A technique for assessing the composition and density of the macroinvertebrate fauna of large stones in streams

A device for quantitatively sampling the macroinvertebrates of large stones in streams is described. In comparison to the usual method for sampling the fauna of large stones, (the lifting of stones upstream of a hand net), the present method accurately samples stonedwelling animals that are good swimmers. Details are given of a reliable method to measure the surface area of stream stones using thin plastic sheeting.     

Species richness in tropical fresh waters of Australia

General ecological expectations about the relationship between latitude and species richness are that at low latitudes (the tropics) species richness is greater than at higher latitudes (temperate and polar regions). Recent work suggests that this may not be the case for several habitat types and biological groups in Australia. Results are conflicting: on present evidence (admittedly sparse) it appears that in Australian tropical fresh waters species richness is generally depressed in zooplankton and littoral microfaunal communities, but not in macroinvertebrate communities in typical streams and in fish communities (and perhaps also in amphibian and reptile communities). The situation is indeterminate for tropical phytoplankton and macrophyte communities.     

Tree species richness in primary and old secondary tropical forest in Singapore

If secondary succession can accumulate species rapidly, then tropical secondary forests may have an important role to play in the conservation of biodiversity. Data on the floristic composition of forest stands in the Central Catchment Nature Reserve, Singapore, have been analysed to investigate the diversity of approximately 100-year-old tropical secondary forest. Classification using TWINSPAN indicated that three floristic communities could be recognized from 59 0.2 ha plots enumerated for trees >30 cm gbh. These were two types of secondary forest, both dominated by Rhodamnia cinerea (Myrtaceae), and dryland primary forest. The secondary forest was developed on land abandoned after use for agriculture at the end of the 19th century. The 16 primary forest plots contained a total of 340 species, more than the 281 recorded from the 43 plots of the two secondary forest types combined. The mean species number per plot in the more diverse of the two secondary forests was only about 60% of the primary forest. Thus the secondary forest, despite a century or so for colonization by species and the presence of contiguous primary forest, was still significantly less diverse than primary forest areas. It is concluded that secondary forest cannot be assumed to accrete biodiversity rapidly in the tropics, and may not be of direct value in conservation. However, other indirect roles, such as providing resources for native animals, and buffering and protecting primary forest fragments may make the protection of secondary forest worthwhile.     

Benthic macroinvertebrate distributions in the riffle-pool communities of two east Texas streams

The benthic macroinvertebrate riffle-pool communities of two east Texas streams were sampled monthly for a period of one year. In contrast to previous studies in primarily upland areas, pools in Alazan Creek and Bernaldo Bayou contained significantly higher densities and biomass, as well as a significantly higher diversity and number of taxa. The majority of taxa collected could be characterized as pool adapted organisms, with the Diptera, Ephemeroptera, Coleoptera, and Odonata having greater numbers in pools. Perlesta, Baetis, and Cheumatopsyche were the only genera to have significant numbers in riffles at both sampling sites. A greater number of taxa were restricted to pools at both sampling locations, and although an unstable, sandy substrate was a factor in limiting colonization of riffles, riffles were not as productive as pools even in the presence of suitable substrate conditions. A cluster analysis based on similarity indices indicated that like habitats between streams were more similar to each other, than were adjacent riffle-pool complexes within streams.     

Concordance between ecotypes and macroinvertebrate assemblages in Mediterranean streams

