2005. Does macrophyte fractal complexity drive invertebrate diversity, biomass and body size distributions? Á/ Oikos 111: 279 Á/290.Habitat structure is one of the fundamental factors determining the distribution of organisms at all spatial scales, and vegetation is of primary importance in shaping the structural environment for invertebrates in many systems. In the majority of biotopes, invertebrates live within vegetation stands of mixed species composition, making estimates of structural complexity difficult to obtain. Here we use fractal indices to describe the structural complexity of mixed stands of aquatic macrophytes, and these are employed to examine the effects of habitat complexity on the composition of freeliving invertebrate assemblages that utilise the habitat in three dimensions. Macrophytes and associated invertebrates were sampled from shallow ponds in southwest England, and rapid digital image analysis was used to quantify the fractal complexity of all plant species recorded, allowing the complexity of vegetation stands to be reconstructed based on their species composition. Fractal indices were found to be significantly related to both invertebrate biomass Á/body size scaling and overall invertebrate biomass; more complex stands of macrophytes contained a greater number of small animals. Habitat complexity was unrelated to invertebrate taxon richness and macrophyte surface area and species richness were not correlated with any of the invertebrate community parameters. The biomass Á/body size scaling relationship of lentic macroinvertebrates matched those predicted by models incorporating both allometric scaling of resource use and the fractal dimension of a habitat, suggesting that both habitat fractal complexity and allometry may control density Á/body size scaling in lentic macroinvertebrate communities.

1. Broad-scale assessments of biodiversity often rely on the use of surrogate taxa, whose reliability has rarely been tested, particularly in freshwater systems. Here we use data from 46 ponds in two regions of the U.K. to explore the performance of macroinvertebrate taxa as surrogates for the rapid assessment of pond biodiversity. For the four dominant taxonomic groups in these ponds (Chironomidae, Coleoptera, Gastropoda and Trichoptera) we explore cross-taxon species richness relationships in each of the two regions, and also determine the degree of concordance between the different taxa in accurately representing the similarity relationships between pond assemblages. 2. Patterns of cross-taxon congruence in species richness were highly variable among taxa and study sites, making the use of a single taxon as a predictor of overall macroinvertebrate species richness problematic. In contrast, all four taxa show >70% congruence with the pattern of community similarity between sites resulting from the entire macroinvertebrate dataset, this result being consistent within and between regions. Canonical correspondence analysis demonstrated that all taxa were related in a similar manner to measured environmental parameters, meaning that limited additional ecological information is gained by including a wider range of pond taxa in rapid site assessment. 3. Single taxonomic groups can, therefore, perform consistently as indicators of community similarity between ponds, and no one taxon dramatically outperforms any other in this respect. The relative merits of the four focal taxa as surrogates for pond invertebrate assemblage composition are discussed with reference to ease of survey, ease of identification and ecological range occupied. 4. It is suggested that Coleoptera have a number of advantages as a surrogate taxon, being diverse, easily sampled, readily identified, taxonomically stable, ecologically well understood and occurring across a wide spectrum of pond types. They are therefore recommended for use as a focal group in rapid pond biodiversity assessments, employing an approach such as ours, which examines patterns of assemblage similarity, rather than species richness alone.

Summary 1.Nestedness is a composite property of many suites of biotas. Such patterns may be driven by dispersal limitation, species-area relationships, hierarchical niche requirements, or occur as an artefact of passive sampling. Despite its widespread occurrence, few studies have explored the factors underlying nested subset structure, and ecological distinctions between nested and non-nested (idiosyncratic) taxa within a region have been largely ignored. 2. Macroinvertebrate assemblages from 45 heathland ponds in south-west England are used to (i) unravel the relative importance of processes driving nested subset structure and (ii) test spatially explicit hypotheses concerning the response of nested and idiosyncratic taxa to parameters shown to structure assemblage-level nestedness. 3. Despite being dominated by taxa with good powers of inter-site dispersal, pond macroinvertebrate assemblages were found to be significantly nested. This nesting was not due to passive sampling, and was best explained by pond area, with habitat parameters and isolation being of secondary importance. 4. The spatial responses of nested and idiosyncratic taxa matched predictions; nested taxa showed strong spatial structure, which was reduced when the effects of pond area and habitat were removed. In contrast, a greater proportion of idiosyncratic taxa were completely spatially random and exhibited weaker responses to factors that structure assemblage level nestedness. 5. Nested and idiosyncratic species tend to differ ecologically: idiosyncratic taxa generally possess broad ecological tolerance and good dispersal capacity, whilst nested species are more likely to have narrow tolerances or limited powers of dispersal. 6. Factors structuring nestedness in ponds can be viewed as probabilistic filters which act to limit the spatial distribution of species with narrow ecological tolerance or low dispersal tendency. Nestedness analysis alone fails to elucidate processes that structure assemblage composition. The additional use of spatially explicit analyses is important if processes that generate nested pattern across a region are to be understood.

ABSTRACT1. A high proportion of ponds are temporary in nature, although the biodiversity value of such sites is still poorly recognized. This work explores the plant and macroinvertebrate communities of 76 temporary and fluctuating water bodies in two regions of southern England (Lizard Peninsula, Cornwall and New Forest, Hampshire) that have retained high pond densities. The ecology and conservation status of sites is examined, and comparisons made with ponds elsewhere in England and Wales.2. Lizard and New Forest ponds supported 119 plant and 165 macroinvertebrate species respectively. Patterns of community similarity for plants and macroinvertebrates were highly concordant, taxa being related in a similar manner to measured environmental variables including pond area, depth, pH and water chemistry.3. Patterns of pond occupancy revealed that most species were locally rare, over half occurring in less than 10% of ponds, and less than 10% being recorded from more than 50% of sites. More than 50% of ponds supported at least one nationally rare plant and almost 75% at least one nationally rare macroinvertebrate. These taxa occupied a wide range of pond types in each region, and did not have predictably different ecologies from common species.4. Comparisons with ponds elsewhere in England and Wales revealed that Lizard and New Forest communities are nationally distinct, being most similar to ponds in areas of low intensity agriculture elsewhere in western Britain. Individual ponds in both regions supported more nationally rare taxa, on average, than ponds sampled in the national survey.5. Ponds in the two areas have high conservation value, both regionally and nationally, supporting almost 75% of the global species richness of temporary ponds sampled across England and Wales. Within each region, many taxa are found in relatively few sites, and effective conservation of pond biodiversity will require a regional management approach.