, and (ii) broad-scale spatial variables. The TWINSPAN classified the lakes into four groups, whereas presence/absence of three taxa was indicative for this classification. The CCA of assemblage composition on environmental variables showed that chironomids respond significantly to altitude and alkalinity. The ordination of composition data on geographical variables revealed strong longitudinal gradient in chironomid distributions. Altitude and alkalinity accounted for 36.2% of the total variation, while the geographic gradient explained 20.5%. As revealed by the variation partitioning procedure, the significant effect of these variables was, in large part, independent of each other. Overall taxonomic richness appeared to be governed by altitude only. Causal ecological and historical factors underlying these results are discussed. This paper may provide a basis for hypothesis testing in future research of the Bohemian Forest lakes.     

Functional characters combined with null models reveal inconsistency in mechanisms of species turnover in lacustrine fish communities

Functional characters have the potential to act as indicators of species turnover between local communities. Null models provide a powerful statistical approach to test for patterns using functional character information. A combined null model/functional character approach provides the ability to distinguish between the effect of competition and environmental filtering on species turnover. We measured 13 functional characters relating directly to resource use for the fish species found in French lakes. We combined this functional character data with a null model approach to test whether co-occurring species overlapped more or less than expected at random for four primary niche axes. We used an environmentally constrained null model approach to determine if the same mechanisms were responsible for species turnover at different sections of the altitudinal gradient. Functional diversity indices were used to examine the variation in functional character diversity with altitude, as a test of the hypothesis that competitive intensity decreases with increasing environmental adversity. The unconstrained null model showed that environmental filtering was the dominant influence on species turnover between lakes. In the constrained null model, there was much less evidence for environmental filtering, emphasising the strong effect of altitude on turnover in functional character values between local communities. Different results were obtained for low-altitude and high-altitude lake subsets, with more evidence for the effect of environmental filtering being found in the high-altitude lakes. This demonstrates that different processes may influence species turnover throughout an environmental gradient. Functional diversity values showed a slight decrease with altitude, indicating that there was only weak evidence that competitive intensity decreased with increasing altitude. Variation resource availability and environmental stress probably cause the observed turnover in functional characters along the altitudinal gradient, though the effects of dispersal limitation and species introductions in high-altitude lakes cannot be ruled out.     

Short-term effect of oxic to anoxic transition on benthic microbial activity and solute fluxes in organic-rich phytotreatment ponds

Macroinvertebrates are one of the key components of lake ecosystems and are required to be monitored alongside other biological groups to define ecological status according to European Union legislation. Macroinvertebrate communities are highly variable and complex and respond to a diverse series of environmental conditions. The purpose of this study was to examine the relative importance of environmental variables in explaining macroinvertebrate abundance. A total of 45 sub-alpine lakes were sampled for macroinvertebrates in the shallow sublittoral. Environmental variables were grouped into four types: (1) aquatic physical and chemical parameters, (2) littoral and riparian habitat, (3) lake morphometric parameters and (4) sediment chemical characteristics. Nonparametric multiplicative regression (NPMR) was used to model the abundance of individual macroinvertebrate taxa. Significant models were produced for nine out of the 24 taxa examined. Sediment characteristics were the group most frequently included in models and also the factors to which taxa abundance was the most sensitive. Aquatic physical and chemical variables were the next group most frequently included in models although chlorophyll a was not included in any of the models and total phosphorus in only one. This indicates that many taxa may not show a direct easily interpretable response to eutrophication pressure. Lake morphometric factors were included in several of the models although the sensitivity of macroinvertebrate abundance tended to be lower than for sediment and aquatic physical and chemical factors. Habitat factors were only included in three models although riparian vegetation was found to have a significant influence on the abundance of Ephemera danica indicating that ecotone integrity is likely to play a role in its ecology. Overall, the models tended to be specific for species with limited commonality across taxa. Models produced by NPMR indicate that the response of macroinvertebrates to environmental variables can be successfully described but further research is required focussing in more detail on the response of key taxa to relevant environmental parameters and anthropogenic pressures.     

