Riku Paavola

Long-term recovery of stream habitat structure and benthic invertebrate communities from in-stream restoration

Headwater streams channelized for water transport of timber in Finland are being restored to their pre-channelization state. The primary motivation is the enhancement of sport fisheries, but restoration probably has profound impacts also on other stream organisms. We assessed how such ‘‘single-goal’’ restorations affect benthic macroinvertebrate communities. We revisited the streams sampled by Laasonen et al. [Aquatic Conservation: Marine and Freshwater Ecosystems 8 (1998)] in the early 1990s when the streams had been recently restored. In 1997, the recovery period of these streams ranged from 4 to 8 years. Habitat structure among the stream types represented a distinct recovery gradient, with streams restored 1 month before sampling and natural streams being the endpoints of a gradient in moss cover (highest in natural, lowest in recently restored streams). Channelized streams supported a characteristic set of indicator species, whereas shifts in species composition between restored and natural streams were more gradual. Macroinvertebrate communities in unmodified streams changed little between the two surveys, whereas communities in restored streams had undergone considerable changes. In-stream restoration is an unpredictable disturbance, to which stream biota cannot have any evolved responses. Therefore, the relatively rapid recovery of habitat structure and macroinvertebrate communities in restored streams is encouraging. However, long-term monitoring of benthic communities in both restored and natural streams is needed to assess whether restoring rivers by these techniques will enhance the recovery of benthic biodiversity in boreal streams. # 2002 Elsevier Science Ltd. All rights reserved.

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.

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.

Twenty years of stream restoration in Finland: little response by benthic macroinvertebrate communities

The primary focus of many in-stream restoration projects is to enhance habitat diversity for salmonid fishes, yet the lack of properly designed monitoring studies, particularly ones with pre-restoration data, limits any attempts to assess whether restoration has succeeded in improving salmonid habitat. Even less is known about the impacts of fisheries-related restoration on other, non-target biota. We examined how restoration aiming at the enhancement of juvenile brown trout (Salmo trutta L.) affects benthic macroinvertebrates, using two separate data sets: (1) a before-after-control-impact (BACI) design with three years before and three after restoration in differently restored and control reaches of six streams; and (2) a space-time substitution design including channelized, restored, and near-natural streams with an almost 20-year perspective on the recovery of invertebrate communities. In the BACI design, total macroinvertebrate density differed significantly from before to after restoration. Following restoration, densities decreased in all treatments, but less so in the controls than in restored sections. Taxonomic richness also decreased from before to after restoration, but this happened similarly in all treatments. In the long-term comparative study, macroinvertebrate species richness showed no difference between the channel types. Community composition differed significantly between the restored and natural streams, but not between restored and channelized streams. Overall, the in-stream restoration measures used increased stream habitat diversity but did not enhance benthic biodiversity. While many macroinvertebrates may be dispersal limited, our study sites should not have been too distant to reach within almost two decades. A key explanation for the weak responses by macroinvertebrate communities may have been historical. When Fennoscandian streams were channelized for log floating, the loss of habitat heterogeneity was only partial. Therefore, habitat may not have been limiting the macroinvertebrate communities to begin with. Stream restoration to support trout fisheries has strong public acceptance in Finland and will likely continue to increase in the near future. Therefore, more effort should be placed on assessing restoration success from a biodiversity perspective using multiple organism groups in both stream and riparian ecosystems.

Characterizing macroinvertebrate assemblage structure in relation to stream size and tributary position

We examined the variability of macroinvertebrate assemblage structure, species identities, and functional feeding group composition in relation to stream size, tributary position, and in-stream factors in a boreal watershed in Finland. Our study included three riffle sites in each of three stream sections in each of three stream size classes. Multi-response permutation procedure, indicator value method, and canonical correspondence analysis revealed clear differences in assemblage structure among the stream size classes, with a gradual increase of species richness as the stream size increased. Significant differences in assemblage structure were also found among the tributary river systems. The functional feeding group composition broadly followed the river continuum concept, i.e., headwaters were dominated by shredders, gatherers, or filterers, whereas scrapers increased in relative abundance with stream size. There was, however, considerable variation in the functional feeding group composition both among and within the headwater stream sections. Our findings refer to a strong influence of stream size on macroinvertebrate assemblages, but also factors prevailing at the scale of individual riffles should be considered in biodiversity conservation of lotic ecosystems.

