Macroalgae,a suitable indicator of the ecological status of coastal rocky communities in the NE Atlantic

Despite the great importance of shallow rocky communities (<30 m deep) due to their contribution to the biodiversity of coastal waters, most efforts in ecological status assessment of marine waters have been carried out in the implementation of soft bottom biotic indices. Therefore, in this paper, a methodology for the environmental evaluation of coastal rocky communities for the Bay of Biscay (NE Atlantic), the CFR index (Quality of Rocky Bottoms), is presented. This index is based on the analysis of seaweed communities throughout the depth gradient, from the intertidal to the shallow subtidal, through the combination of a multimetric approach, which combines the richness of characteristic macroalgae populations, their total cover, the presence of opportunistic species and the physiological condition of the whole macroalgae community. Quality thresholds for these indicators are based on both ecological data and expert judgment. Finally, a preliminary application of this metric to sites with theoretically good and bad environmental conditions is used to analyze the suitability of the index. As a result, the four selected indicators responded in a significant way to the pressure gradient tested, proving to be appropriate for the type of pressure analysed and obtaining coherent results in the final quality assessment with the CFR index. In conclusion, the CFR index has proved to be an effective tool for the assessment of the ecological quality of coastal rocky communities, based on the analysis of macroalgae assemblages.     

Ability of invertebrate indices to assess ecological condition on intertidal rocky shores

The implementation of directives such as the European Water Framework Directive (WFD) and the Marine Strategy Framework Directive (MSFD) has promoted the development of several tools and methods for assessing the ecological health of marine ecosystems. Within the scope of the WFD and in terms of rocky shores, several multimetric tools were developed based on the macroalgae biological quality element (BQE), in addition to those based on macroinvertebrates.The WFD requires member states to assess each BQE separately. The present work aimed to test the ability of ecological indices to distinguish sites within anthropogenic disturbance gradients caused by organic enrichment, using macroinvertebrate communities on intertidal rocky shores. Owing to the lack of more specific indices (for rocky shore), indices based on abundance, diversity and/or taxonomic composition were selected from several widely used indices in ecological studies and/or developed for soft-bottom macroinvertebrate communities.Present findings reveal several indices based on diversity and/or taxonomic composition able to distinguish sites within the disturbance gradients, showing increasing quality from the site nearest the source of organic enrichment to that farthest from it, especially indices calculated using biomass data, and in the summer season. Such results open good perspectives for the use of intertidal macroinvertebrate communities from rocky shores, and also help add the perspective of this biological quality element in the ecological quality assessment of coastal waters.     

Ants and plants as indicators of biodiversity,ecosystem services,and conservation value in constructed grasslands

Grasslands are constructed for soil and wildlife conservation in agricultural landscapes across Europe and North America. Constructed grasslands may mitigate habitat loss for grassland-dependent animals and enhance ecosystem services that are important to agriculture. The responses of animal species richness and abundance to grassland habitat quality are often highly variable, however, and monitoring of multiple taxa is often not feasible. We evaluated whether multiple animal taxa responded to variation in constructed grassland habitats of southwest Ohio, USA, in ways that could be predicted from indicators based on quality assessment indices, Simpson diversity, and the species richness of ants and plants. The quality assessment indices included a widely used Floristic Quality Assessment (FQA) index, and a new Ant Quality Assessment (AntQA) index, both based on habitat specificity and species traits. The ant and plant indicators were used as predictor variables in separate general linear models of four target taxa—bees, beetles, butterflies and birds—with response variables of overall species richness and abundance, and subsets of taxa that included the abundance of ecosystem-service providers and grassland-associated species. Plant Simpson diversity was the best-fitting predictor variable in models of overall bee and beetle abundance, and the abundance of bees classified as ecosystem-service (ES) providers. FQA and plant richness were the best predictors of overall butterfly species richness and abundance. Ant species richness was the best predictor of overall bird species richness and abundance as well as the abundance of ES birds, while the AntQA index was the best predictor for the abundance of grassland bird and butterfly species. Thus, plant Simpson diversity and ant species richness were the most effective indicators for complementary components of grassland animal communities, whereas quality assessment indices were less robust as indicators and require more knowledge on the habitat specificity of individual ant and plant species.     

