Filters and templates: stonefly (Plecoptera) richness in Ouachita Mountains streams, U.S.A.

1.  We collected adult stoneflies periodically over a 1-year period at 38 sites in two headwater catchments in the Ouachita Mountains, Arkansas, U.S.A. The 43 species collected were a subset of the Ozark-Ouachita fauna and the much larger fauna of the eastern U.S.A. We estimated 78–91% species coverage in the two catchments using jackknife extrapolation of species richness from our survey.
2.  Many streams, especially small ones, lacked surface water for months, but others, both small and large, flowed permanently.
3.  Using published regional presence–absence and coarse ecological data in a discriminant function analysis (DFA), we identified stream size (negative) and regional frequency of occurrence (positive) as predictors of presence in these headwater catchments. For the combined catchments, the extrapolated richness (51 spp.) was similar to an estimate (48 spp.) based on predicted absences from DFA and the Ouachita provincial total of known stonefly species (57 spp.).
4.  Local species richness (1–27 spp. per site) was correlated strongly with stream size (catchment area) but was independent of stream drying. Generic richness was correlated negatively with stream drying and positively, but less strongly, with stream size.
5.  Regionally endemic stoneflies dominated in drying streams, and widely distributed species dominated in more permanent streams. The composition of stonefly assemblages was associated with regional factors (species pools, regional abundance, evolution of tolerant endemic species, regional climate) and local factors (drying, stream size).     

Patterns of species richness for blackflies (Diptera: Simuliidae) in the Nearctic and Neotropical regions

Abstract.  1. Patterns of simuliid species richness were examined over a variety of scales at 532 stream sites in the Nearctic (394) and Neotropical (138) regions. In Nearctic streams, species richness of immature blackflies both within and across ecoregions and over two seasons was examined. Stream variables at each site included seston, width, depth, velocity, discharge, conductivity, pH, dissolved oxygen, water temperature, dominant streambed-particle size, canopy cover, and riparian vegetation. These variables were subjected to a principal component analysis and derived principal components were related back to richness, using regression analysis. At the level of the stream reach, richness was not highly correlated with single-point measurements of stream conditions.
2. Using data from both Nearctic and Neotropical sites, the effect of regional richness on local richness was examined. As regional richness increased, local diversity reached an asymptote in which further increases in regional richness were not matched by increases in local richness. Hence, simuliid communities are best described as saturated (type II) communities, consistent with the current view of lotic communities as non-equilibrium systems.
3. The well-established pattern of greater species richness in tropical regions was not observed in this study. To the contrary, blackfly richness is higher in temperate streams than in tropical streams at both local and regional scales.     

Macroinvertebrate diversity in headwater streams: a review

1. Headwater streams are ubiquitous in the landscape and are important sources of water, sediments and biota for downstream reaches. They are critical sites for organic matter processing and nutrient cycling, and may be vital for maintaining the 'health' of whole river networks.
2. Macroinvertebrates are an important component of biodiversity in stream ecosystems and studies of macroinvertebrate diversity in headwater streams have mostly viewed stream systems as linear reaches rather than as networks, although the latter may be more appropriate to the study of diversity patterns in headwater systems.
3. Studies of macroinvertebrate diversity in headwater streams from around the world illustrated that taxonomic richness is highly variable among continents and regions, and studies addressing longitudinal changes in taxonomic richness of macroinvertebrates generally found highest richness in mid-order streams.
4. When stream systems are viewed as networks at the landscape-scale, α-diversity may be low in individual headwater streams but high β-diversity among headwater streams within catchments and among catchments may generate high γ-diversity.
5. Differing ability and opportunity for dispersal of macroinvertebrates, great physical habitat heterogeneity in headwater streams, and a wide range in local environmental conditions may all contribute to high β-diversity among headwater streams both within and among catchments.
6. Moving beyond linear conceptual models of stream ecosystems to consider the role that spatial structure of river networks might play in determining diversity patterns at the landscape scale is a promising avenue for future research.     

Comparing species richness among assemblages using sample units: why not use extrapolation methods to standardize different sample sizes?

