Macroinvertebrate communities of non-glacial high altitude intermittent streams

1. Macroinvertebrate assemblages of five non‐glacial intermittent high altitude headwater streams (above 1400 m – Serra da Estrela, Portugal), with dry periods of different lengths (0–3 months), were investigated in nearly undisturbed conditions to (i) examine spatial differences and identify environmental variables responsible for the observed invertebrate patterns, (ii) assess the association of dry period length with invertebrate community structure and (iii) determine the influence of using different taxonomic identification levels (order, family and genus) to assess invertebrate community patterns. 2. More than 100 macroinvertebrate genera were identified. Insects clearly dominated these communities with more than 95% of total captures and around 95% of the total richness. Diptera were the most rich and abundant group with chironomid occurrences comprising over 70% of macroinvertebrate captures. 3. The highest taxon richness, diversity, EPT (Ephemeroptera + Plecoptera + Trichoptera) and OCH (Odonata + Coleoptera + Heteroptera) genus richness, the greatest number of exclusive and characteristic taxa identified by the Indicator Value (IndVal), and a distinct community structure shown by Canonical Correspondence Analyses (CCA), were found in the only stream that was never totally dry, with pools lasting over summer. Environmental gradients that spatially structured the macroinvertebrate communities were always related to flow variations. 4. Over time, the highest abundances found in these systems were also related to flow variations and maximum genus richness occurred in the connected pools or in isolated pools. Streams with longer dry periods presented a distinct recolonization phase, with higher abundance of the stonefly larvae Nemoura sp. and the presence of the chironomid larvae Krenosmittia sp., possibly arriving from the hyporheos. 5. Taxonomic level of invertebrate identification was vital for recognizing the characteristic taxa (IndVal) of streams yet was not critical for identifying streams with the highest macroinvertebrate richness/diversity or structuring environmental gradients. 6. Overall, this study emphasizes the variability of high altitude intermittent streams macroinvertebrate communities, despite spatial proximity. This variability was probably related to flow intermittency and hydrologic permanence, different vegetation covers and riverbed substrata. Consequently, the establishment of reference conditions should involve long‐term data collections and more detailed physical characterization. Also, these findings have significant implications for accurately predicting the ecological consequences of future climate change in high altitude scenarios.     

Functional redundancy of stream macroconsumers despite differences in catchment land use

1. The conversion of forested landscapes to agriculture and, increasingly, to suburban and urban development significantly affects the structure and function of both terrestrial and aquatic ecosystems. While a growing body of research is examining how biotic communities change in response to human alteration of landscapes, less is known about how these changes in community structure affect biotic interactions. 2. The objective of this study was to examine top‐down control by macroconsumers (fish and crayfish) across a human‐impacted landscape. We predicted that changes in stream macroconsumers and physicochemical characteristics associated with increased catchment development (e.g. decreased abundance of fish that are obligate benthic invertivores, increased sedimentation) would diminish top‐down control of benthic insects. We expected that effects on algal assemblages would be more variable, with increased top‐down control at sites dominated by algivorous fish and diminished control elsewhere. To test these predictions, we experimentally excluded fish and crayfish from areas of the bed of five streams whose catchments ranged from 100% to <50% forested, and examined the effects of exclusion on benthic insects and algae. 3. Despite cross‐site differences in physical, chemical and biological characteristics, the outcome of our experiments was consistent across five sites representing a range of catchment development. Across all sites, macroconsumers reduced total insect biomass, largely due to decreases in Chironomidae and Hydropsychidae larvae. Macroconsumers also affected algal assemblages, reducing chlorophyll‐a and the proportion of upright and filamentous diatoms (e.g. Melosira, Cymbella) but increasing the proportion of adnate diatoms (e.g. Achnanthes) across all sites. 4. We expected that differences in factors such as macroconsumer assemblage composition, nutrient and light availability and sedimentation would result in variable responses to macroconsumer exclusion in the five streams. Contrary to these expectations, only one response variable (ash‐free dry mass) showed a statistically significant interaction (i.e. site × exclusion) effect. Most responses to exclusion were relatively consistent, suggesting functional redundancy in assemblages of macroconsumers among the sites despite differences in catchment land use.     

