Jani Heino

Climate change and freshwater biodiversity: detected patterns, future trends and adaptations in northern regions

Current rates of climate change are unprecedented, and biological responses to these changes have also been rapid at the levels of ecosystems, communities, and species. Most research on climate change effects on biodiversity has concentrated on the terrestrial realm, and considerable changes in terrestrial biodiversity and species’ distributions have already been detected in response to climate change. The studies that have considered organisms in the freshwater realm have also shown that freshwater biodiversity is highly vulnerable to climate change, with extinction rates and extirpations of freshwater species matching or exceeding those suggested for better‐known terrestrial taxa. There is some evidence that freshwater species have exhibited range shifts in response to climate change in the last millennia, centuries, and decades. However, the effects are typically species‐specific, with cold‐water organisms being generally negatively affected and warm‐water organisms positively affected. However, detected range shifts are based on findings from a relatively low number of taxonomic groups, samples from few freshwater ecosystems, and few regions. The lack of a wider knowledge hinders predictions of the responses of much of freshwater biodiversity to climate change and other major anthropogenic stressors. Due to the lack of detailed distributional information for most freshwater taxonomic groups and the absence of distribution‐climate models, future studies should aim at furthering our knowledge about these aspects of the ecology of freshwater organisms. Such information is not only important with regard to the basic ecological issue of predicting the responses of freshwater species to climate variables, but also when assessing the applied issue of the capacity of protected areas to accommodate future changes in the distributions of freshwater species. This is a huge challenge, because most current protected areas have not been delineated based on the requirements of freshwater organisms. Thus, the requirements of freshwater organisms should be taken into account in the future delineation of protected areas and in the estimation of the degree to which protected areas accommodate freshwater biodiversity in the changing climate and associated environmental changes.

Lentic macroinvertebrate assemblage structure along gradients in spatial heterogeneity, habitat size and water chemistry

Littoral zones of small water bodies are spatially heterogeneous habitats, harbouring diverse biotic communities. Despite this apparent heterogeneity, many studies have stressed the importance of water chemistry in determining the structure of littoral macroinvertebrate assemblages. The purpose of this study was to consider the relative importance of several spatial and water chemistry variables in explaining the patterns in the structure of macroinvertebrate assemblages in 21 lentic water bodies in northeastern Finland. Water bodies were selected to represent various habitat conditions ranging from small permanent bog ponds to small forest lakes. According to canonical correspondence analysis (CCA), the most important environmental factors related to assemblage composition were water body area, moss cover, total nitrogen and water hardness. In general, species composition in small bog ponds tended to differ from that in larger lakes with forested shoreline. Total species richness was best explained by a composite variable (PCA) describing physical habitat heterogeneity, species richness being lowest in small bog lakes with simple bottom structure and low amount of aquatic plants. Species numbers in dominant functional feeding groups were related to different environmental factors. Shredder species richness was best explained by a regression model incorporating total nitrogen and the amount of organic matter, both of which were negatively related to the number of shredder species. The number of gatherer species increased with mean substratum particle size. Scraper species richness was negatively affected by the abundance of detritus and positively affected by depth, and a model including both variables explained most of the variation. Variation in the number of predatory species was best explained by a regression model including moss cover and lake area.

The importance of metacommunity ecology for environmental assessment research in the freshwater realm.

Most bioassessment programs rest on the assumption that species have different niches, and that abiotic environmental conditions and changes therein determine community structure. This assumption is thus equivalent to the species sorting perspective (i.e. that species differ in their responses to environmental variation) in metacommunity ecology. The degree to which basing bioassessment on the species sorting perspective is reasonable is likely to be related to the spatial extent of a study and the characteristics of the organism groups (e.g. dispersal ability) with which the effects of anthropogenic changes are assessed. Recent findings in metacommunity research have stressed that community structure is determined not only by local abiotic environmental conditions but also by biotic interactions and dispersal‐related effects. For example, dispersal limitation may prevent community structure recovery from the effects of a putative stressor, as organisms may not be able to disperse to all sites in a region. Mass effects (i.e. the presence of species in environmentally suboptimal sites due to high dispersal rates from environmentally suitable sites) may, in turn, obscure the effects of a stressor, as dispersal from source sites (e.g. an unaltered site) allows persistence at sink sites (e.g. an anthropogenically altered site). Better bioassessment should thus take both niche‐ and dispersal‐related processes simultaneously into consideration, which can be accomplished by explicitly modelling spatial location as a proxy for dispersal effects. Such an integrated approach should be included in bioassessment programs using general multivariate approaches, predictive modelling, and multimetric indices.