Comparisons of species richness among assemblages using different sample sizes may produce erroneous conclusions due to the strong positive relationship between richness and sample size. A current way of handling the problem is to standardize sample sizes to the size of the smallest sample in the study. A major criticism about this approach is the loss of information contained in the larger samples. A potential way of solving the problem is to apply extrapolation techniques to smaller samples, and produce an estimated species richness expected to occur if sample size were increased to the same size of the largest sample. We evaluated the reliability of 11 potential extrapolation methods over a range of different data sets and magnitudes of extrapolation. The basic approach adopted in the evaluation process was a comparison between the observed richness in a sample and the estimated richness produced by estimators using a sub-sample of the same sample. The Log-Series estimator was the most robust for the range of data sets and sub-sample sizes used, followed closely by Negative Binomial, SO-J1, Logarithmic, Stout and Vandermeer, and Weibull estimators. When applied to a set of independently replicated samples from a species-rich assemblage, 95% confidence intervals of estimates produced by the six best evaluated methods were comparable to those of observed richness in the samples. Performance of estimators tended to be better for species-rich data sets rather than for those which contained few species. Good estimates were found when extrapolating up to 1.8-2.0 times the size of the sample. We suggest that the use of the best evaluated methods within the range of indicated conditions provides a safe solution to the problem of losing information when standardizing different sample sizes to the size of the smallest sample.     

Species richness and spatial distribution of blackflies (Diptera: Simuliidae) in streams of Central Amazonia, Brazil

1. The spatial distribution and species richness of blackflies were evaluated at 58 stream sites in Central Amazonia, Brazil. Samples were taken along a north–south axis of approximately 130 km and a east–west axis of approximately 220 km.
2. Based on stream-site characteristics, the occurrence of larvae of the six most frequently collected species was highly predictable (79.3–91.5% accuracy in prediction of occurrence). The predictive value of stream size and the presence of impoundments agrees with results of similar work in the Holarctic Region, suggesting a general responses of blackflies to environmental parameters.
3. Although only 19.0% of interstream variation in species richness was explained by a regression model, results suggested that species richness was greater in larger, cooler, faster, covered streams with rocky beds than in smaller, warmer, slower, open streams with sandy bottoms. Overall, the species richness of blackflies (11 species in total) was lower than in the temperate zone suggesting, for some taxa at least, that aquatic communities do not follow the terrestrial pattern of greater species richness in the tropics.     

Abundance, diversity and body size: patterns from a grassland arthropod community

1. The empirical relationships among body size, species richness and number of individuals may give insight into the factors controlling species diversity and the relative abundances of species. To determine these relationships, we sampled the arthropods of grasslands and savannahs at Cedar Creek, MN using sweep nets (90 525 individuals of 1225 species) and pitfall traps (12 721 individuals of 92 species). Specimens were identified, enumerated and measured to determine body size.
2. Both overall and within abundant taxonomic orders, species richness and numbers of individuals peaked at body sizes intermediate for each group. Evolution could create unimodal diversity patterns by random diversification around an ancestral body size or from size-dependent fitness differences. Local processes such as competition or predation could also create unimodal diversity distributions.
3. The average body size of a species depended significantly on its taxonomic order, but on contemporary trophic role only within the context of taxonomic order.
4. Species richness ( S _i) within size classes was related to the number of individuals ( I _i) as S _i =  I i^0·5. This relationship held across a 100 000-fold range of body sizes. Within size classes, abundance distributions of size classes were all similar power functions. A general rule of resource division, together with similar minimum population sizes, is sufficient to generate the relationship between species richness and number of individuals.
5. Smaller bodied species had slightly shallower abundance distributions and may, in general, persist at lower densities than larger species.
6. Our results suggest there may be fewer undescribed small arthropod species than previously thought and that most undescribed species will be smaller than arthropods.     