1. According to the guidelines of the European Water Framework Directive, assessment of the ecological quality of streams and rivers should be based on ecotype-specific reference conditions. Here, we assess two approaches for establishing a typology for Mediterranean streams: a top-down approach using environmental variables and bottom-up approach using macroinvertebrate assemblages.
2. Classification of 162 sites using environmental variables resulted in five ecotypes: (i) temporary streams; (ii) evaporite calcareous streams at medium altitude; (iii) siliceous headwater streams at high altitude; (iv) calcareous headwater streams at medium to high altitude and (v) large watercourses.
3. Macroinvertebrate communities of minimally disturbed sites ( n  = 105), grouped using UPGMA (unweighted pair-group method using arithmetic averages) on Bray–Curtis similarities, were used to validate four of the five ecotypes obtained using environmental variables; ecotype 5, large watercourses, was not included as this group had no reference sites.
4. Analysis of similarities ( anosim ) showed that macroinvertebrate assemblage composition differed among three of the four ecotypes, resulting in differences between the bottom-up and top-down classification approaches. Siliceous streams were clearly different from the other three ecotypes, evaporite and calcareous ecotypes did not show large differences in macroinvertebrate assemblages and temporary streams formed a very heterogeneous group because of large variability in salinity and hydrology.
5. This study showed that stream classification schemes based on environmental variables need to be validated using biological variables. Furthermore, our findings indicate that special attention should be given to the classification of temporary streams.     

Spatial variability in macroinvertebrate assemblages along and among neighbouring equatorial glacier‐fed streams

1. During the past two decades, understanding of the structure and function of glacier‐fed stream ecosystems at temperate latitudes has increased substantially. In contrast, information on their tropical counterparts is very limited. We studied three neighbouring glacier‐fed streams in the tropical Andes of Ecuador. Our main goals were (i) to determine overall longitudinal patterns in density, taxon richness and the composition of macroinvertebrate assemblages and driving factors in equatorial glacial streams and (ii) to examine variability among replicate streams in faunal metrics and assemblages, and stream‐specific effects of supposed environmental key factors. 2. We measured four geographical and 17 environmental factors and collected five Surber samples (500 cm²) of macroinvertebrates at each of nine sites, three sites along three streams. The streams were located 1–5 km apart. In each stream, the three sites were placed at comparable distances from the glacier and were grouped as ‘upper’ (50–200 m), ‘middle’ (1.5 km) and ‘lower’ sites (3.5–5.6 km). 3. In total, 2200 individuals (64% chironomids) were collected and 47 taxa (30 dipterans, 18 of these Chironomidae) identified. Density ranged from 176 to 372 ind. m^?2, and the number of taxa ranged from 2 to 6 at the upper sites and 868–3044 ind. m^?2 and 21–27 taxa at the lower sites. Density, number of taxa, rarefied richness and axis‐1 coordinates from a MDS ordination increased logarithmically with distance from the glacier. These faunal metrics were equally related to altitude and glacier per cent of catchment and correlated with maximum conductivity, mean temperature, mean daily maximum temperature and a channel stability index. As expected, the mean difference in distance decay in similarity was higher at the upper (47% km^?1) than at the lower reaches (20% km^?1) of the streams. 4. The number of taxa varied among sites within the upper and middle groups, but not among the lower sites. In contrast, but in accordance with our expectation, assemblage composition did not differ among upper sites but did so at middle and lower sites, following a supposed decrease in environmental harshness along the streams. Relationships between faunal metrics and the four environmental variables mean temperature, the stability index, chlorophyll a and coarse particulate organic matter also varied among the three streams. Generalised linear model analyses revealed that temperature interacted with stream on macroinvertebrate density, while chlorophyll a had a significant effect on the number of taxa in interaction with stream and stability. 5. The basic predictions of the Milner et al. (2001a) , model regarding longitudinal faunal patterns and temperature and stability as main driving factors were met by our three replicate equatorial glacial streams. Qualitative departures from the model were mainly because of zoogeographical differences. We demonstrated that variability in assemblages between comparable sites in closely situated streams was considerable, and the effect of key environmental factors varied among streams and interacted with other factors. Quantifying spatial variation in benthic assemblages may help us foresee possible consequences for biodiversity as a result of glacial retreat.     