Habitat heterogeneity drives the geographical distribution of beta diversity: the case of New Zealand stream invertebrates

To define whether the beta diversity of stream invertebrate communities in New Zealand exhibits geographical variation unexplained by variation in gamma diversity and, if so, what mechanisms (productivity, habitat heterogeneity, dispersal limitation, disturbance) best explain the observed broad-scale beta diversity patterns. We sampled 120 streams across eight regions (stream catchments), spanning a north–south gradient of 12° of latitude, and calculated beta diversity (with both species richness and abundance data) for each region. We explored through a null model if beta diversity deviates from the expectation of stochastic assembly processes and whether the magnitude of the deviation varies geographically. We then performed multimodel inference analysis on the key environmental drivers of beta diversity, using Akaikes information criterion and model and predictor weights to select the best model(s) explaining beta diversity. Beta diversity was, unexpectedly, highest in the South Island. The null model analysis revealed that beta diversity was greater than expected by chance in all eight regions, but the magnitude of beta deviation was higher in the South Island, suggesting differences in environmental filtering and/or dispersal limitation between North and South Island. Habitat heterogeneity was the predominant driver of beta diversity of stream macroinvertebrates, with productivity having a secondary, and negative, contribution. This is one of the first studies accounting for stochastic effects while examining the ecological drivers of beta diversity. Our results suggest that local environmental heterogeneity may be the strongest determinant of beta diversity of stream invertebrates, more so than regional- or landscape-scale variables.

Macroinvertebrate community structure and species diversity in humic streams of Finnish Lapland

The majority of theories o n stream bemhic community structure idemifY site productivity and/or disturbance as the major factors underlying observed patterns (WARD & STANFORD 1983, HILDREW & ToWNSEND 1987, HILDREW & GJLLER 1994). Nevertheless, water chemistry is also known to influence species richness and composition of lotic macroinvertebrate communities. For example, the key role of acidity in regulating benthic assemblages has been documented repeatedly in both large-scale surveys (WRIGHT et al. 1984, Rurr et al. 1990) and in smaller-scale (catchmem-wide) studies (ToWNSEND et al. 1983, ÜRMEROD & EDWARDS 1987). Many other chemical factors have received much less attention, although MALMQVIST & MAKI (I 994) recendy found water colour to be strongly associated with the distribution of macroinvertebrate assemblages in northern Swedish streams. Water colour may contribute significandy to benthic community structure in boreal streams draining peadand areas. However, as pointed out by PETERSEN et al. (1995), studies on macroinvertebrate communities in boreal headwater streams are few, and the role of different factors in regulating these near-pristine communities are largely unknown. WARD (I 998) recendy urged for more investigations on natural river systems for the purposes of biodiversity assessment and conservation. Such studies provide background information for appropriately directed mitigation activities in damaged river systems. The aim of this study is to provide such information by idemifYing the major environmental gradients associated with stream macroinvertebrate communities in differendy sized streams in northeastern Finland.

Diatoms: unicellular surrogates for macroalgal community structure in streams?

As wholesale biodiversity assessment is often impractical, the use of surrogates that reflect the assemblage structure and diversity of other taxa has attracted increased attention. We sampled 47 boreal streams for diatoms and macroalgae and examined their assemblage patterns along major environmental gradients. Our main intention was to examine whether diatoms might be useful surrogates for macroalgae in boreal streams. We also assessed whether taxon richness and community composition provided similar insights into the patterns of cross-taxon concordance. According to canonical correspondence analysis, diatom distribution was most strongly related to water pH, conductivity, latitude and longitude, and macroalgal distribution to water pH and iron content, latitude and bed instability. In Mantel’s test, diatoms and macroalgae showed significant cross-taxon concordance. However, there was no significant correlation between taxon richness of the two algal groups, likely reflecting their differing responses to environmental variables. We found evidence that although diatoms and macroalgae are partly controlled by different environmental factors, they are segregated rather similarly along latitude and a few environmental gradients such as water pH and iron content. We conclude that, at least at broad geographical extents and in small streams, diatoms reflect the structure of the macroalgal community and are therefore useful surrogates for cost-effective biomonitoring of algal communities in streams.

Effects of changing climate on European stream invertebrate communities: a long-term data analysis

Long-term observations on riverine benthic invertebrate communities enable assessments of the potential impacts of global change on stream ecosystems. Besides increasing average temperatures, many studies predict greater temperature extremes and intense precipitation events as a consequence of climate change. In this study we examined long-term observation data (10-32years) of 26 streams and rivers from four ecoregions in the European Long-Term Ecological Research (LTER) network, to investigate invertebrate community responses to changing climatic conditions. We used functional trait and multi-taxonomic analyses and combined examinations of general long-term changes in communities with detailed analyses of the impact of different climatic drivers (i.e., various temperature and precipitation variables) by focusing on the response of communities to climatic conditions of the previous year. Taxa and ecoregions differed substantially in their response to climate change conditions. We did not observe any trend of changes in total taxonomic richness or overall abundance over time or with increasing temperatures, which reflects a compensatory turnover in the composition of communities; sensitive Plecoptera decreased in response to warmer years and Ephemeroptera increased in northern regions. Invasive species increased with an increasing number of extreme days which also caused an apparent upstream community movement. The observed changes in functional feeding group diversity indicate that climate change may be associated with changes in trophic interactions within aquatic food webs. These findings highlight the vulnerability of riverine ecosystems to climate change and emphasize the need to further explore the interactive effects of climate change variables with other local stressors to develop appropriate conservation measures.