Response of macroalgae and macroinvertebrates to anthropogenic disturbance gradients in rocky shores

The compliance of macroalgal and macroinvertebrate assemblages to anthropogenic disturbance gradients (e.g., nutrient enrichment) was investigated at intertidal rocky shores. Macroalgae and macroinvertebrates presented parallel behavior, both showing shifts in the communities’ structural variation along the gradients, in which an higher number of opportunistic species (and higher abundances) were found in more stressful sites (close to the disturbance source), in contrast to less disturbed sites (far from the disturbance source), which showed higher presence of more sensitive species (and higher abundance of several of them).The macroinvertebrate abundance and taxonomic composition, which are parameters required by the Water Framework Directive (WFD) to be included in tools for the ecological quality status assessment, responded to the disturbance gradient. Results suggest that the macroinvertebrate biological element might be considered an indicator of disturbance in intertidal rocky shores as good as the macroalgae, and therefore the development of a specific methodology based solely on benthic macroinvertebrates of rocky shores, presently a gap in the ecological quality status assessment for the WFD, seems feasible.     

Transitional and coastal waters ecological status assessment: advances and challenges resulting from implementing the European Water Framework Directive

The derivation, performance, sensitivity and inherent uncertainty of ecological quality indicators have become major topics in developing tools for the management of marine, transitional and coastal waters. In reviewing the advances in these waters, related to an ecological status assessment, we show the future challenges to be addressed within the European Water Framework Directive (WFD). Using new analyses carried out under the research project ‘Water Bodies in Europe: Integrative Systems to Assess Ecological status and Recovery’, we provide a complete set of assessments for the biological quality elements (BQEs) (phytoplankton, macroalgae-seagrasses, macroinvertebrates and fish) to be assessed, as well as the validation of existing indicators and multimetric indices and, in some cases, the development of new assessment indices. We show that these indices respond differently to different human pressures and they each have challenges in defining reference conditions against which future changes are judged. In investigating good ecological potential, as the response to heavily modified water bodies, we show that there are flaws in the Directive, not least in its definitions. Our analyses have also focussed on uncertainty in using the indices and we emphasise the problems of defining ecological class boundaries based on indices which themselves may be combined indices (multimetrics). The analysis shows that some of those multimetrics are redundant and/or are inter-correlated and thus may reduce the sensitivity in defining ecological class boundaries. If this is related to the drivers-pressures-state change-impacts-response approach then there are lessons for management measures aimed at achieving good ecological status and even the potential for legal challenges to decisions based on uncertain indices under the WFD. Hence, we conclude the continued need for advances in assessing pressures and gradients, and defining reference conditions for state change, index development, impact assessment and the validation of indices for each BQE.     

An appraisal of a biocontamination assessment method for freshwater macroinvertebrate assemblages; a practical way to measure a significant biological pressure?

Freshwater invasive or alien species (IAS) can have a major impact on benthic macroinvertebrate assemblage structure and diversity. This has implications for accurate biological monitoring, the assessment of the ecological quality status of rivers and achievement of Water Framework Directive (WFD) objectives. Although IAS constitutes a major biological pressure to WFD objectives, current approaches to ecological status assessment tend to ignore their presence. This problem is compounded as biotic indices such as the Biological Monitoring Working Party (BMWP) score do not distinguish between native and IAS, when IAS tend to be more tolerant of organic pollution than the natives they replace. Biocontamination is the presence of an IAS in a system, and we tested a new method of biocontamination assessment, designed to be used alongside current routine water quality monitoring techniques, by applying it to biological monitoring data from the river monitoring programme of a small Island, The Isle of Man. Although 54% of monitoring sites exhibited no biocontamination, 19% showed low or moderate biocontamination and 27% high or severe biocontamination. Richness contamination was low (only two contaminated families being recorded), but abundance contamination was high in some sites (87% of individuals being IAS). Sites with a greater relative abundance of IAS individuals exhibited lower BMWP water quality. Within invaded sites BMWP monitoring was not responsive to changing chemical water quality, whereas within uninvaded sites it was. In invaded sites, the relative abundance of IAS increased as ammonia and BOD₅ increased. Our study shows current monitoring approaches mask the presence of AIS within assemblages, with some highly biocontaminated sites registering high BMWP biological quality. This new index represents a simple way to integrate the IAS biological pressure into established WFD monitoring programmes, to produce more comprehensive estimates of ecological quality status than are currently being realised.     