Species richness and assemblage structure of Trichoptera in Danish streams

• 1 Species richness and assemblages of Trichoptera from 157 ‘pristine’ Danish lowland stream sites were analyzed, for dependence on geographical position of the sites and simple physical variables, using two way indicator species analysis and detrended correspondence analysis.
• 2 A total of 106 species were recorded, representing ≈ 90% of the species pool known from Danish streams. Only seven species occurred at more than half the sites, whereas an additional 11 species were found at more than a quarter of the sites.
• 3 Although sites showed significant regional differences in environmental variables (stream order, width, slope and presence/absence of riparian forest), species richness and assemblages were primarily correlated with stream order, width and slope. Maximum richness was found at the largest (5th order) stream sites.
• 4 Regional differences in species assemblages were found, with several species being absent from the islands Funen and Bornholm. Species assemblages also differed between forested and non‐forested stream sites.
• 5 We found evidence that stream temperature may be of only minor importance in determining Trichoptera species richness and assemblage composition in Danish streams compared to other size‐related physical factors.

     

Drivers of beta diversity of macroinvertebrate communities in tropical forest streams

1. There has recently been increasing interest in patterns of beta diversity but we still lack a comprehensive understanding of these patterns in various regions (e.g. the tropics), ecosystems (e.g. streams) and organism groups (e.g. invertebrates). 2. Our aim was to investigate the patterns of beta diversity of stream macroinvertebrates in relation to key environmental (i.e. stream size, pH and habitat degradation) and geographical variables (i.e. latitude, longitude, altitude) in a tropical region. We surveyed a total of 8–10 riffle sites in each of 34 streams (altogether 337 riffle sites were sampled) in Peninsular Malaysia to examine variation in macroinvertebrate community composition at within‐stream and among‐stream scales. 3. Based on test of homogeneity of dispersion, we found that the streams studied differed significantly in within‐stream variation in community composition (i.e. among‐site variation of within stream beta diversity). The patterns were similar based on Bray–Curtis coefficient on abundance data, Sorensen coefficient on presence–absence data and Simpson coefficient on presence–absence data. We also found that within‐stream beta diversity was significantly related to stream size, pH and latitude, with each of these variables individually accounting for around 20% of the variation in beta diversity in simple regressions, while the total variation explained by the three significant variables amounted to around 50% in multiple regressions. By contrast, habitat degradation, longitude and altitude were not significantly related to beta diversity. We also found that the factor drainage basin accounted for much of the variation in beta diversity in general linear models, suppressing the effects of environmental variables. 4. We concluded that within‐stream beta diversity is mainly related to a combination of the identity of a drainage basin and stream environmental factors. Our findings provide important background for stream environmental assessment and conservation planning by emphasising that (i) macroinvertebrate communities within streams are not homogeneous, but show considerable beta diversity, (ii) streams differ in their degree of within‐stream beta diversity, (iii) stream size and water pH should be considered in applied contexts related to within‐stream beta diversity and (iv) historical effects may be different in different drainage basins and may affect present‐day patterns of within‐stream beta diversity.     