Diatoms as indicators of river quality in the Nepalese Middle Hills with consideration of the effects of habitat-specific sampling

1. Using a replicated survey design at the catchment scale, we compared the composition and diversity of diatom communities in three stream groups in the Middle Hills of Nepal: semi-natural reference (Arun Valley), enriched by agricultural runoff (Likhu Khola) and grossly polluted by sewage (Kathmandu Valley). We sampled riffles at all sites; in the Kathmandu Valley and Likhu Khola we also sampled diatoms in pools and on vegetation.
2. Species richness and diversity H' were significantly higher in agricultural streams than in either organically polluted streams or references. Community composition, as shown by principal components analysis, varied significantly between all three stream types due to differences in the abundances of species characteristic of organic enrichment (Kathmandu Valley), moderate enrichment (Likhu Khola) or undisturbed hillstreams (Arun Valley).
3. Few species varied in relative abundance between pools, riffles and vegetation so that variations in community composition and diversity were stronger between stream groups than between habitats. Samples from any one habitat produced only 75.7% (± 8.6 SD) of the species from three habitats, so that surveys aimed at recording biodiversity may need more comprehensive habitat coverage than surveys for biomonitoring.
4. We conclude that diatom communities can indicate different sources of pollution in Nepalese streams, and advocate further studies to develop this indicator potential over a wider area of the Himalaya. Comparisons between replicate groups of streams selected a priori helped to clarify effects which were sometimes obscured by other survey designs.     

Response of aquatic hyphomycete communities to enhanced stream retention in areas impacted by commercial forestry

1. Aquatic hyphomycetes are an important component of detritus processing in streams. Their response to enhanced stream retentiveness was tested by manipulating three streams located in Kielder Forest (northern England), a large plantation of exotic conifers, and two streams in Montagne Noire (south-west France) dominated by native broadleaf woodland. Treatment was by placement of logs or plastic litter traps into a 10–20 m stream section. Fungal spores were collected from stream water upstream and downstream of the treated sections over 1–2 years.
2. The average concentration of fungal spores in reference sections was nearly 10× greater in the French streams than in the English streams. The number of hyphomycete species was also higher in the French streams. These differences between regions were probably a consequence of the much lower standing stock and diversity of leaf litter in the English streams.
3. Despite these large regional differences, the treatment had a clear effect in all streams. Detrital standing stocks were enhanced in treated sections by up to 90% in French streams and 70% in English streams.
4. Mean spore density below treated sections increased by 1.8–14.8% in French streams and 10.2–28.9% in the naturally less retentive English streams. The number of fungal species increased significantly below the treated sections of the English streams, although not the French ones.
5. In biologically impoverished plantation streams, input of woody debris can increase detritus retention and enhance hyphomycete diversity and productivity. This may have consequent benefits for detritus processing and macroinvertebrate production.     

Physico-chemistry and aquatic insects of a glacier-fed and a spring-fed alpine stream

1. Physico-chemical conditions and benthic macroinvertebrates were studied in two adjacent alpine streams in the Tyrolean Alps, Austria, for 2 years, and aquatic insect emergence was recorded for 1 year.
2. In the spring-fed system, maximum discharge and increased concentrations of suspended solids, nitrate and particulate phosphorus occurred during snowmelt in June. In the glacier-fed stream, high discharge and strong diel fluctuations in flow and concentrations of suspended solids created a harsh and unstable environment during summer. Glacial ablation, variation in groundwater inflow, and water inputs from tributaries draining calcareous rocks caused water chemistry to vary both seasonally and longitudinally in glacier-fed Rotmoosache.
3. A total of 126 aquatic or semi-aquatic invertebrate taxa were collected, 94 of which were found in the glacier-fed stream and 120 in the spring-fed stream. Chironomid abundance was 2–8 times and taxa richness 2–3 times lower in the glacier-fed stream than in the spring-fed stream, as was the number of chironomid taxa (72 versus 93 total).
4. These results broadly support the conceptual model by Milner & Petts (1994) concerning glacier-fed stream systems. However, single samples and seasonal means showed relatively high invertebrate abundance and richness, especially during winter, indicating a considerable degree of spatial and temporal variability.
5. We suggest that the seasonal shifts from harsh environmental conditions in summer to less severe conditions in autumn and a rather constant environment in winter are an important factor affecting larval development, life-history patterns and the maintenance of relatively high levels of diversity and productivity in glacier-fed streams.     