A comparative approach of assessing ecological status in two coastal areas of Eastern Mediterranean

This work presents an assessment of the ecological quality status of two marine coastal areas in the Aegean Sea (Greece, Eastern Mediterranean) based on the benthic macroinvertebrate quality element. S. Evvoikos and Thessaloniki gulfs, two coastal areas subjected to slight and heavier anthropogenic pressures, were selected to test the biotic index Bentix developed for the assessment of the ecological status. Other ecological indicators, such as the Shannon diversity index (H′), the species richness (S) and the AMBI biotic index were also applied and evaluated comparatively. Faunistic data were also used to interpret results. The resulting classification was validated with the use of physicochemical parameters and pressure information. This work also provides an insight into the structure of the Bentix classification scheme within the scope of its use in Water Framework Directive (WFD) implementation.     

Effects of pollution on macroinvertebrates and water quality bio-assessment

Many large rivers in China have an inflow of contaminated water. Water pollution caused by urban sewage and agriculture, and occasionally pollution events from industries have become a significant stress on aquatic ecology. Pollution affects the biodiversity of the aquatic community and the species composition changes from natural species to tolerant species. The species composition of aquatic animals may reflect water pollution level. Extremely non-uniform distributions of functional feeding groups occurred as a result of high nutrient levels. A combination of chemical and biological methods constitutes the best approach for biological monitoring studies that measure water quality. Macroinvertebrates were used as bio-assessment indicator to determine the environmental quality of given water body. In this study, samples of water and macroinvertebrates were taken from several dozen sites in 14 rivers in China with different pollution levels, including the Yangtze, East, Weihe, Songhua, Yongding, and Panlong rivers. Macroinvertebrates were identified to genus or family level. Water samples were classified into different water quality grades according to the concentration of different substances. Five biological indices: taxa richness (S), density (D), total BMWQ score (t-BMWQ), average BMWQ score (a-BMWQ), and the family biotic index (FBI) were used for biological assessment of water quality. Analyzing macroinvertebrates’ occurrence in different water quality levels, taxa-specific indicators, which are defined as the taxa of macroinvertebrates that live in a certain water quality level but do not exist in other water quality levels were proposed for water quality bio-assessment. Leptophlebiidae, Siphlonuridae, Arctopsychidae, Perlidae, and Antocha sp. are the taxa-specific indicators for very good or good water quality; Chironomidae, Lymnaeidae, Tubifex sp., Limnodrilus sp., Limnoperna lacustris, Corbicula sp., Macrobrachium sp., Planorbidae, Glossiphoniidae, and Branchiura sp. are the taxa-specific indicators for very poor water quality; and Psychomyiidae and Hydroptilidae are the taxa-specific indicators for moderate water quality.     

Multi-scale approach using phytoplankton as a first step towards the definition of the ecological status of reservoirs

The growing need to analyse the present state of ecosystems and predict their rate of change has triggered a demand to explore species environment relationships for assessing alterations under anthropogenic influence. The Water Framework Directive (WFD) requires the definition of different types of water bodies which are of relevance when assessing their ecological status. The main aim of this study was to define of the types of Portuguese reservoirs located in the North and Centre of Portugal and to assess their ecological status using phytoplankton as water quality indicators. In this study, sampling was carried out in 34 reservoirs during four seasons (spring, summer, autumn and winter), through a period of 8 years (1996–2004).Two groups of reservoirs could be distinguished, from the multivariate statistical analysis based on environmental variables and on phytoplankton assemblages: G1, lowland reservoirs located in the main rivers (Douro and Tagus), with a very low residence time, characterized by higher water mineral content (hardness and conductivity), higher concentrations of nutrients (namely, nitrates), dominated by Bacillariophyta and Chlorophyta and characterized by the presence of tolerant of poor environmental conditions species, mainly associated with meso and eutrophic states of water bodies; G2, deeper high altitude reservoirs, largely located in tributaries, with high residence time, presenting a specific species composition under reference conditions, with higher species richness. The transition from deeper and colder reservoirs (reference sites) to shallow and warmer reservoirs (impaired sites), was evident in G2, contrarily to G1, and was mostly positively correlated to organic pollution and mineral gradients. The results presented here are fundamental for the development of a routine for monitoring ecological status according to the WFD.     