Correspondence between regional delineations and spatial patterns in macroinvertebrate assemblages of boreal headwater streams

Geographical stratification may provide a useful framework for stream management programs, yet most studies testing the utility of such stratifications have been conducted in temperate regions. We studied the correspondence between regional delineations (5 ecoregions, 11 subecoregions), environmental characteristics, and benthic macroinvertebrate assemblages in 156 boreal headwater streams in Finland, using a combination of principal components analysis, nonmetric multidimensional scaling, and discriminant function analysis (DFA). Both stream characteristics and macroinvertebrate assemblage structure showed a closer correspondence to ecoregions than to subecoregions, a pattern partly attributable to the smaller number of classes at the ecoregion level. Based on environmental variables, DFA classified correctly 84.0% of the streams by ecoregion and 74.4% by subecoregion. For macroinvertebrate assemblages, 64.7% of the streams were correctly classified at the ecoregion, and 51.9% at the subecoregion level. Arctic-alpine and north boreal ecoregions supported the most distinct benthic assemblages. There were significant differences among ecoregions in mean species number, with lowest species richness in the middle boreal and highest in the south boreal streams. However, no clear latitudinal gradients in local species richness were detected. Relative abundances of functional-feeding groups also exhibited regional differences. Shredders dominated the middle boreal brown-water streams, whereas scrapers attained their highest abundances in the arctic-alpine and north boreal clear-water streams. Despite some congruence among ecoregions and macroinvertebrate assemblages, regional delineations alone do not provide a suitable background for stream biomonitoring programs. A combination of regional delineations and local characteristics probably provides a better framework for the classification of boreal headwater streams, and for the prediction of their benthic fauna.

Concordance of species richness patterns among multiple freshwater taxa: a regional perspective

Geographical gradients in species richness and the degree to which different taxa show congruent patterns remain unknown for many taxonomic groups. Here, I examined broad-scale species richness patterns in five groups of freshwater organisms; macrophytes, dragonflies, stoneflies, aquatic beetles and fishes. The analyses were based on provincial distribution records in Denmark, Norway, Sweden and Finland. In general, variation in species richness across provinces was concordant among the groups, but stoneflies showed weaker negative relationships with the other taxonomic groups. Species richness in most groups decreased with increasing latitude and altitude, and a considerable part of the variation was explained by mean July temperature. However, stoneflies showed a reversed pattern, with species richness correlating positively, albeit more weakly, with mean provincial altitude. Nevertheless, combined species richness of all five taxa showed a strong relationship with mean July temperature, accounting for 74% of variation in provincial species richness alone. Such temperature-controlled patterns suggest that regional freshwater biodiversity will strongly respond to climate change, with repercussions for local community organization in freshwater ecosystems in Fennoscandia.

Metacommunity structuring in stream networks: roles of dispersal mode, distance type, and regional environmental context

Within a metacommunity, both environmental and spatial processes regulate variation in local community structure. The strength of these processes may vary depending on species traits (e.g., dispersal mode) or the characteristics of the regions studied (e.g., spatial extent, environmental heterogeneity). We studied the metacommunity structuring of three groups of stream macroinvertebrates differing in their overland dispersal mode (passive dispersers with aquatic adults; passive dispersers with terrestrial adults; active dispersers with terrestrial adults). We predicted that environmental structuring should be more important for active dispersers, because of their better ability to track environmental variability, and that spatial structuring should be more important for species with aquatic adults, because of stronger dispersal limitation. We sampled a total of 70 stream riffle sites in three drainage basins. Environmental heterogeneity was unrelated to spatial extent among our study regions, allowing us to examine the effects of these two factors on metacommunity structuring. We used partial redundancy analysis and Morans eigenvector maps based on overland and watercourse distances to study the relative importance of environmental control and spatial structuring. We found that, compared with environmental control, spatial structuring was generally negligible, and it did not vary according to our predictions. In general, active dispersers with terrestrial adults showed stronger environmental control than the two passively dispersing groups, suggesting that the species dispersing actively are better able to track environmental variability. There were no clear differences in the results based on watercourse and overland distances. The variability in metacommunity structuring among basins was not related to the differences in the environmental heterogeneity and spatial extent. Our study emphasized that (1) environmental control is prevailing in stream metacommunities, (2) dispersal mode may have an important effect on metacommunity structuring, and (3) some factors other than spatial extent or environmental heterogeneity contributed to the differences among the basins.

Searching for biodiversity indicators in running waters: do bryophytes, macroinvertebrates, and fish show congruent diversity patterns?

