Scales of spatiotemporal variability in macroinvertebrate abundance and diversity in monsoonal streams: detecting environmental change

1. We quantified spatial and temporal variability in benthic macroinvertebrate species richness, diversity and abundance in six unpolluted streams in monsoonal Hong Kong at different scales using a nested sampling design. The spatial scales were regions, stream sites and stream sections within sites; temporal scales were years (1997–99), seasons (dry versus wet seasons) and days within seasons. 2. Spatiotemporal variability in total abundance and species richness was greater during the wet season, especially at small scales, and tended to obscure site‐ and region‐scale differences, which were more conspicuous during the dry season. Total abundance and richness were greater in the dry season, reflecting the effects of spate‐induced disturbance during the wet season. Species diversity showed little variation at the seasonal scale, but variability at the site scale was apparent during both seasons. 3. Despite marked variations in monsoonal rainfall, inter‐year differences in macroinvertebrate richness and abundance at the site scale during the wet season were minor. Inter‐year differences were only evident during the dry season when streams were at base flow and biotic interactions may structure assemblages. 4. Small‐scale patchiness within riffles was the dominant spatial scale of variation in macroinvertebrate richness, total abundance and densities of common species, although site or region was important for some species. The proportion of total variance contributed by small‐scale spatial variability increased during the dry season, whereas temporal variability associated with days was greater during the wet season. 5. The observed patterns of spatiotemporal variation have implications for detection of environmental change or biomonitoring using macroinvertebrate indicators in streams in monsoonal regions. Sampling should be confined to the dry season or, in cases where more resources are available, make use of data from both dry and wet seasons. Sampling in more than one dry season is required to avoid the potentially confounding effects of inter‐year variation, although variability at that scale was relatively small.     

The influence of flow and season upon leaf-litter breakdown in monsoonal Hong Kong streams

We investigated the effects of water extraction and flow regime on leaf litter breakdown in 10 Hong Kong streams experiencing varying degrees of extraction: five with severe extraction (>85% downstream flow removed), four with moderate extraction (20–85% removed), and one without extraction (~0% removed). Breakdown rates, macroinvertebrate assemblages and nutrients were compared in upstream (reference) reaches and downstream reaches with reduced flow in each stream during high-flow (wet season) and low-flow (dry season) conditions to determine whether responses to flow were influenced by Hong Kong’s monsoonal climate. Litter breakdown was slower in downstream reaches during the wet season, and discharge was the main determinant of breakdown rate in both reaches across the 10 study streams. In contrast, inter-reach differences in breakdown during the dry season were small and unrelated to flow. Instead, breakdown rates among streams were influenced by the abundance of detritivorous shredders and chironomids (Chironominae), as well as phosphate concentrations, indicating the importance of biological processing of litter under low-flow conditions. Although litter breakdown rates appear to offer a promising functional indicator of flow-reduction impacts during the wet season, the pattern seen reflected the physical effects of flow rather than any underlying biological processes, and the lack of response to flow reduction during the dry season was notable.     

Dynamics of habitats and macroinvertebrate assemblages in rivers of the Australian dry tropics

1. The dry tropics are characterised by episodic summer rainfall such that the majority of annual river flow occurs in a short period of time. This dryland hydrological cycle leads to variably connected channels and waterholes along the length of a river bed. 2. We investigated the seasonal changes in biophysical characteristics and macroinvertebrate assemblage composition in dry‐tropics rivers at 15 sites on four rivers, each sampled five times (representing one annual hydrological cycle), in the Burdekin catchment, north Queensland, Australia. 3. Assemblages and their temporal trajectories differed among seasons, sites and habitats, even within the same habitat and/or river. Wet season flooding did not appear to ‘reset’ assemblages, with post‐wet season assemblages differing between years. 4. We found no consistent pattern in taxonomic richness over time, and sites within rivers showed no consistent convergence or divergence (i.e. turnover) in macroinvertebrate assemblage composition. However, biophysical variables associated with the rigours of the late dry season had significant effects on macroinvertebrate assemblages, highlighting the variable and often harsh conditions of dry‐tropics rivers. Underlying these patterns were different resistance and resilience traits of invertebrates (such as colonisation and establishment abilities), as well as the local‐scale effects of biophysical variables. 5. The dynamic nature of dryland rivers presents major challenges to monitoring programmes, and our results suggest a more complex scenario for monitoring and management than previously described.     