Influence of sampling effort on metrics of fish-based indices for the assessment of estuarine ecological quality

Multimetric fish-based indices have been increasingly gaining importance in Europe, as the Water Framework Directive (WFD) requires fish fauna, and particularly its composition and abundance, to be taken into account in the assessment of the ecological quality of continental surface waters, including transitional waters. These indices are composed of several metrics, mostly related with structural and functional characteristics of fish communities, such as species richness, the role of nursery areas, or trophic web structure. Therefore, ecological quality assessments should ensure that these structural and functional characteristics of fish communities were covered by the sampling methods used. In the present work, the influence of sampling effort on several metrics of the Estuarine Fish Assessment Index (EFAI) was studied. Pseudo-random samples were generated from data of four Portuguese estuaries and bootstrap cycles were performed, in order to obtain metrics’ means and standard deviations per number of hauls analysed. The number of hauls necessary for the means to level off differed with the metrics considered. Generally, for metrics on percentages (percentage of marine migrants, percentage of estuarine residents and percentage of piscivores) the curve levelled off with less than 20 hauls, both for the estuary as a whole and for different estuarine salinity zones. On the other hand, metrics on species richness required much larger samples. In order to decrease to −5% the current estimated bias of metrics, the WFD sampling costs would have to be more than 3 times higher than they currently are. The findings in the present study are of great importance for an effective assessment of estuarine ecological quality and particularly in the context of the WFD, as the metrics studied are common to other Member State indices.     

Diversity measures in macroinvertebrate and zooplankton communities related to the trophic status of subtropical reservoirs: Contradictory or complementary responses?

Macroinvertebrate communities have been widely used as a tool for assessing the environmental quality of freshwater ecosystems, whereas zooplankton communities have been to some extent neglected. However, the importance of using different indicators to achieve a more comprehensive framework of assessment regarding water quality has been recognized. This study compared estimates of species richness (number of species) and the Shannon–Wiener index for data on macroinvertebrate and zooplankton communities in tropical reservoirs and related them to their trophic state. The trop+hic classification was obtained by applying the Carlson index (1977) modified by Toledo et al. (1983), and the index of the Brazilian Society of the Environmental Technology Agency. The comparative response of the different indicators was analyzed using a series of bivariate correlations (Draftsman’s plot). The results illustrate that diversity measures, namely species richness, responded differently when related to the trophic classification of reservoirs, depending on the community considered. The species richness of zooplankton was positively related to hypereutrophic conditions, due to the higher number of rotifer species, including tolerant generalist species and at the same time, as a result of the exclusion of species from other groups, whereas for macroinvertebrates, species richness was negatively related to hypereutrophic conditions. Melanoides tuberculatus, which exhibits a high tolerance and competitive ability under such conditions, was the dominant species in macroinvertebrate communities, which excluded endemic species and reduced local richness and diversity. The same indicators applied to the zooplankton and macroinvertebrate communities might therefore provide contradictory responses regarding ecological quality assessment in tropical reservoirs, which suggest that zooplankton should be taken into account among the biological quality elements considered in the ecological quality assessment, management, and restoration of water bodies.     

Assessing ecological quality of shallow lakes: Does knowledge of transparency suffice?

The European Water Framework Directive (WFD) requires that all aquatic ecosystems in their member states should reach ‘good’ ecological quality by 2015. To assess ecological quality, the WFD requires the definition of reference conditions using biological, physical and chemical indicators and the assignment of each water body to one of five quality classes using these indicators. Elaborate assessment schemes using large sets of variables are now being developed. Here we address the question whether all this is really needed and what the simplest assessment approach would be for the case of shallow lakes. We explore the relationships between the quality class assigned to a lake by experts in shallow lake ecology and a rich set of biological, physical, and chemical data. Multinomial logistic regression analyses were carried out based on data from 86 shallow lakes throughout Europe that were sampled in 2000 and/or 2001. Ecological quality of shallow lakes judged by experts was strongly correlated to physical and chemical variables associated with light regime and nutrients and much less to biological variables.Our regression model showed that ecological quality of this set of shallow lakes judged by experts could be predicted quite well from water transparency expressed as Secchi depth and that other variables did not contribute to it significantly. According to the WFD, lakes should at least have a ‘good’ ecological quality. Quality judged by experts and predicted quality were similar for 78% of the lakes with respect to meeting this standard. As a cautionary note we stress that Secchi depth alone will be a less useful indicator if effects of stressors other than eutrophication (e.g. lake acidification and toxic pollution) are to be considered.     