The degree to which different taxonomic groups show congruence in diversity patterns has attracted increased attention, yet such studies on stream biota are lacking. We examined environmental correlates of and congruence in the species richness patterns of bryophytes, macroinvertebrates, and fish in 101 boreal streams in Finland. Congruence in species richness among the taxonomic groups was generally low, mainly because of their differing responses to major environmental gradients. Bryophytes and macroinvertebrates showed the strongest degree of congruence, but even this relationship had a relatively weak predictive power. Bryophyte diversity showed the strongest relationship with water colour, followed by habitat stability, and stream size. Macroinvertebrate diversity increased with stream size, and further variation was accounted for by water colour and acidity. Fish species richness showed a weak and complex relationship with geographical location, stream size, and in-stream habitat characteristics. The regression models explained 23, 45, and 26% of the variation in species richness of bryophytes, macroinvertebrates, and fish, respectively. Our results suggest that indicator taxa may be of limited value in stream biodiversity inventories. Habitat-based approaches are suggested as an alternative surrogate measure in the conservation evaluation of lotic biodiversity.

DEFINING MACROINVERTEBRATE ASSEMBLAGE TYPES OF HEADWATER STREAMS: IMPLICATIONS FOR BIOASSESSMENT AND CONSERVATION

We used data from 235 boreal headwater streams in Finland to examine whether macroinvertebrate assemblages constitute clearly definable types, and how well biologically defined assemblage types can be predicted using environmental variables. Two- way indicator species analysis produced 10 assemblage types, which differed significantly from each other (multiresponse permutation procedure, MRPP). However, based on MRPP and nonmetric multidimensional scaling, there was wide variation among sites within each assemblage type, and high degrees of overlap among assemblage types. Such continuous variation was also evidenced by the low number of effective indicator taxa (indicator value method) for each assemblage type. Furthermore, discriminant function analysis based on environmental variables showed a moderate yet variable prediction success (59.6% of sites correctly predicted; range 0-96% per group). Canonical correspondence analysis indicated that variation in assemblage structure was primarily related to latitude and water chemistry, especially pH and water color. The absence of discrete macroinvertebrate assemblage types in boreal headwater streams may stem from a number of reasons: (1) macroinvertebrate taxa inhabiting boreal streams tend to exhibit individualistic, taxon-specific responses to environmental gradients, (2) they have wide environmental tolerances and geographic dis- tributions, and (3) boreal streams are characterized by frequent, unpredictable disturbances. Our results suggest that local filters in headwater streams are relatively weak, resulting in poorly distinguishable assemblage types. By contrast, the major latitudinal gradients in macroinvertebrate assemblage structure suggest that regional-scale filters, e.g., temperature, exert strong control over taxon distributions in headwater streams. We suggest that a tiered approach combining regional stratification and subsequent prediction of assemblage struc- ture could provide a suitable framework for the bioassessment and conservation of boreal headwater streams.

Control of stream insect assemblages: roles of spatial configuration and local environmental factors

Abstract 1. Current views in ecology emphasise that community structure is the sum of multiple processes, with imprints of both regional and local drivers. However, the degree to which stream insect assemblages are structured by spatial configuration (complying with the dispersal‐based neutral hypothesis) and local environmental features (complying with the niche‐based species sorting hypothesis) has not been rigorously examined based on surveys in multiple years.

A comparative analysis reveals weak relationships between ecological factors and beta diversity of stream insect metacommunities at two spatial levels

The hypotheses that beta diversity should increase with decreasing latitude and increase with spatial extent of a region have rarely been tested based on a comparative analysis of multiple datasets, and no such study has focused on stream insects. We first assessed how well variability in beta diversity of stream insect metacommunities is predicted by insect group, latitude, spatial extent, altitudinal range, and dataset properties across multiple drainage basins throughout the world. Second, we assessed the relative roles of environmental and spatial factors in driving variation in assemblage composition within each drainage basin. Our analyses were based on a dataset of 95 stream insect metacommunities from 31 drainage basins distributed around the world. We used dissimilarity-based indices to quantify beta diversity for each metacommunity and, subsequently, regressed beta diversity on insect group, latitude, spatial extent, altitudinal range, and dataset properties (e.g., number of sites and percentage of presences). Within each metacommunity, we used a combination of spatial eigenfunction analyses and partial redundancy analysis to partition variation in assemblage structure into environmental, shared, spatial, and unexplained fractions. We found that dataset properties were more important predictors of beta diversity than ecological and geographical factors across multiple drainage basins. In the within-basin analyses, environmental and spatial variables were generally poor predictors of variation in assemblage composition. Our results revealed deviation from general biodiversity patterns because beta diversity did not show the expected decreasing trend with latitude. Our results also call for reconsideration of just how predictable stream assemblages are along ecological gradients, with implications for environmental assessment and conservation decisions. Our findings may also be applicable to other dynamic systems where predictability is low.