The Effect of Riffle-Scale Environmental Variability on Macroinvertebrate Assemblages in a Tropical Stream

In a tropical stream (at the Soberaní;a National Park, Panama), different environmental factors were quantified in riffle habitats (water characteristics: velocity, depth, turbulence, and direction; stone characteristics: surface area, sphericity, and degree of burial; and others: substrate type, and canopy cover). Characteristics of macroinvertebrate assemblages (mean density of individuals, mean taxon richness, and cumulative taxon richness in three stones at each riffle) were related to both mean values and variability of these environmental factors at riffle scale. Macroinvertebrate density was higher in shallow, fast flowing, stony riffles, with low variability in dominant substrate type. Taxon richness was also higher in shallow riffles with loose, not buried stones, and water direction more or less parallel to the bank. Environmental variability resulted as important as mean values of environmental factors to explain variation in macroinvertebrate assemblages. This is the first study, to our knowledge, that quantifies substratum variability and demonstrates its influence on macroinvertebrate assemblages in a tropical stream.     

Macroinvertebrate survival during cessation of flow and streambed drying in a lowland stream

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Effects of Tea Plantations on Stream Invertebrates in a Global Biodiversity Hotspot in Africa

Tropical stream ecosystems in montane forest watersheds are important centers of endemism and diversity and provide essential ecosystem services. These habitats are subject to a variety of stressors, including the conversion of adjacent terrestrial habitats from forest to agriculture, but the impacts of these anthropogenic effects are largely unknown because of the paucity of studies in these systems. In montane habitats in the wet tropics, large-scale cultivation of tea is common and can represent an important source of income at local and national scales. However, little is known about how tea cultivation impacts adjacent stream ecosystems. In this study, we examine stream macroinvertebrate assemblages in a biodiversity hotspot the East Usambara Mountains, Tanzania. Specifically, we compare diversity of macroinvertebrate assemblages found on cobbles in stream riffles in watersheds dominated by forest with those surrounded by tea cultivation. We found that streams surrounded by tea were characterized by significantly lower dissolved oxygen and had lower total estimated species richness and number of families. Furthermore, the richness of invertebrate taxa known to be sensitive to anthropogenic disturbance were substantially reduced in tea streams and general assemblage-level analysis shows significant differences in the composition of macroinvertebrate assemblages between tea and forested streams. Our results suggest that tea cultivation may reduce stream habitat quality and biodiversity in the East Usambaras. Further research is needed to evaluate the effects of tea cultivation on streams over longer times scales and to address methods for minimizing negative effects of agriculture on montane stream communities.     

Geometry of biodiversity patterning: assemblages of benthic macroinvertebrates at tributary confluences

We assessed whether tributaries in upland catchments (=watersheds) affected assemblages of benthic macroinvertebrates in mainstems, as has been reported in northern hemisphere systems. Eight confluences of small to medium streams (stream orders 1–4, 2.2–10.8 m wide) were studied in the Acheron River basin in Victoria, Australia. For each confluence, two transects were sampled at each of five zones relative to the confluence: two zones upstream in the mainstem, one zone upstream in the tributary, one zone at the confluence and one zone downstream in the mainstem. Surveys were conducted in both high-flow and low-flow conditions. In mainstems, there was no change in macroinvertebrate density, taxonomic richness or functional feeding group composition downstream relative to upstream of the confluences. While tributaries statistically had distinctive benthic macroinvertebrate assemblages compared to mainstems, these distinctions were small. In low flows, densities in tributaries were substantially lower than in mainstems, but densities during high flows were more similar (albeit only about one-third as high as in low flow) in tributaries and mainstems. An inverse pattern was evident for taxonomic richness, where richness in tributaries and mainstems was similar in low flows but was greater in mainstems than in tributaries in high flows. We found little evidence of tributary effects in macroinvertebrate assemblages in this basin, which is at odds with some previous results from other continents. To explain this divergence, we suggest a conceptual model outlining factors that control variation in effects of tributaries on assemblages of benthic macroinvertebrates in mainstems.     