Macrophytes as indicators of stream condition in the wet tropics region,Northern Queensland,Australia

This study investigates the use of aquatic macrophytes as indicators of stream condition in catchments with varied land use and levels of riparian disturbance in the Wet Tropics region of North Queensland (Australia), a region of global significance in terms of faunal and floral diversity. In a paired catchment design spatial variations in macrophyte assemblage structure were characterised using multivariate and univariate techniques. Seven metrics were trialled: total macrophyte cover, species richness, % alien taxa, % native taxa, % submerged taxa, % emergent taxa and % Poaceae. Forty-four macrophyte taxa were recorded from the study area. Poaceae, Cyperaceae and mosses were the most frequently recorded taxa. Upper catchment areas in all tributaries surveyed were dominated by mosses and Cladopus queenslandicus (Domin) C.D.K. Cook (Podestemaceae). This assemblage occurred in areas with intact riparian canopy cover and good overall riparian condition. Macrophyte assemblages in lower catchment areas were distributed along gradients of riparian disturbance. Simultaneous autoregression model coefficients indicated that riparian condition had a negative influence on macrophyte cover, species richness and the proportions of alien taxa, emergent taxa and Poaceae present at sites in the Wet Tropics. Macrophyte metrics were not strongly influenced by the types of land use or water quality. These findings suggest that a riparian condition assessment would provide an adequate first assessment of the state of aquatic macrophyte assemblages in Wet Tropics streams.     

Richness‐dependence of phylogenetic diversity indices

Phylogenetic diversity indices are widely used to characterize the structure and diversity of ecological communities. Most indices are based on a metric that is expected to vary with species richness, so they are standardized to remove this richness‐dependence. With this standardization, values of 0 are consistent with random phylogenetic structure, while phylogenetic clustering is associated with either negative or positive values (depending on the index). One common interpretation of phylogenetic clustering is that it indicates some combination of environmental and biological filtering that restricts the species that can be present in a community. Increasingly, studies are comparing phylogenetic indices along environmental gradients to infer differences in the factors structuring communities. This comparison implicitly assumes that index values are comparable among communities with different numbers of species. Using a set of simulations, I show here that this assumption is incorrect. Holding the strength of filtering constant, communities composed of more species show larger absolute index values. This problem is most pronounced when the environmental filter favors a moderate‐sized clade strongly over others and when using the net relatedness index (NRI) to measure clustering. This bias potentially casts doubt on studies studying phylogenetic index patterns along gradients where richness also varies. Fortunately, the arising generality that more stressful environments have lower species richness and stronger clustering is opposite to this bias and therefore robust. I also show that a simple rarefaction can remove the richness‐dependence of these indices, at the expense of increased error.     

Central European water quality indices applied to long-term data from peri-alpine lakes: test and possible improvements

We tested the applicability of two available Water Framework Directive (WFD) compliant phytoplankton indices on three large peri-alpine lakes to analyze their suitability for trophic classification. We show that the indices vary in their final resulting Ecological Quality Ratios (EQRs) and are only appropriate to roughly distinguish lakes of different water quality according to OECD criteria (OECD, 1982). Annual mean TP concentration in Lake Geneva was 0.03 mg l^?1 in 2006 (Lazzarotto &; Rapin, 2007), which marks mesotrophic conditions. According to the tested indices, the lake is of ‘good’ (0.60–0.80) quality after the German WFD method (PSI) and of ‘moderate’ quality according to the Austrian WFD method (BI). We prove that the way how to derive the per annum value of the metric ‘Brettum Index’ (BI) in the Austrian WFD method significantly influences the resulting index and the scatter of long-term data. We focused on improvement strategies for this index in terms of calculation and sampling frequency. Contrary to the tested bin, the modified index presents no significant differences between four and six sampling dates per year. In order to improve the significance of the available indices, we propose to modify the way of per annum index calculation for the Austrian WFD method as well as to focus on species composition to achieve a high relative proportion of indicator species. A phytoplankton index alone may not be sensitive enough to track the changes that occur within a lake. The results confirm the need to take into account other biological elements such as fish, macrophytes, attached diatoms and macrozoobenthos as suggested by the WFD.     