Vegetation and flow regime in lowland streams

1. The hydrological regime is important to the distribution of benthic organisms in streams. The objective of this study was to identify relationships between hydrological variables, describing the flow regime, and macrophyte cover, species richness, diversity and community composition in Danish lowland streams.
2. We quantified macrophyte vegetation in 44 Danish streams during summer by cover, species richness and diversity. Flow regime was characterized by 18 non-intercorrelated variables describing magnitude, frequency and duration of low and high flow events, timing or predictability of flow and general flow variability.
3. We found support in the stepwise multiple regressions analysis for our expectation that macrophyte cover is lowest in streams with high flow variability and highest in streams with long duration of low flow and low flow variability. We found support for the intermediate disturbance hypothesis as there were significant quadratic relationships between species richness and diversity as functions of disturbance frequency. There was poor discrimination in a detrended correspondence analysis (DCA) analysis of macrophyte community composition between four twinspan groups separating streams with different hydrological properties. Moreover, we did not find any relationship between the presence of disturbance-tolerant species and hydrological disturbance, suggesting that plant community composition developed independently of stream hydrology.     

Effects of snow cover on the benthic fauna in a glacier-fed stream

1. Alpine streams above the tree line are covered by snow for 6–9 months a year. However, winter dynamics in these streams are poorly known. The annual patterns of macroinvertebrate assemblages were studied in a glacial stream in the Austrian Alps, providing information on conditions under the snow.
2. Snow cover influenced water temperature, the content of benthic organic matter and insect development. Taxa richness and abundance of macroinvertebrates did not show a pronounced seasonal pattern. The duration of the autumn period with stable stream beds was important in determining the abundance and composition of the winter fauna.
3. There were significant differences in species composition between summer and winter. Two potential strategies in larval survival were evident: adaptation to the extreme abiotic conditions in summer (e.g. Diamesa spp.) or avoidance of these conditions and development during winter (e.g. Ephemeroptera and Plecoptera).
4. A comparison of a stream reach with continuous snow cover and a stream reach that remained open throughout winter showed that conditions under snow are suboptimal. At the open stream site, with higher water temperatures and greater food supply (benthic organic matter content), abundance and taxa richness was higher and larval growth was faster. Several taxa were found exclusively at this site.
5. Winter conditions did not provide an entirely homogeneous environment, abiotic conditions changed rapidly, especially at the onset of snowfall and at snowmelt. Continuous monitoring is necessary to recognize spatial and temporal heterogeneity in winter environments and the fauna of alpine streams.     

Diversity, distribution and larval habitats of North Swedish blackflies (Diptera: Simuliidae)

1. Blackfly species richness and community structure were analysed at fifty-six sites in northern Sweden in two seasons. The sites were situated in a wide range of streams and rivers from small springbrooks, bog streams and lake-outlet streams to medium-sized forest rivers and large rivers draining montane regions.
2. Thirty-nine blackfly species were found, with between two and thirteen species per site. Neither species richness nor abundance could be related to the environmental variables measured.
3. An analysis of labral fan size of blackflies indicated a clear trend for the prevalence of larvae with small fans in large rivers and larvae with larger fan size in small streams. Similarly, fan size related to current velocities so that large fans were associated with slow current velocities and small fans with high velocities.
4. A strong relationship existed between species composition and habitat, as seen in ordination by non-metric multidimensional scaling. The relationship found between fan size and habitat size-related variables, such as channel width, depth, velocity and substratum particle size, along with longitude and altitude, in partial least squares regression analysis offered an explanation of the species composition–habitat relationship.
5. In addition to testing that distributions of blackfly larvae reflect morphological traits, we tested two general hypotheses pertaining to distribution patterns: (a) that blackfly communities show bimodal distributions; and (b) that their distributions are nested. Neither of these two hypotheses was supported by our observations. However, widespread blackfly species were locally more abundant than those found at relatively few sites, thus showing a positive abundance–occupancy relationship.     

Leaf litter breakdown rates in boreal streams: does shredder species richness matter?