Flagellate algae (Chrysophyceae, Dinophyceae, Cryptophyceae) in 48 high mountain lakes of the Northern and Southern slope of the Eastern Alps: biodiversity, taxa distribution and their driving variables

Survey samplings on late summer phytoplankton communities were carried out on 48 high mountain lakes located on the Austrian and Italian slopes of the Eastern Alps. The lakes of North Tyrol (A) and South Tyrol (I) were sampled in 2000 as part of the EU project EMERGE (EVK1-CT-1999- 00032). The lakes of Trentino (I) were investigated in other research projects during 1996 and 1997 (Adamello mountain range) and 2000 (catchment of the River Avisio), respectively. The objectives of this paper are: (1) to study taxonomy and biodiversity of Chrysophyceae, Dinophyceae and Cryptophyceae in high altitude lakes of the Eastern Alps; (2) to identify functional flagellate groups characterising lakes with similar habitat properties, (3) to identify the environmental variables driving abundance and distribution of the three selected algal groups, thus contributing to the selection of sensitive bioindicator taxa. The lakes investigated show rather wide morphological, chemical and trophic state gradients. Flagellate algae account for a median relative abundance (R.A.) of 68%. Chrysophyceae are the most important group in terms of biodiversity and R.A. Special flagellate associations could be related to lake features, like catchment geology, mineralization level and nutrient concentrations. However, the distribution of flagellate algae did not allow a complete geographical separation of the lakes studied in the different districts. Multivariate canonical analyses indicate that the distribution of Chrysophyceae is mainly driven by NO₃-N concentration and thermal conditions, while Dinophyceae are driven by a combination of alkalinity, altitude, thermal condition and, less importantly, nutrient concentration. Physical properties of the lakes, such as thermal condition and lake depth, represent the principal driving variables for Cryptophyceae. The responses to the different environmental variables suggest that the three flagellate groups analysed might be used as indicators for environmental changes in high mountain lakes of the Eastern Alps.     

Relationships between aquatic plants and environmental factors along a steep Himalayan altitudinal gradient

We examined the relationships of aquatic plants with physical–chemical characteristics among 28 lakes within a steep altitudinal gradient ranging from tropical (77 m) to high alpine (4750 m) in the Himalayas of Nepal. Species richness and diversity showed an approximately linear decrease with increasing altitude. The study region exhibits a relatively high proportion of monocotyledonous helophytes, as is typical of aquatic plants on the Indian subcontinent. A canonical correspondence analysis of the entire altitudinal gradient (CCA-1) suggested that the strongest abiotic influences on the distribution of aquatic plants are associated with water temperature, substrate quality, altitude, pH, transparency and conductivity. Two more-restricted CCA analyses examined a shorter altitudinal gradient of 70–1500 m. The CCA-2 analysis (all aquatic plants) and CCA-3 (only euhydrophytes) found that the most important abiotic influences were associated with temperature, lake surface area, suspended solids, bicarbonate and dissolved phosphorus. These results suggest that relatively local influences are different from those that have a regional basis, but that climate-related influences are key along altitudinal gradients. The temperature gradient in the CCA distinguished montane and alpine Arcto-tertiary floristic elements of the highest altitudinal regions from the more widely distributed temperate and tropical species of the lower regions.     

Zooplankton (Cladocera) in assessments of biologic integrity and reference conditions: application of sedimentary assemblages from shallow boreal lakes

Zooplankton are potentially powerful proxies for the assessments of biologic integrity. The paleolimnological perspective and use of fossil Cladocera also provide the means to reconstruct reference conditions and natural long-term community dynamics. Unfortunately, the use of zooplankton in lake quality assessments is currently underexploited. We studied a surface sediment dataset of 41 lakes in Finland to examine the relationship between Cladocera remains and environmental variables. Of the examined environmental variables, total phosphorus availability was found to be the most important variable in explaining the Cladocera community composition. Following the tests on species environment relations, we selected a lake trophic typology as the most suitable environmental variable for developing a new tool for limnoecological quality assessments. A test of the model on a modern and historic sample from a eutrophied lake showed that the test lake has proceeded from “mesotrophic/poor” to “eutrophic/bad” limnoecological state in agreement with previous independent evidence. The model developed here showed favorable performance that can be used to provide reliable estimates of ecological and environmental state of lakes.     