Effects of channelisation on stream habitat in relation to a spate and flow refugia for macroinvertebrates in northern Japan

1. The effects of channelisation on macroinvertebrates were examined in relation to a spate and flow refugia. Habitat components that can function as flow refugia were identified in a small, low‐gradient stream in northern Hokkaido, Japan. 2. Macroinvertebrates and their habitat characteristics (depth, current velocity and substratum) were sampled and measured in natural and channelised sections on three occasions: before, during and immediately after a spate. For macroinvertebrate sampling and habitat measurements, five (riffle, glide, pool, backwater and inundated habitats) and three (channelised‐mid, channelised‐edge and inundated habitats) habitat types were classified in the natural and channelised section, respectively. 3. The rate of velocity increase with discharge was compared among habitat types to determine which habitat types were less affected by increased discharge. The rate was the highest in riffles followed by glides and channelised‐mids. Backwaters maintained low current velocity even at high flow. In addition, current velocity in both natural and channelised inundated habitats was low relative to other habitat types during the spate. 4. Through the spate, total density of macroinvertebrates in channelised‐mids and taxon richness in both channelised‐mids and edges decreased. In the natural section, however, such a significant decrease was not found except for taxon richness in pools. This indicated that the spate had a greater impact on assemblages in the channelised section. Riffle assemblages exhibited a rapid recovery immediately after the spate, suggesting the existence of flow refugia in the natural section. Among the habitat types we examined, backwaters and inundated habitats appeared to have acted as flow refugia, because these habitats accumulated macroinvertebrates during the spate. 5. The lower persistence of the macroinvertebrate assemblage in the channelised section was attributable to the lower availability of flow refugia such as backwaters and inundated habitats. Our results emphasised the importance of considering flow fluctuations and refugia in assessing the effects of channelisation. In addition, the lateral heterogeneity of stream channels should be considered in stream restoration and management.     

High spatial variability biases the space-for-time approach in environmental monitoring

The space-for-time approach is widely used in fundamental and applied ecology but assemblages from some habitats are highly variable. For example, streams may show marked spatio-temporal changes in the taxonomic composition of the macroinvertebrate assemblages. We exemplify the effect of the temporal component ‘season’ on some assemblage-derived stream quality assessment metrics under the assumptions of the space-for-time and the replicated samples approaches. Benthic macroinvertebrates were sampled in spring, summer, and fall from two stream types, namely streams in the Pleistocene sediments of the alpine foothills and small fine substrate dominated siliceous highland rivers in southern and central Germany. As exemplified for ASPT and the German multimetric index (MMI), the data showed no effect of season when samples were regarded as independent, whereas stream quality decreased between spring and fall in the replicated samples approach. The transformation of MMI to rank-ordered stream quality classes depicted a decrease in perceived stream quality in 29% and 54% of the sites by summer and early fall, respectively, when compared to spring samples. We thus suggest (1) to test seemingly robust metrics in a repetitive measures approach for other stream types and regions, and (2) to standardize the sampling season for ecological quality assessment. Based on this example, we assume that many subtle, but significant, environmental trends are still to be detected in highly heterogeneous habitats from various ecosystems.     

The influence of reduced flow during a drought on patterns of variation in macroinvertebrate assemblages across a spatial hierarchy in two lowland rivers

1. With the aim of determining whether patterns of variation in macroinvertebrate assemblage composition across the hierarchy of spatial units in two lowland rivers changed during a supra‐seasonal drought (1997–2000), patterns during a reduced flow season (1999–2000) were compared with those during two preceding higher flow seasons (1997–98 and 1998–99) using samples from the Glenelg and Wimmera Rivers, two lowland regulated rivers in western Victoria, Australia. 2. We hypothesised that (i) differences between reaches would increase during the reduced flow season owing to decreased hydrological connectivity, (ii) differences between the habitats would decrease because the cessation of flow in run habitats should cause them to become more similar to pool habitats and (iii) differences between microhabitats would also decrease because of reduced scour of inorganic substrata and large woody debris. 3. During each season, macroinvertebrates were sampled from three microhabitats (sand/silt substratum, large woody debris and macrophytes) that were hierarchically nested within a run or pool habitat and within one of three reaches within each river. A range of physico‐chemical variables was also sampled. 4. Analysis of similarity showed that assemblage composition in both rivers during the higher flow seasons differed more among microhabitats than other spatial units. However, during the reduced flow season, assemblage composition in the Wimmera River differed most among reaches. This change in pattern was associated with the combined effects of decreased flow and longitudinal increases in salinity. In contrast, the fauna of the Glenelg River appeared to be resistant to the effects of the reduced flow season, owing to limited decline in water quality despite lower river discharge. 5. As salinisation and poor water quality in the Wimmera River result from human activities in the catchment, these results support the idea that human impacts on rivers can change macroinvertebrate scaling patterns and exacerbate the effects of drought beyond the tolerance of many riverine macroinvertebrates.     