Influence of macroinvertebrate sample size on bioassessment of streams

In order to standardise biological assessment of surface waters in Europe, a standardised method for sampling, sorting and identification of benthic macroinvertebrates in running waters was developed during the AQEM project. The AQEM method has proved to be relatively time-consuming. Hence, this study explored the consequences of a reduction in sample size on costs and bioassessment results. Macroinvertebrate samples were collected from six different streams: four streams located in the Netherlands and two in Slovakia. In each stream 20 sampling units were collected with a pond net (25×25 cm), over a length of approximately 25 cm per sampling unit, from one or two habitats dominantly present. With the collected data, the effect of increasing sample size on variability and accuracy was examined for six metrics and a multimetric index developed for the assessment of Dutch slow running streams. By collecting samples from separate habitats it was possible to examine whether the coefficient of variation (CV; measure of variability) and the mean relative deviation from the “reference” sample (MRD; measure of accuracy) for different metrics depended only on sample size, or also on the type of habitat sampled. Time spent on sample processing (sorting and identification) was recorded for samples from the Dutch streams to assess the implications of changes in sample size on the costs of sample processing. Accuracy of metric results increased and variability decreased with increasing sample size. Accuracy and variability varied depending on the habitat and the metric, hence sample size should be based on the specific habitats present in a stream and the metric(s) used for bioassessment. The AQEM sampling method prescribes a multihabitat sample of 5 m. Our results suggest that a sample size of less than 5 m is adequate to attain a CV and MRD of ≤ 10% for the metrics ASPT (Average Score per Taxon), Saprobic Index and type Aka+Lit+Psa (%) (the percentage of individuals with a preference for the akal, littoral and psammal). The metrics number of taxa, number of individuals and EPT-taxa (%) required a multihabitat sample size of more than 5 m to attain a CV and MRD of ≤ 10%. For the metrics number of individuals and number of taxa a multihabitat sample size of 5 m is not even adequate to attain a CV and MRD of ≤ 20%. Accuracy of the multimetric index for Dutch slow running streams can be increased from ≤ 20 to ≤ 10% with an increase in labour time of 2 h. Considering this low increase in costs and the possible implications of incorrect assessment results it is recommended to strive for this ≤ 10% accuracy. To achieve an accuracy of ≤ 10% a multihabitat sample of the four habitats studied in the Netherlands would require a sample size of 2.5 m and a labour time of 26 h (excluding identification of Oligochaeta and Diptera) or 38 h (including identification of Oligochaeta and Diptera). Electronic supplementary material Electronic supplementary material is available for this article at and accessible for authorised users.     

Role of transported particulate organic matter in the macroinvertebrate colonization of litter bags in streams

1. The exposure of mesh litter bags has been widely used to investigate the role of benthic macroinvertebrates in leaf processing in freshwaters. In this sense, several studies have related litter bag breakdown rates to the presence of colonizing invertebrates. A possible confounding factor in such experiments is that the litter bags trap suspended particulate organic matter that can itself attract invertebrate colonists, irrespective of the intended experimental treatment.
2. We attempted to quantify the accumulation of particulate organic matter (POM) within litter bags and to investigate its possible impact on macroinvertebrate density and richness. In seven headwater forested streams we exposed mesh bags filled either with beech leaves ( Fagus sylvatica ) or with plastic strips of an equal surface area.
3. Principal component analysis (PCA) showed that bag type and stream were the main explanatory variables for invertebrate colonization and POM accumulation within the bags. In contrast, there was little variation among sampling dates (6.4% of the total inertia).
4. The accumulated POM within the bags was substantial (up to 8.83 g ash-free dry mass (AFDM)) but highly variable among sites (mean from 0.32 to 4.58 g AFDM). At each of the seven sites, both richness and abundance of invertebrates were positively correlated with the mass of accumulated POM in bags. Macroinvertebrate colonization (notably taxon richness) was directly linked with the quantity of POM accumulated.
5. Our findings provide evidence of a potential pitfall in linking invertebrates to litter processing in mesh bags, particularly when large amounts of POM, entrained in stream flow, accumulate within the bags. An evaluation of the POM mass trapped in litter bags could account for the erratic patterns sometimes observed in their colonization by invertebrates.     

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.     

Prediction of neotropical tree and liana species richness from soil and climatic data

We present an analysis of local species richness in neotropical forests, based on a number of 0.1 ha samples of woody plants collected by the late Alwyn Gentry. For each of 69 forests, soils were analysed and climatic data were collated. Using transformed independent variables and interaction terms, multiple regression equations were developed that explained the greatest possible amount of variation in species richness, and the best equations were selected on the basis of regression diagnostics. The best models are presented for (a) all neotropical forests, (b) forests west of the Andes (transandean) and (c) east of the Andes (cisandean), and for various subsets based on elevation and annual rainfall. For the whole dataset, and for most subsets, annual rainfall and rainfall seasonality were the most important variables for explaining species richness. Soil variables were correlated with precipitation — drier forests have more nutrient-rich soils. After the inclusion of rainfall variables, available soil nutrient concentrations contributed little to explaining or accounting for additional variation in species numbers, indicating that tropical forest species richness is surprisingly independent of soil quality. The results are consistent with the hypothesis that plants in mature tropical forests may obtain nutrients through the process of direct cycling, in which mineral nutrients are extracted from litterfall before they enter the soil. The strong relationship between community species richness and rainfall patterns has implications for biodiversity conservation. Wet forests with an ample year-round moisture supply harbour the greatest number of woody plant species and should be a focus of conservation efforts.Died 3 August 1993.     