Environmental Variability, Historical Contingency, and the Structure of Regional Fish and Macroinvertebrate Faunas in Ouachita Mountain Stream Systems

In 1990–1992, the United States Forest Service sampled six hydrologically variable streams paired in three different drainage basins in the Ouachita Mountains, Arkansas, U.S.A. Fishes, macroinvertebrates, and stream environmental variables were quantified for each stream. We used these data to examine the relationship between regional faunas (based on taxonomy and trophic affiliation of fishes and macroinvertebrates) and measured environmental variables. Because fishes are constrained to their historically defined drainage basins and many insect taxa are able to cross basin barriers, we anticipated that both groups would respond differently to environmental variability. Fishes were influenced more by environmental variability that was unique to their historical drainage basins, but macroinvertebrates were associated more strongly with environmental variability that was independent of drainage basins. Thus, the individual drainage basins represented a historical constraint on regional patterns of fish assembly. For both fishes and macroinvertebrates, groupings based on taxonomy and trophic affiliation showed a similar response to environmental variability and there was a high degree of association between taxonomic and trophic correlation matrices. Thus, trophic group structure was highly dependent on the taxonomic make-up of a given assemblage. At the basin-level, fish and macroinvertebrate taxa were associated more strongly with environmental variability than the trophic groups, and these results have implications for basin-level studies that use trophic groupings as a metric to assess ecological patterns. Trophic categories may not be a useful ecological measure for studies at large spatial scales.     

Concordance between ecotypes and macroinvertebrate assemblages in Mediterranean streams

1. According to the guidelines of the European Water Framework Directive, assessment of the ecological quality of streams and rivers should be based on ecotype-specific reference conditions. Here, we assess two approaches for establishing a typology for Mediterranean streams: a top-down approach using environmental variables and bottom-up approach using macroinvertebrate assemblages.
2. Classification of 162 sites using environmental variables resulted in five ecotypes: (i) temporary streams; (ii) evaporite calcareous streams at medium altitude; (iii) siliceous headwater streams at high altitude; (iv) calcareous headwater streams at medium to high altitude and (v) large watercourses.
3. Macroinvertebrate communities of minimally disturbed sites ( n  = 105), grouped using UPGMA (unweighted pair-group method using arithmetic averages) on Bray–Curtis similarities, were used to validate four of the five ecotypes obtained using environmental variables; ecotype 5, large watercourses, was not included as this group had no reference sites.
4. Analysis of similarities ( anosim ) showed that macroinvertebrate assemblage composition differed among three of the four ecotypes, resulting in differences between the bottom-up and top-down classification approaches. Siliceous streams were clearly different from the other three ecotypes, evaporite and calcareous ecotypes did not show large differences in macroinvertebrate assemblages and temporary streams formed a very heterogeneous group because of large variability in salinity and hydrology.
5. This study showed that stream classification schemes based on environmental variables need to be validated using biological variables. Furthermore, our findings indicate that special attention should be given to the classification of temporary streams.     