Stream macroinvertebrate assemblage uniformity and drivers in a tropical bioregion

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Seasonal variation of black fly (Diptera: Simuliidae) species diversity and community structure in tropical streams of Thailand

Seasonal variation of the physicochemical conditions of streams plays an important role in aquatic insect diversity and community structure. Asian tropical streams profoundly change between seasons due to the effects of monsoons. However, little is known about how these changes affect aquatic insect diversity and community structure. The objectives of this study are to examine seasonal variations of the physicochemical conditions in tropical streams in Thailand and to assess the effects of these changes on black fly community structure and diversity. Black flies were sampled and physicochemical conditions recorded at eight sites between December 2007 and December 2008. A total of 10 black fly species were found. Comparisons of the streams between seasons revealed that physical conditions related to rainfall rate were significantly different. Canonical correspondence analysis differentiated sampling sites from each season. Streams in the rainy season were faster and deeper, with higher discharge and conductivity than those of the cold and dry seasons. Species richness was significantly higher in the rainy season than in the cold and dry seasons ( F  = 6.23, P  = 0.004). Community structure profoundly changed between the low-flow season (cold and dry) and high-flow season. Black fly species found predominantly in the low-flow season ( Simulium siamense "cytoform A", S. aureohirtum ) decreased dramatically during the high-flow season. In contrast, species found at high frequency during the high-flow season ( S. nakhonense , S. angulistylum ) disappeared in the low-flow season. The study demonstrates the importance of seasonal variation of stream conditions on black fly community structure and diversity.     

Seasonal patterns of stream macroinvertebrate communities in response to anthropogenic stressors in monsoonal Taiwan

Freshwater ecosystems are affected by a variety of anthropogenic stressors. Temporal variability of biotic communities in these ecosystems makes it difficult to accurately assess the impacts of specific stressors, which has seldom been considered in understudied regions of Asia. We studied the seasonal effects of anthropogenic stressors on stream macroinvertebrates based on sampling every three months over two years at five stream sites in central Taiwan. Several macroinvertebrate metrics (taxon richness, Shannon diversity index, and relative abundance of Trichoptera) were lower during the wet season than the dry season. The presence of dams caused changes in the structure of macroinvertebrate communities, decreased the seasonal variability in relative abundances of Trichoptera, as well as resulted in lower dissolved inorganic nitrogen concentrations and larger substrate size. The presence of urban areas had less or no influence on environmental factors and structural changes. However, significantly lower total abundance, taxon richness, and relative abundance of Trichoptera occurred in the presence of either dams and/or urban areas. One key management implication from the present study is that bioassessment utilizing macroinvertebrates should be facilitated by awareness of the potential role of temporal factors on the effects of anthropogenic stressors, especially in monsoonal Asia.     

Seasonal dynamics of macroinvertebrate communities in a semiarid saline spring stream with contrasting environmental conditions

Semiarid saline streams are rare aquatic ecosystem types. Their constituent biota is expected to have adapted evolutionarily to strong hydrological variability and salinity stress; however, their ecology is not well known. In this study, we quantify the seasonal changes in the structure of the macroinvertebrate community in the Reventón Rambla (south-eastern Spain), a permanent saline spring stream which is included in a drainage system consisting of ephemeral dry channels (so-called “ramblas”). Seasonal patterns of community structure were studied in two reaches with contrasting environmental regimes using univariate and multivariate statistics. The upstream site showed more stable environmental conditions than the downstream site, and both sites also differed with regard to species richness, and structural and functional group attributes. On a seasonal basis, community dissimilarity was high during periods when both sites were isolated during summer droughts but dissimilarity decreased when both sites were connected through surface flow. Furthermore, the communities tended to show cyclical trajectories in multivariate ordination space. Rather than being related to salinity stress, these patterns seemed to track the hydrological disturbance regime of this rambla system. Spates tended to disrupt communities, while signs of recovery were evident during low-flow periods. Results suggest that salinity fluctuation does not pose a severe abiotic constraint to these adapted macroinvertebrate communities. Their suits of functional properties provide them with the necessary traits to recover quickly from natural disturbance. While human-caused salinization of streams severely impacts communities eventually reducing their recovery potential, our results suggest that communities in natural saline streams may show similar responses to hydrological disturbance as communities from non-saline streams.     