Biological traits of stream macroinvertebrates from a semi-arid catchment: patterns along complex environmental gradients

1. The relationships between biological traits of macroinvertebrates and environmental characteristics were investigated in streams with contrasting physical, chemical or landscape level attributes. We used an ordination technique, RLQ analysis, which links an environmental table (R) with traits table (Q) through an abundance table (L) to investigate the relationship between habitat characteristics and biological traits.
2. A major environmental axis explaining the distribution of species and their distinctive biological features was obtained. This axis included variables of anthropogenic pressure (agricultural and urban uses) and natural variability (climatic and geologic) that are strongly intercorrelated in the study area, with a clear spatial component.
3. The attributes of species from frequently disturbed systems (small size, multivoltinism, diapause, ovoviviparity, etc.) were associated with semi-arid areas whereas traits common in more stable and favourable environments (large body size, semi-voltinism, isolated eggs, etc.) were found in upland forested areas.
4. The natural climatic variation was proposed as a disturbance axis of a theoretical habitat templet (driven by the intense hydrological disturbances typical of semi-arid streams), while anthropogenic pressure (mainly intensive agriculture) and high salinity, a natural consequence of geology, was proposed as an adversity axis. Different life-histories associated with contrasting environmental features were superimposed in this habitat templet.
5. The ecological–evolutionary scenario in which stream macroinvertebrates have evolved and by which their communities are organized, is closely linked to disturbance, environmental harshness and human pressure.     

Synthesis of a 13C-Labeled Tracer for Stream DOC: Labeling Tulip Poplar Carbon with 13CO2

Ecosystem tracer-level additions would benefit from a stable isotope-labeled source of complex organic molecules. We tested a method to label tree C with ¹³C and create a stable isotope tracer for stream dissolved organic carbon (DOC) using tulip poplar (Liriodendron tulipifera L.) seedlings. In 2000, seedlings were grown with 0.82 moles of ¹³CO₂ to assess the distribution and level of ¹³C enrichment in the tree tissues. In 2001, seedlings were grown with 25 times more ¹³CO₂ to generate tissues with a ¹³C signal strong enough for a ¹³C-DOC stream tracer addition. ¹³C enrichment in the trees varied in each year and by tissue age and type. Tissues formed during labeling (new) were more enriched in ¹³C than tissues established prior to the ¹³CO₂ injection (old). Stems were most enriched in ¹³C in both new and old tissues. A higher percentage of ¹³CO₂ was incorporated into seedlings in 2000 (59% ±1) than 2001 (43% ±0). Percent ¹³C incorporation among tree tissue types paralleled biomass distributions. Although tree C and ¹³C were equally soluble in both years, a greater percentage of tree C went into solution in 2001 (30%) than 2000 (20%). The water-soluble tree C accounted for approximately 12% of the injected ¹³CO₂ and had both humic and polysaccharide components. Results from a whole-stream ¹³C-DOC tracer addition demonstrated that tree C could be sufficiently labeled with ¹³CO₂ to create a stream DOC isotope tracer with some polymeric constituents.     

Habitat characteristics and macroinvertebrate assemblages in boreal forest streams: relations to catchment silvicultural activities

We compared the stream habitat characteristics and macroinvertebrate assemblages of boreal headwater streams in both the Finnish and the Russian parts of a single river basin, the Koitajoki River. Over the last 50 years, the Finnish side of the catchment has been managed using modern forestry techniques, whereas Russian side has remained nearly unexploited and is near to its natural state. Differences in silvicultural activities were observed to contribute to differences in habitat structure. The channel habitats were in fairly natural state in the Russian reference streams, whereas the impacted Finnish sites were cleared and straightened. In comparison with the impacted channels, the abundance of coarse woody debris (CWD) was 10–100-fold higher in the reference streams. Implications on the forestry-induced deterioration of water quality were also observed. On the contrary, only small differences in macroinvertebrate assemblages were detected. Despite the lower amount of retentive structures (CWD), significantly higher relative abundance of shredders was observed in the forestry-impacted streams. Otherwise the zoobenthic communities were quite similar in the two subcatchments. We suggest that several mechanisms may explain this similarity: (1) community structure is controlled by naturally acidic conditions, (2) the adverse impacts of forestry on habitat structure and water quality of streams may be compensated by increased input of deciduous litter and organic compounds from drained, structurally young riparian forests and (3) macroinvertebrate species have flexible feeding habits and may thus readily adapt to changing conditions.     