Community structure of Ephemeroptera in Siberian streams

In recent decades, the relationships between environmental conditions and community structures of stream macroinvertebrates have been investigated in many parts of the world. However, knowledge about assemblages of mayflies and other stream macroinvertebrates in Siberia (northern Asia) and Asia is limited. In fact, the patterns in mayfly species richness and assemblage structure in relation to environmental parameters have not been previously examined in western Siberia. The aim of the present study was to examine the relationship between Ephemeroptera community structure and physical parameters along a river altitude/longitude gradient in Siberia. The results showed that maximum species richness was at relatively low altitudes, high water temperatures, slow current velocities, medium stream widths, medium‐small substrate particle size, and the presence of macrophytes. The mayfly assemblage was separated using TWINSPAN classification into eight distinct groups, which differed significantly with respect to at least one measured environmental factor. Multivariate ordination (detrended correspondence analysis) revealed that mayfly assemblages are structured by a single dominant gradient of altitude‐related environmental variables; altitude and water temperature were the best predictors. Ordination further revealed that mayfly assemblages are structured by altitude‐related environmental factors at high elevations, whereas in the lowlands these factors are less important.     

Monitoring change in aquatic invertebrate biodiversity: sample size, faunal elements and analytical methods

Replication is usually regarded as an integral part of biological sampling, yet the cost of extensive within-wetland replication prohibits its use in broad-scale monitoring of trends in aquatic invertebrate biodiversity. In this paper, we report results of testing an alternative protocol, whereby only two samples are collected from a wetland per monitoring event and then analysed using ordination to detect any changes in invertebrate biodiversity over time. Simulated data suggested ordination of combined data from the two samples would detect 20% species turnover and be a cost-effective method of monitoring changes in biodiversity, whereas power analyses showed about 10 samples were required to detect 20% change in species richness using ANOVA. Errors will be higher if years with extreme climatic events (e.g. drought), which often have dramatic short-term effects on invertebrate communities, are included in analyses. We also suggest that protocols for monitoring aquatic invertebrate biodiversity should include microinvertebrates. Almost half the species collected from the wetlands in this study were microinvertebrates and their biodiversity was poorly predicted by macroinvertebrate data.     

Macroinvertebrate stream communities along regional and physico-chemical gradients in Western Greenland

1. Macroinvertebrates were collected and physico‐chemical variables measured at 16 stream sites in Western Greenland during July 1999. Eight sites were located on Disko Island in an arctic oceanic climate and eight sites in the Kangerlussuaq area close to the icecap where the climate is arctic continental. The streams had different water sources (glacial, groundwater, snowmelt and lake water). 2. The streams showed pronounced differences in water temperature (2.2–17.3 °C), concentrations of suspended solids (0–2400 mg L^?1), and conductivity (10–109 μS cm^?1). Principal component analysis (PCA) analysis of the physico‐chemical variables separated the Disko Island sites into a distinct group, whereas the sites in the Kangerlussuaq area were more dispersed. 3. A total of 56 macroinvertebrate species were found, including 31 species of Chironomidae, the most abundant of which was Orthocladius thienemanni. Diamesa sp. was only the sixth most abundant chironomid taxon. Species composition varied between sites, and abundance varied from about 20 individuals m^?2 in a glacier fed stream to more than 16 000 m^?2 in a lake outlet. 4. The macroinvertebrate communities of the 16 streams were separated into five TWINSPAN groups reflecting water source, irrespective of region. Lake outlets and ground‐water‐fed streams had the highest species richness and abundance, temperature and bed stability, while glacier‐fed streams were characterized by low species richness, abundance, temperature, bed stability and high concentrations of suspended solids. Macroinvertebrate species richness was positively correlated with water temperature and negatively with bed stability. Conductivity was positively correlated with invertebrate abundance. 5. The results of this study suggest that the source of stream water can be used to predict invertebrate community composition in Greenlandic streams and thus the effects of changes in water balance and flow regime, and to identify sites of special conservation interest.     

