The intermediate disturbance hypothesis has been influential in the development of ecological theory and has important practical implications for the maintenance of biodiversity but has received few rigorous tests. WC tested the hypothesis that maximum taxon richness of macroinvertebrates will occur in communities subject to intermediate levels of disturbance at 54 stream sites that differed in the frequency and intensity of flood-related episodes of bed movement. Our results support the intermediate disturbance hypothesis, with both highly mobile and relatively sedentary taxa conforming to the predicted bell-shaped curve. Taxon richness was not related to habitat area (stream width), distance from the headwater, or the diversity of microhabitats (particle size categories) but was significantly and negatively related to the proportion of the substratum made up of small particles. Of all the factors measured, however, bed disturbance was by far the best at accounting for variation in taxonomic richness. We also quantified several kinds of potential refugia for invertebrates and found a positive relationship between richness and a refugia axis that combines amount of dead space with proportion of large substratum particles.
1. The habitat templet approach depends on defining templet axes appropriate to the organism(s) of interest, predicting the traits of species associated with different parts of the templet, and testing these predictions in a range of habitats whose positions in the templet have been determined. 2. In this study of thirty‐five benthic insect taxa at fifty‐four tributary sites of the Taieri River on the South Island of New Zealand, we chose as the temporal axis the intensity/frequency of disturbance, defined in terms of bed movement during high discharge events. As the spatial axis, we postulated that three features would provide refugia and therefore ameliorate disturbance—percentage of the bed with low shear stress, percentage of the bed made up of large substratum particles and availability of interstitial space in the bed—from which we derived a combined multivariate refugium axis. 3. More disturbed communities contained a significantly higher percentage of individuals possessing the following traits: small size, high adult mobility, habitat generalist (each predicted to confer resilience in response to disturbance), clinger, streamlined/flattened and with two or more life stages outside the stream (each predicted to confer resistance in the face of disturbance). When analyses were performed on the percentage of taxa having particular traits, the predicted positive relationships with average bed movement were found for high adult mobility and habitat generalist traits. 4. The percentage of variance in trait scores explained by intensity of disturbance was generally higher in sites with less refugia available and lower in sites further from the headwaters. The percentage of variance explained was higher in sites recently subject to a major high discharge disturbance, suggesting that disturbances tend to strengthen the pattern of preponderance of resilience/resistance traits. 5. We mapped insect taxa onto the two‐dimensional templet, following Grime et al.’s triangular terrestrial plant classification. The full variety of resistance and resilience traits were represented in insect species throughout the templet, but taxa associated with more disturbed conditions generally displayed a larger number of resilience and resistance traits, combined, than taxa associated with more stable stream beds.
1. The relationship between land use and stream conditions was investigated, including physicochemistry, the availability of primary food resources and species richness, species composition and trophic structure of stream macroinvertebrate communities. The survey involved eight subcatchments of the Taieri River (New Zealand) encompassing reasonably homogeneous examples of four major land uses: native forest, native tussock grassland, plantations of introduced pine and agricultural pasture. 2. Each land use was represented by two subcatchments, each subcatchment by two to four tributaries, and each tributary by two to three sampling sites. These three sampling scales each represent typical designs for stream community studies. By recording responses at all scales, it can be determined explicitly whether the scale of sampling influences interpretation of community structure. 3. Elevation, riffle length, proportion of large substrata in the bed, total phosphorus and alkalinity were significantly related to land use, as were canopy cover and the relative abundance of leaves and wood in the streams. Principal components analysis of invertebrate density data identified nine orthogonal community types, the distributions of two of which were significantly related to land use. The role played by browsers and shredders in the stream community depended on land use. 4. Primary analysis was at the level of the tributary. When it focused on sites within tributaries, more variables were related to land use and at a higher level of significance. This was largely a result of enhanced statistical power due to increased replication. When whole subcatchments were the focus of attention, statistical power was so low, even with six to eleven subsamples to generate overall means, that few significant patterns could be identified. However, the community patterns that were revealed were similar whatever the scale of sampling.
1. The physical characteristics of two contrasting streams, and habitat types within these streams, are described in terms of a two-dimensional physical habitat templet in which disturbance frequency and the availability of spatial refugia are the temporal and spatial axes. 2. It is predicted that habitats experiencing a high disturbance frequency and low refuge availability will be characterized by a low invertebrate spedes diversity, a low biomass of epilithic algae and particulate organic matter and a community made up of mobile, weedy species. Habitats having a low disturbance frequency and high refuge availability will be characterized by a diverse community containing sedentary and specialist spedes, with high algal and particulate organic matter levels. 3. A lower median substrate particle size and higher shear stress regime in Timber Creek were indicative of a higher disturbance frequency than in the Kyebum. Substrate diversity was lower in Timber Creek than in the Kyeburn and indicated that the availability of refugia was lower in Timber Creek. In both streams, pools were found to have a higher disturbance frequency and iower availability of refugia than riffles. 4. Invertebrate spedes diversity, the biomass of epilithic algae and particulate organic matter and the representation of sedentary spedes, filter feeders and shredders were higher in the more temporally stable and spatially heterogeneous Kyeburn. The community of Timber Creek, frequently disturbed and having low refuge availability, had a high proportion of mobile and weedy spedes, with the highly mobile, generalist-feeding Deleatidium spp. (Ephemeroptera; Leptophlebiidae) being the most dominant organisms. 5. The predictions made about stream community structure and spedes characteristics in relation to disturbance frequency and the availability of spatial refugia are generally supported. Now a larger scale investigation is required to test the generality of the predictions. We conclude that the habitat templet approach offers a sound framework within which to pose questions in stream ecology. and their adaptations (Wiens, 1976). Therefore, as
1. Persistence and stability of lotic invertebrate communities were determined at an annual time scale over a 9‐year period (1990–98) at 26 river sites over the northern half of New Zealand. In addition, a number of water quality variables were measured monthly and flow information collected continuously over the same period at each site. 2. The aim of the study was to describe the levels of interannual variability in invertebrate communities, and relate community changes to variability in environmental conditions. The consequences of this temporal variability for the sensitivity of predictive models were also investigated. 3. Levels of change in environmental conditions varied significantly between years, but were relatively similar across sites. In contrast, community persistence (similarity between years in species assemblage composition), and stability (similarity between years with respect to relative abundance of species in the community) both varied significantly between sites, but changes between years were similar. Community stability was highest at sites with relatively harsh flow conditions (high coefficient of variation, high relative size of floods), and was also greater in communities dominated by Ephemeroptera. 4. Relationships between change in environmental conditions and changes in community composition and structure were relatively weak for most individual sites. However, when average levels of change for each of the 26 sites were used, communities showed greater persistence under conditions where flow conditions remained relatively constant. Water quality changes had no significant effect on community persistence when assessed for all 26 sites combined. 5. Results from this study suggest that lotic invertebrate communities fluctuate around a relatively stable state, at least over a 9‐year period. However, the extent of interannual variation in community composition and structure observed, along with the relatively low degree of cluster fidelity observed within a single region, suggests that predictive models based on reference site conditions extrapolated over several years should be applied with caution in New Zealand streams.
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