Variability in stream macroinvertebrates at multiple spatial scales

1. We intensively sampled 16 western Oregon streams to characterize: (1) the variability in macroinvertebrate assemblages at seven spatial scales; and (2) the change in taxon richness with increasing sampling effort. An analysis of variance (ANOVA) model calculated spatial variance components for taxon richness, total density, percent individuals of Ephemeroptera, Plecoptera and Trichoptera (EPT), percent dominance and Shannon diversity.
2. At the landscape level, ecoregion and among‐streams components dominated variance for most metrics, accounting for 43–72% of total variance. However, ecoregion accounted for very little variance in total density and 36% of the variance was attributable to differences between streams. For other metrics, variance components were more evenly divided between stream and ecoregion effects.
3. Within streams, approximately 70% of variance was associated with unstructured local spatial variation and not associated with habitat type or transect position. The remaining variance was typically split about evenly between habitat and transect. Sample position within a transect (left, centre or right) accounted for virtually none of the variance for any metric.
4. New taxa per stream increased rapidly with sampling effort with the first four to eight Surber samples (500–1000 individuals counted), then increased more gradually. After counting more than 50 samples, new taxa continued to be added in stream reaches that were 80 times as long as their mean wetted width. Thus taxon richness was highly dependent on sampling effort, and comparisons between sites or streams must be normalized for sampling effort.
5. Characterization of spatial variance structure is fundamental to designing sampling programmes where spatial comparisons range from local to regional scales. Differences in metric responses across spatial scales demonstrate the importance of designing sampling strategies and analyses capable of discerning differences at the scale of interest.     

Assessing the performance of volunteers in monitoring streams

1. Citizens are concerned about the quality of water resources and many participate in monitoring activities, though doubts remain about the quality of the data volunteers collect. We trained volunteers to collect benthic macroinvertebrates using professional protocols. Of the seven stream sites sampled by volunteer crews, six sites were also sampled by professional crews.
2. In the laboratory, volunteers used morphological features to identify as many different taxa as possible within the major insect orders; their identification was approximately to family. Volunteers calculated five metrics: total taxon richness, richness of three key groups (Ephemeroptera, Plecoptera and Trichoptera), and percentage dominance of the three most abundant taxa. All metrics were strongly correlated with (a) the percentage of urbanized area in the catchment and (b) the metrics derived from a more complete taxonomic identification by a professional scientist. Taxon richness metrics declined with urban development, while percent dominance increased.
3. An overall summary multimetric index was used to compare the field and laboratory procedures of volunteers and professionals. Using an ANOVA model, we detected no significant difference between field samples collected by volunteers and professionals. The variance of index values associated with differences between crews was zero. The ability of the index to detect significant differences among sites (statistical power) improved by only 13% for assessments based on professional laboratory identification instead of volunteer laboratory identification.
4. Citizen volunteers, when properly trained, can collect reliable data and make stream assessments that are comparable to those made by professionals. Data collected by volunteers can supplement information used by government agencies to manage and protect rivers and streams.     

Diurnal variations in the carbon chemistry of two acidic peatland streams in north-east Scotland