Benthic indices and ecological quality of shallow Algeria fine sand community

This study applies six macrozoobenthos-based biotic indices in the shallow coastal waters along the Algerian coast (southern Mediterranean Sea) to establish a reference situation for future use. These shallow fine sand communities were sampled in seven bays along the Algerian coast during the 1980s and the beginning of the 1990s. For the first time, some of the benthic indices used nowadays in Europe for the implementation of the WFD and/or in North America for the Clean Water Act were used to analyze the data collected twenty years ago in order to assess the biological quality status of the Algerian shallow fine sand communities and to provide ecological classifications for the fine sand community along the Algerian coast. The faunal composition showed that the fine sand communities belonged to a Well-Calibrated Fine Sand (WCFS) biocenosis characterized by high species richness and high H′ Shannon-Weaver diversity, with moderate abundance levels only. In the bays of Fetzara, Jijel, Bejaia, Bou Ismail and Arzew (though not in bays of Algiers and Oran), H′ is >4.5, which implies highly diversified communities. The results of the six benthic indices (H′, AMBI, M-AMBI, BENTIX, BOPA and ITI) suggest that good and high quality status assessments are prevalent in all seven bays for the benthic shallow sand communities along the Algerian coast. Nevertheless, the effect of pollution was observed at a small number of sites in the Arzew, Oran and Algiers bays. For example, in Algiers Bay, an east-west quality gradient revealed the effect of organic matter input.     

Response of epilithic diatom assemblages to water pollution in rivers in the Tokyo Metropolitan area, Japan

1. During the spring of 1992, fifty-two quantitative diatom samples were collected from twenty-eight rivers located in the Tokyo Metropolitan area, Japan, to study the response of the diatom assemblages to water pollution (assessed using physical and chemical data determined monthly from April 1987 to March 1992). 2. Species composition was analysed by means of biotic indices (Pantle and Buck's saprobic index) and multivariate analyses [two-way indicator species analysis (TWINSPAN) for classification and canonical correspondence analysis (CCA) for ordination]. Species-abundance relationships were analysed using diversity indices (species richness, Shannon's diversity index and Pielou's evenness index) and rank-abundance patterns (rank-abundance curves). 3. CCA revealed two major gradients. The first corresponded to organic pollution and eutrophication. The second corresponded to variables related to geographical location. Four main station groups were determined by TWINSPAN. The location of the indicator species of groups 1–3 along the CCA axis 1 is consistent with their known pollution tolerance characteristics. Indicator species for group 4 had larger scores on CCA axis 2, and are representative of brackish water environments. 4. Species richness tended to be higher in the intermediate range of water pollution. Pielou's evenness index and Shannon's diversity index followed the same tendency but only weakly. 5. The rank-abundance patterns of diatom assemblages were more or less constant in all stations. The curves were very similar in shape, differing only in length and gradient (directly related to species richness and evenness, respectively). 6. The results of this study indicate that the response of diatom assemblages to environmental change can be observed in species compositional variation. Multivariate analyses and pollution indices revealed this response and are to be preferred to species diversity measures.     

Do biotic interactions modulate ecosystem functioning along stress gradients? Insights from semi-arid plant and biological soil crust communities

Climate change will exacerbate the degree of abiotic stress experienced by semi-arid ecosystems. While abiotic stress profoundly affects biotic interactions, their potential role as modulators of ecosystem responses to climate change is largely unknown. Using plants and biological soil crusts, we tested the relative importance of facilitative–competitive interactions and other community attributes (cover, species richness and species evenness) as drivers of ecosystem functioning along stress gradients in semi-arid Mediterranean ecosystems. Biotic interactions shifted from facilitation to competition along stress gradients driven by water availability and temperature. These changes were, however, dependent on the spatial scale and the community considered. We found little evidence to suggest that biotic interactions are a major direct influence upon indicators of ecosystem functioning (soil respiration, organic carbon, water-holding capacity, compaction and the activity of enzymes related to the carbon, nitrogen and phosphorus cycles) along stress gradients. However, attributes such as cover and species richness showed a direct effect on ecosystem functioning. Our results do not agree with predictions emphasizing that the importance of plant–plant interactions will be increased under climate change in dry environments, and indicate that reductions in the cover of plant and biological soil crust communities will negatively impact ecosystems under future climatic conditions.