1. Leaf litter breakdown rates were assessed in 23 boreal streams of varying size (first–seventh order) in central and northern Sweden. 2. Shredders were most abundant in small streams, while shredder species richness showed a hump-shaped relationship with stream order, with most species in fourth order streams. 3. In a partial least-squares regression analysis, year, water temperature, shredder species richness and shredder abundance were those factors correlating most strongly with leaf breakdown rates. Shredder species richness was more strongly correlated with leaf litter breakdown rates than shredder abundance, and shredder biomass showed no such correlation. 4. These data suggest that shredder species richness is an important variable in terms of leaf litter dynamics in streams.     

Relationships Between Plant Assemblages and Water Flow Across a Boreal Forest Landscape: A Comparison of Liverworts,Mosses, and Vascular Plants

The distribution of water across landscapes affects the diversity and composition of ecological communities, as demonstrated by studies on variation in vascular plant communities along river networks and in relation to groundwater. However, non-vascular plants have been neglected in this regard. Bryophytes are dominant components of boreal flora, performing many ecosystem functions and affecting ecosystem processes, but how their diversity and species composition vary across catchments is poorly known. We asked how terrestrial assemblages of mosses and liverworts respond to variation in (i) catchment size, going from upland-forest to riparian settings along increasingly large streams and (ii) groundwater discharge conditions. We compared the patterns found for liverworts and mosses to vascular plants in the same set of study plots. Species richness of vascular plants and mosses increased with catchment size, whereas liverworts peaked along streams of intermediate size. All three taxonomic groups responded to groundwater discharge in riparian zones by maintaining high species richness further from the stream channel. Groundwater discharge thus provided riparian-like habitat further away from the streams and also in upland-forest sites compared to the non-discharge counterparts. In addition, soil chemistry (C:N ratio, pH) and light availability were important predictors of vascular plant species richness. Mosses and liverworts responded to the availability of specific substrates (stones and topographic hollows), but were also affected by soil C:N. Overall, assemblages of mosses and vascular plants exhibited many similarities in how they responded to hydrological gradients, whereas the patterns of liverworts differed from the other two groups.     

Diatoms as indicators of river quality in the Nepalese Middle Hills with consideration of the effects of habitat-specific sampling

1. Using a replicated survey design at the catchment scale, we compared the composition and diversity of diatom communities in three stream groups in the Middle Hills of Nepal: semi-natural reference (Arun Valley), enriched by agricultural runoff (Likhu Khola) and grossly polluted by sewage (Kathmandu Valley). We sampled riffles at all sites; in the Kathmandu Valley and Likhu Khola we also sampled diatoms in pools and on vegetation.
2. Species richness and diversity H' were significantly higher in agricultural streams than in either organically polluted streams or references. Community composition, as shown by principal components analysis, varied significantly between all three stream types due to differences in the abundances of species characteristic of organic enrichment (Kathmandu Valley), moderate enrichment (Likhu Khola) or undisturbed hillstreams (Arun Valley).
3. Few species varied in relative abundance between pools, riffles and vegetation so that variations in community composition and diversity were stronger between stream groups than between habitats. Samples from any one habitat produced only 75.7% (± 8.6 SD) of the species from three habitats, so that surveys aimed at recording biodiversity may need more comprehensive habitat coverage than surveys for biomonitoring.
4. We conclude that diatom communities can indicate different sources of pollution in Nepalese streams, and advocate further studies to develop this indicator potential over a wider area of the Himalaya. Comparisons between replicate groups of streams selected a priori helped to clarify effects which were sometimes obscured by other survey designs.     