Benthic macroinvertebrate communities of intermittent streams in the middle reaches of the Guadiana Basin (Portugal)

The Guadiana River has an irregular hydrological regime, with severe droughts and floods, but little is known about how aquatic fauna respond to these natural events. Macroinvertebrate data and environmental information were collected at seven sites from three tributaries in the middle reaches of the Guadiana River, approximately every 3 months from April 1995 to April 1997. Despite considerable annual variation in discharge (related to duration of flood and drought periods), the number of macroinvertebrates found was consistently high. Diptera represented the major proportion of the benthic fauna (73.2%) followed by Ephemeroptera (10.3%), Coleoptera (4.1%) and Trichoptera (3.1%). Canonical correspondence analysis (CCA) was used to evaluate the relationships between taxa density and habitat variables. Generally, Plecoptera and Ephemeroptera were found in the upstream sampling sites. Wider and deeper sites were associated with the presence of Diptera and were least diverse. High values for both the Shannon-Wiener diversity index and the average score per taxon were usually found at upstream sites where Ephemeroptera and Plecoptera predominated. The data suggest that macroinvertebrates have a great capacity to recover rapidly from severe drought periods, both in terms of taxonomic diversity and number of individuals.     

Seasonal dynamics of macroinvertebrate assemblages in the benthos and associated with detritus packs in two low-order streams with different riparian vegetation

• 1 The seasonal dynamics of the benthic macroinvertebrate assemblage, and the subset of this assemblage colonising naturally formed detritus accumulations, was investigated in two streams in south‐west Ireland, one draining a conifer plantation (Streamhill West) and the other with deciduous riparian vegetation (Glenfinish). The streams differed in the quantity, quality and diversity of allochthonous detritus and in hydrochemistry, the conifer stream being more acid at high discharge. We expected the macroinvertebrate assemblage colonising detritus to differ in the two streams, due to differences in the diversity and quantity of detrital inputs.
• 2 Benthic density and taxon richness did not differ between the two streams, but the density of shredders was greater in the conifer stream, where there was a greater mass of benthic detritus. There was a significant positive correlation between shredder density and detritus biomass in both streams over the study period.
• 3 Detritus packs in the deciduous stream were colonised by a greater number of macroinvertebrates and taxa than in the conifer stream, but packs in both streams had a similar abundance of shredders. The relative abundance of taxa colonising detritus packs was almost always significantly different to that found in the source pool of the benthos.
• 4 Correspondence analysis illustrated that there were distinct faunal differences between the two streams overall and seasonally within each stream. Differences between the streams were related to species tolerances to acid episodes in the conifer stream. Canonical correspondence analysis demonstrated a distinct seasonal pattern in the detrital composition of the packs and a corresponding seasonal pattern in the structure of the detritus pack macroinvertebrate assemblage.
• 5 Within‐stream seasonal variation both in benthic and detritus pack assemblages and in detrital inputs was of similar magnitude to the between‐stream variation. The conifer stream received less and poorer quality detritus than the deciduous stream, yet it retained more detritus and had more shredders in the benthos. This apparent contradiction may be explained by the influence of hydrochemistry (during spate events) on the shredder assemblage, by differences in riparian vegetation between the two streams, and possibly by the ability of some taxa to exhibit more generalist feeding habits and thus supplement their diets in the absence of high quality detritus.