Influence of flow permanence on headwater macroinvertebrate communities in a Cumberland Plateau watershed,USA

Forested headwater systems provide critical habitat needs for diverse macroinvertebrate faunas globally. This study compared macroinvertebrate community structure between ten temporary and perennial stream channels in a Cumberland Plateau (USA) watershed. Macroinvertebrates were sampled in winter and spring over a 2-year period. Several macroinvertebrate taxa displayed distinct abundance patterns with either increasing or decreasing stream permanence. There were no individual taxa, however, that was common or abundant in one flow regime yet absent in the other. Of the 108 taxa totally collected, only nine and eight were not obtained from the temporary and perennial channels, respectively. There were several functional differences revealed between flow regimes, namely higher densities in the perennial channels for scrapers and filtering-collectors and five of six richness variables except shredders. Overall, this study revealed taxonomic similarity of macroinvertebrate assemblages between temporary and perennial channels but also subtle downstream functional changes that are typical of forested headwater systems. This implies that the longitudinal hydrologic gradient was relatively shallow and that most taxa persist across the flow regime.     

Macroinvertebrate communities in Phragmites australis (Cav.) Trin. ex Steud. reed beds and open bank habitats in central victorian streams in Australia

Reed invasion is a common phenomenon of open streams with disturbed riparian vegetation in river catchments. Knowledge of the effects of such vegetation change on aquatic communities is fundamental to river management. Macroinvertebrate fauna in Phragmites australis (Cav.) Trin. ex Steud. and open bank habitats were examined in three rivers in central Victoria in order to understand the effect of such littoral habitat on macroinvertebrates. Data were analysed using Partially Nested Factorial ANOVA with season, river and habitats as main effects. Habitat structure had a significant effect (p<0.05) on macroinvertebrate species richness, however this was not seasonally consistent across the three rivers. There was a significant increase (p<0.05) in macroinvertebrate taxa richness in Phragmites habitats during winter and spring seasons. Total abundance of taxa showed no consistent significant differences in the two habitats. Results of Canonical Analysis of Principle Coordinates indicated significant differences (p<0.05) in macroinvertebrate assemblages between Phragmites and bare bank habitats in all seasons. Habitat selection by taxa could be related to the microphysical environment of the habitats. This study suggests that reed beds create important littoral habitat structures which support diverse macroinvertebrate assemblages.     

Natural flow regime,temperature and the composition and richness of invertebrate assemblages in streams of the western United States

1. We tested how strongly aquatic macroinvertebrate taxa richness and composition were associated with natural variation in both flow regime and stream temperatures across streams of the western United States. 2. We used long‐term flow records from 543 minimally impacted gauged streams to quantify 12 streamflow variables thought to be ecologically important. A principal component analysis reduced the dimensionality of the data from 12 variables to seven principal component (PC) factors that characterised statistically independent aspects of streamflow: (1) zero flow days, (2) flow magnitude, (3) predictability, (4) flood duration, (5) seasonality, (6) flashiness and (7) base flow. K‐means clustering was used to group streams into 4–8 hydrologically different classes based on these seven factors. 3. We also used empirical models to estimate mean annual, mean summer and mean winter stream temperatures at each stream site. We then used invertebrate data from 63 sites to develop Random Forest models to predict taxa richness and taxon‐specific probabilities of capture at a site from flow and temperature. We used the predicted taxon‐specific probabilities of capture to estimate how well predicted assemblages matched observed assemblages as measured by RIVPACS‐type observed/expected (O/E) indices and Bray–Curtis dissimilarities. 4. Macroinvertebrate taxon richness was only weakly associated with streamflow and temperature variables, implying that other factors more strongly influenced taxa richness. 5. In contrast to taxa richness, taxa composition was strongly associated with streamflow and temperature. Predictions of taxa composition (O/E and Bray–Curtis) were most precise when both temperature and streamflow PC factors were used, although predictions based on either streamflow PC factors or temperature alone were also better than null model predictions. Of the seven aspects of the streamflow regime we examined, variation in baseflow conditions appeared to be most directly associated with invertebrate biotic composition. We were also able to predict assemblage composition from the conditional probabilities of hydrological class membership nearly as well as Random Forests models that were based directly on continuous PC factors. 6. Our results have direct implication for understanding the relative importance of streamflow and temperature in regulating the structure and composition of stream assemblages and for improving the accuracy and precision of biological assessments.     