1. Two acidic peatland upland streams in north‐east Scotland draining catchments of 1.3 and 41.4 km² were sampled each season for 2 years to investigate diurnal variations in dissolved and gaseous forms of carbon. Stream metabolism, alkalinity, discharge, pH, air and water temperatures were measured to aid data interpretation. 2. Free CO₂ showed marked diurnal variation with lowest concentrations during the period from late morning to early afternoon and highest during the hours of darkness. Although alkalinity and pH also showed some diurnal fluctuations, in comparison with other more productive alkaline systems, variation was small. Dissolved organic carbon (DOC) showed no significant diurnal pattern. However, significant changes in stream discharge influenced DOC concentrations, as well as over‐riding diurnal patterns of free CO₂, alkalinity and pH. 3. The highest diurnal ratios (maximum concentration/minimum concentration) in CO₂, gross primary productivity (GPP) and community respiration (CR) occurred in spring and summer and the lowest in autumn and winter. Variation in biotic in‐stream processes caused changes in CO₂ concentrations and temperature affected both the solubility of CO₂ and changes in up‐stream CO₂ inputs. There was no significant difference in diurnal fluctuations between the two orders of stream studied. 4. The mean GPP (as CO₂) was 0.81 g CO₂ m^?2 day^?1 and mean CR 2.67 g CO₂ m^?2 day^?1. The mean primary production/respiration (P/R) ratio was 0.26 ± 0.09 and 0.33 ± 0.15 in the first and second order streams, respectively. These values are low compared with published data because these heterotrophic headwater streams are dominated by benthic respiration and upstream allochthonous inputs with little autotrophic metabolism, particularly during the colder autumn and winter months. 5. The results have implications for the calculation of dissolved inorganic carbon (DIC) fluxes in streamwater. Samples taken during daylight hours tend to have lower concentrations of free CO₂ and HCO₃^? than samples taken during darkness. During spring, concentrations of free CO₂ were measured up to 2.4 (annual mean 1.8) times higher at night than during the day at a similar discharge. It is suggested that fluxes based on daytime measurements alone will under‐estimate the annual flux of these determinands in streamwater by as much as 40%.     

Evaluation of methods for estimating macroinvertebrate species richness using individual stones in tropical streams

1. The most straightforward way to assess diversity in a site is the species count. However, a relatively large sample is needed for a reliable result because of the presence of many rare species in rich assemblages. The use of richness estimation methods is suggested by many authors as a solution for this problem in many cases.
2. We examined the performance of 13 methods for estimating richness of stream macroinvertebrates inhabiting riffles both at local (stream) and regional (catchment) scales. The evaluation was based on (1) the smallest sub-sample size needed to estimate total richness in the sample, (2) constancy of this size, (3) lack of erratic behaviour in curve shape and (4) similarity in curve shape through different data sets. Samples were from three single stream sites (local) and three from several streams within the same catchment basin (regional). All collections were made from protected forest areas in south-east Brazil.
3. All estimation methods were dependent on sub-sample size, producing higher estimates when using larger sub-sample sizes. The Stout and Vandermeer method estimated total richness in the samples with the smallest sub-sample size, but showed some erratic behaviour at small sub-sample sizes, and the estimated curves were not similar among the six samples. The Bootstrap method was the best estimator in relation to constancy of sub-sample sizes, but needed an unacceptably large sub-sample to estimate total richness in the samples. The second order Jackknife method was the second best estimator both for minimum sub-sample size and constancy of this size and we suggest its use in future studies of diversity in tropical streams. Despite the inferior performance of several other methods, some produced acceptable results. Comments are made on the utility of using these estimators for predicting species richness in an area and for comparative purposes in diversity studies.     

Herbivory and growth in terrestrial and aquatic populations of amphibious stream plants

1. Many amphibious plant species grow in the transition between terrestrial and submerged vegetation in small lowland streams. We determined biomass development, leaf turnover rate and invertebrate herbivory during summer in terrestrial and aquatic populations of three amphibious species to evaluate advantages and disadvantages of aerial and submerged life.
2. Terrestrial populations had higher area shoot density, biomass and leaf production than aquatic populations, while leaf turnover rate and longevity were the same. Terrestrial populations experienced lower percentage grazing loss of leaf production (average 1.2–5.1%) than aquatic populations (2.9–17.3%), while the same plant dry mass was consumed per unit ground area.
3. Grazing loss increased linearly with leaf age apart from the youngest leaf stages. Grazing loss during the lifetime of leaves was therefore 2.4–3.1 times higher than mean apparent loss to standing leaves of all ages. The results imply that variation in density of grazers relative to plant production can account for differences in grazing impact between terrestrial and aquatic populations, and that fast leaf turnover keeps apparent grazing damage down.
4. We conclude that the ability of amphibious plants to grow submerged permits them to expand their niche and escape intense competition on land, but the stream does not provide a refugium against grazing and constrains plant production compared with the terrestrial habitat.