Temporal variability of stream bioassessments using benthic macroinvertebrates

1. When using benthic macroinvertebrate communities for bioassessment, temporal variation may influence judgement as to whether or not a site is degraded.
2. In a survey of sixteen reference and sixteen test sites in the upper Thames River catchment area (UTRCA) in south-western Ontario, Canada, consistent differences between summer and winter samples were found for taxon richness (increase; P = 0.06) and the Family Biotic Index (decrease; P = 0.11). A bioassessment based on these results would indicate better water quality in the same streams in winter relative to summer. No consistent pattern of seasonal difference was detected for Simpson's Diversity and Equitability, or percentage Dominant Taxon.
3. The Reference Condition Approach to bioassessment uses predictive modelling to explain variation in reference communities with the environmental conditions at these sites as predictors. The community at a test site is compared with that predicted by the model. Several predictive models were constructed using simple geographic and habitat characteristics (i.e. catchment area, distance to source, stream width, substrate and habitat diversity) as predictors. By including season of sampling in the models, we increased their predictive power and the ability of the bioassessment to detect degradation. The best results were achieved when separate predictive models were built for each sampling season.     

Disruption of a longitudinal pattern in environmental factors and benthic fauna by a glacial tributary

SUMMARY 1. In the upper Rhône catchment (Swiss Alps), modifications in the longitudinal pattern of environmental conditions and the benthic macroinvertebrate fauna were investigated in a glacier-fed stream (Rhône) at its confluence with a smaller glacier-fed tributary (Mutt) in June, August and September 1998. The distance to the source glacier was greater for the Mutt than for the Rhône.
2. Environmental conditions were harsher for the biota in the main stream upstream of the confluence than in the tributary. The tributary upstream of the confluence was characterised by higher taxonomic richness and abundance of the zoobenthos than the Rhône upstream.
3. Although environmental conditions in the main stream were little modified by the tributary, the fauna was richer and more diverse below the confluence. During the period of ice melt, colonisation from the Mutt led to the occurrence of faunal elements atypical of glacial streams in the main glacial stream upstream of the confluence, where water temperature remains below 4 °C.
4. Although contributing an average of only 10% to the Rhône discharge, the Mutt tributary is suggested to be the faunal driver of the system.     

Longitude variability of macrobenthos in the streams of Central European Russia

The changes in macrobenthos community depended on changes in stream discharge (from 0.001 m3/sec to 100 m3/sec) were studied on the basis of data from 254 stations of 180 streams located in the centre of European part of Russia. While discharge of stream increases community composition changes considerably. Species diversity increases from 12 to 47 species per station with the increase in stream size. Moreover, the diversity of almost all groups, discerned according habitat or trophic position and main taxonomic groups (with the exception of Plecoptera and Diptera) increases with the increase in stream discharge. The diversity of lithorheophilic species and species inhabiting soft substrates is higher in small streams, whereas phitophilic species are more diverse in large streams. With the increase in stream size the percent of algophagous and filter feeders increases, percent of carnivorous remains the same and the percent of detritophages decreases. The relative role of invertebrates macrotaxons change with the stream size: in the smallest streams the diversity is the highest among Diptera, Trichoptera and Plecoptera, in medium-size streams--among Trichoptera, Ephemeroptera and Diptera, in large streams--in Ephemeroptera, Gastropoda and Trichoptera. In some groups some species replace other (close-related) in parallel with change in stream discharge however in general this tendency is weak. The index of saprobic conditions increases with the increase in stream size from 1.6-1.7 to 2.3-2.4. In small streams the saprobic conditions are very different whereas large streams have tendency to be at beta-mesosaprobic level. Cluster analysis reveal four size classes of streams. Peculiarities of habitat and macrobenthos fauna of these classes are described.     

Colonization by Macroinvertebrates of Experimentally Disturbed Stones in Three Tropical Streams Differing in Size

We experimentally disturbed stones in three streams of different sizes and followed the macroinvertebrate colonization process in terms of abundance, species richness and similarity over 64 days. We hypothesized that colonization in the smallest and in the largest streams would be slower than in the medium‐sized stream. The small upstream pool of colonists available in the smallest stream could restrict colonization, while in the largest stream predation by the diverse fish assemblage could restrict drifting colonists. The medium‐sized stream did not have these two constraints. We found similar colonization patterns in all three streams, leading to the rejection of the stated hypotheses. Lack of support of the original hypothesis might be due to the weakness of the two hypothesized restrictions on colonization. In addition, colonization by crawling species from undisturbed nearby patches might be of significant importance. (© 2004 WILEY‐VCH Verlag GmbH & Co. KGaA, Weinheim)