Analysis of the reproductive strategy of <Emphasis Type="Italic">Jenynsia multidentata</Emphasis> (Cyprinodontiformes,Anablepidae) with focus on sexual differences in growth,size, and abundance

The habitat heterogeneity generated and sustained by the connectivity of floodplain habitats, the seasonal flood pulse, and the variability of the physical structures typically found in floodplains of large rivers results in a variable space–time mosaic of water sources that results in a high biodiversity of the river-floodplain system. In order to assess the implications of natural connectivity and the heterogeneity on the patterns of macroinvertebrate assemblages at different spatial scales, monthly samplings in six different mesohabitats (lakes with different hydrological connection and secondary channels with permanently and intermittent flow) of the Paraná River floodplain were performed from April 2005 to March 2006. The mesohabitats had different granulometry and detritus composition of their bottom sediments. They also had different conductivity, transparency, and depth in relation to the different connectivity degrees. Mesohabitats differed in the abundance of macroinvertebrates of different taxonomic groups and diversity. The environmental variables were correlated to the patterns of macroinvertebrate abundance, with dominance of different species of annelids and mollusks at the patch, mesohabitats, and island scales. An alpha diversity gradient from the isolated lake (65 taxonomic units) to the secondary channels (25 taxonomic units) was obtained. The analyzed mesohabitats showed a high taxa turnover, with high values not only among the mesohabitats located in the different islands, but also among the mesohabitats in relation to different connectivity degrees. The mesohabitats showed negative co-occurrence of macroinvertebrate assemblages. The spatial heterogeneity, sustained by the connectivity degree, played a key role in structuring benthic assemblages at different scales, positively influencing the regional diversity.     

Weak altitudinal pattern of overall chironomid richness is a result of contrasting trends of subfamilies in high-altitude ponds

The decline of species richness with altitude is one of the most obvious patterns in ecology and results from the combination of ecological and evolutionary mechanisms. In harsh high-altitude environments, the effect of altitude usually overrules other environmental variables related to biodiversity. Studies using species richness along altitude gradients in high altitude are relatively numerous for lakes, but not for ponds. However, due to their special features, such as small size, high isolation and regional variability, ponds have been proved to be different systems compared to lakes. In high-altitude waterbodies, species of the family Chironomidae often dominate in benthic invertebrate communities and thus serve as an ideal model to study aquatic community changes along an altitude gradient. However, due to the time-consuming processing and expertise needed to identify the species, chironomids are often excluded from regular surveys. In the present study, we sampled 66 Tatra ponds over a 1100-m altitude gradient for benthic invertebrates, with special attention to chironomids. Out of the total 122 taxa collected, Chironomidae constituted the richest group with 58 taxa, being present in all the study ponds. The most diverse pond supported 13 chironomid taxa, and mean diversity was 6 taxa/pond. While total invertebrate richness decreased with altitude, chironomid richness showed only a weak negative response to altitude. The proportion of total chironomid diversity made up of Tanypodinae and Chironominae subfamilies decreased with altitude, while the opposite trend was recorded for the proportion of Diamesinae and Orthocladiinae.     

Littoral benthic macroinvertebrates of alpine lakes (Tatra Mts) along an altitudinal gradient: a basis for climate change assessment

Littoral benthic macroinvertebrates were studied in three alpine lakes in the High Tatra Mountains (Slovakia) located at different elevations: 2157, 1940 (alpine zone) and 1725 m (sub-alpine zone). The study sites were selected in order to obtain a gradient in thermal regimes and particular organic matter (POM). Differences in the faunal composition of lakes were tested for the ability of these differences to indicate climatic changes, and species/taxa were identified that could be used for the purposes of monitoring and climate change assessment. Macroinvertebrates were sampled quantitatively during the ice-free seasons of 2000 and 2001, and lake surface water temperature (LSWT) and POM were measured. LSWT and POM were negatively correlated with elevation, whereas ice cover was positively correlated with elevation. A total of 60 oligostenothermic macroinvertebrate species/taxa were collected belonging to ten higher taxonomic groups. Statistical analysis showed trends in several biotic metrics with altitude. More specifically, there was a clear increase in the number of species/taxa, genera, and higher taxonomic groups, as well as an increase in the Shannon–Wiener diversity with decreasing altitude. On the contrary, evenness and density of benthic macroinvertebrates did not show any clear relationship with altitude. Gatherers of detrital particles dominated the assemblages’ trophic structures, but no distinct changes in the proportion of functional feeding groups along the altitudinal gradient were found. While the non-insect fauna of the lakes was rather uniform across the elevational gradient, the insect fauna composition was highly correlated with altitude, as confirmed by Detrended Correspondence Analysis. Aquatic insects, in particular chironomids and caddisflies, can therefore be used as good indicators of temperature changes. Our results suggest that under warmer conditions, non-insect benthic macroinvertebrates will remain more or less stable, while aquatic insects will undergo an increase in the number of thermophilic species typical for lower altitudes. These colonizers will increase the diversity of alpine lakes, while the extinction of cold stenothermal species will lead to impoverishment of the native fauna. An indirect impact on benthic macroinvertebrates through changes in food sources is likely, and changes in trophic structure of the littoral assemblages can be expected.     