Macroinvertebrate Assemblages of an Andean High‐Altitude Tropical Stream: The Importance of Season and Flow

We studied the Piburja stream, a high‐altitude tropical stream in Ecuador. Our main goals were to determine whether the macroinvertebrate community composition and abundance differed between seasons, reaches and velocity patches. Likewise, we aimed to examine the importance of the hydrological regime in determining these differences. Flow was significantly higher in the wet season, but the stability of flow was higher in the dry season. There was a strong increase in macroinvertebrate community metrics (richness, density and diversity) in the dry season. Seasons and velocity patches better explained the community composition. Reaches did not show differences at the community level, but some taxa showed significant differences among reaches. Our findings differed from those published in previous studies that have suggested that mountain tropical streams are non‐seasonal. (© 2011 WILEY‐VCH Verlag GmbH & Co. KGaA, Weinheim)     

A preliminary investigation of temporal patterns in semiarid ant communities: Variation with habitat type

Abstract This study reports on preliminary findings of habitat‐contingent temporal variability in ant assemblages in Purnululu National Park in northern Australia's semiarid tropics, by sampling at the end of the dry season (October 2004) and the end of the wet season (April 2005). Six grids of 15 pitfall traps were established in each of the spinifex, sandplain and gorge habitats. Community composition was dominated by behaviourally dominant ants (Iridomyrmex spp.) and climate specialists (Melophorus and Meranoplus spp.). Ant activity was higher in the wet season sampling period, with greater species richness and abundance. Interestingly, temporal variation in ant assemblage richness, abundance and composition varied markedly with habitat type. While there were large differences between sampling periods for the spinifex and sandplain habitat, this was not the case in the gorges. These temporal changes in ant assemblages are postulated to be linked with major environmental differences between the two sampling periods, driven by seasonal climatic conditions. It is likely that these changes influenced the ant assemblages through species differences in physiological tolerance levels, ecological requirements and competitive ability. This study demonstrates the need, in highly seasonal environments, to consider the temporal context of studies in relation to habitat type, particularly when undertaking biodiversity surveys and monitoring.     

Thermal and hydrologic responses to climate change predict marked alterations in boreal stream invertebrate assemblages

Air temperature at the northernmost latitudes is predicted to increase steeply and precipitation to become more variable by the end of the 21st century, resulting in altered thermal and hydrological regimes. We applied five climate scenarios to predict the future (2070–2100) benthic macroinvertebrate assemblages at 239 near‐pristine sites across Finland (ca. 1200 km latitudinal span). We used a multitaxon distribution model with air temperature and modeled daily flow as predictors. As expected, projected air temperature increased the most in northernmost Finland. Predicted taxonomic richness also increased the most in northern Finland, congruent with the predicted northwards shift of many species’ distributions. Compositional changes were predicted to be high even without changes in richness, suggesting that species replacement may be the main mechanism causing climate‐induced changes in macroinvertebrate assemblages. Northern streams were predicted to lose much of the seasonality of their flow regimes, causing potentially marked changes in stream benthic assemblages. Sites with the highest loss of seasonality were predicted to support future assemblages that deviate most in compositional similarity from the present‐day assemblages. Macroinvertebrate assemblages were also predicted to change more in headwaters than in larger streams, as headwaters were particularly sensitive to changes in flow patterns. Our results emphasize the importance of focusing protection and mitigation on headwater streams with high‐flow seasonality because of their vulnerability to climate change.