Patterns of fish species richness in China's lakes

Aim To document the patterns of fish species richness and their possible causes in China's lakes at regional and national scales. Location Lakes across China. Methods We compiled data of fish species richness, limnological characteristics and climatic variables for 109 lakes across five regions of China: East region, Northeast region, Southwest region, North‐Northwest region, and the Tibetan Plateau. Correlation analyses, regression models and a general linear model were used to explore the patterns of fish species richness. Results At the national scale, lake altitude, energy availability (potential evapotranspiration, PET) and lake area explained 79.6% of the total variation of the lake fish species richness. The determinants of the fish richness pattern varied among physiographic regions. Lake area was the strongest predictor of fish species richness in the East and Southwest lakes, accounting for 22.2% and 82.9% of the variation, respectively. Annual PET explained 68.7% of the variation of fish richness in the Northeast lakes. Maximum depth, mineralization degree, and lake area explained 45.5% of the fish variation in the lakes of the North‐Northwest region. On the Tibetan Plateau, lake altitude was the first predictor variable, interpreting 32.2% of the variation. Main conclusions Lake altitude was the most important factor explaining the variation of fish species richness across China's lakes, and accounted for 74.5% of the variation. This may stem in part from the fact that the lakes investigated in our study span the largest altitudinal range anywhere in the world. The effects of the lake altitude on fish species richness can be separated into direct and indirect aspects due to its collinearity with PET. We also found that the fish diversity and its determinants were scale‐dependent. Fish species richness was probably energy‐determined in the cold region, while it was best predicted by the lake area in the relatively geologically old region. The independent variables we used only explained a small fraction of the variations in the lake fish species richness in East China and the Tibetan Plateau, which may be due to the effects of human activity and historical events, respectively.     

Plant species and growth form richness along altitudinal gradients in the southwest Ethiopian highlands

Questions: Do growth forms and vascular plant richness follow similar patterns along an altitudinal gradient? What are the driving mechanisms that structure richness patterns at the landscape scale? Location: Southwest Ethiopian highlands. Methods: Floristic and environmental data were collected from 74 plots, each covering 400 m². The plots were distributed along altitudinal gradients. Boosted regression trees were used to derive the patterns of richness distribution along altitudinal gradients. Results: Total vascular plant richness did not show any strong response to altitude. Contrasting patterns of richness were observed for several growth forms. Woody, graminoid and climber species richness showed a unimodal structure. However, each of these morphological groups had a peak of richness at different altitudes: graminoid species attained maximum importance at a lower elevations, followed by climbers and finally woody species at higher elevations. Fern species richness increased monotonically towards higher altitudes, but herbaceous richness had a dented structure at mid‐altitudes. Soil sand fraction, silt, slope and organic matter were found to contribute a considerable amount of the predicted variance of richness for total vascular plants and growth forms. Main Conclusions: Hump‐shaped species richness patterns were observed for several growth forms. A mid‐altitudinal richness peak was the result of a combination of climate‐related water–energy dynamics, species–area relationships and local environmental factors, which have direct effects on plant physiological performance. However, altitude represents the composite gradient of several environmental variables that were interrelated. Thus, considering multiple gradients would provide a better picture of richness and the potential mechanisms responsible for the distribution of biodiversity in high‐mountain regions of the tropics.