Activity and satiation state in Pterostichus melanarius: an experiment in different agricultural habitats

1. A capture–recapture experiment was conducted to examine the effects of satiation state on the activity and foraging efficiency of the carabid beetle Pterostichus melanarius Ill., in four different habitats (a hedge, a transition zone between the hedge and the adjacent wheat crop, the margin of the wheat crop, and the centre of the wheat crop) in an agricultural ecosystem in France. 2. Prey biomass and density decreased from the centre of the hedge to the crop margin and to the centre of the crop, where prey density was low but prey biomass was moderate. Starved and satiated beetles were released in circular enclosures and recaptured using pitfall traps for 6 days. 3. Immediately after release, starved P. melanarius were significantly more active than satiated beetles, but this difference disappeared after the first day. At each recapture date, starved individuals had gained weight whereas satiated individuals had lost weight. The analysis of activity and weight variation at the end of the first day indicated that the centre of the crop seemed to be the most favourable habitat for starved individuals to find food rapidly, whereas the behaviour of satiated individuals was identical in all habitats. 4. This experiment shows that habitat use of P. melanarius is flexible. The results are discussed in the context of the classical view of foraging behaviour in carabid beetles. The activity and foraging efficiency in P. melanarius in different habitats can be explained as the result of the interaction between individual satiation state, prey availability in the habitat, and particular preferences of the species.     

Effects of Fragmentation and Sea-Level Changes upon Frog Communities of Land-Bridge Islands off the Southeastern Coast of Brazil

We investigate the composition of anuran communities of land-bridge islands off the southeastern coast of Brazil. These islands provide natural long-term experiments on the effects of fragmentation in the Brazilian Atlantic Forest (BAF). We hypothesize that Pleistocene sea-level changes, in combination with other abiotic variables such as area and habitat diversity, has affected anuran species richness and community composition on these islands. Data from the literature and collections databases were used to produce species lists for eight land-bridge islands and for the mainland adjacent to the islands. We assess the effects of area, number of breeding habitats and distance to the mainland upon anuran species richness on land-bridge islands. Additionally we use nestedness analysis to quantify the extent to which the species on smaller and less habitat-diverse islands correspond to subsets of those on larger and more diverse ones. We found that area has both direct and indirect effects on anuran species richness on land-bridge islands, irrespective of distance to the mainland. However, on islands with comparable sizes, differences in species richness can be attributed to the number and quality of breeding habitats. Anuran communities on these islands display a nested pattern, possibly caused by selective extinction related to habitat loss. Common lowland pond-breeders were conspicuous by their absence. In the BAF, the conservation of fragments with a high diversity of breeding habitats could compensate for the generally negative effect of small area upon species richness. We suggest that sea-level changes have an important role in shaping composition of anuran species on coastal communities.     

Opposing deer and caterpillar foraging preferences may prevent reductions in songbird prey biomass in historically overbrowsed forests

Overbrowsing by ungulates decimates plant populations and reduces diversity in a variety of ecosystems, but the mechanisms by which changes to plant community composition influence other trophic levels are poorly understood. In addition to removal of avian nesting habitat, browsing is hypothesized to reduce bird density and diversity through reduction of insect prey on browse‐tolerant hosts left behind by deer. In this study, we excluded birds from branches of six tree species to quantify differences in songbird prey removal across trees that vary in deer browse preference. Early in the breeding season, birds preyed on caterpillars at levels proportional to their abundance on each host. Combining these data with tree species composition data from stands exposed to experimentally controlled deer densities over 30 years ago, we tested whether overbrowsing by white‐tailed deer reduces prey biomass long after deer densities are reduced. Our analysis predicts total prey availability in the canopy of regenerating forests is fairly robust to historic exposure to high deer densities, though distribution of prey available from host species changes dramatically. This predicted compensatory effect was unexpected and is driven by high prey abundance on a single host tree species avoided by browsing deer, Prunus serotina. Thus, while we confirm that prey abundance on host trees can act as a reliable predictor for relative prey availability, this study shows that quantifying prey abundance across host trees is essential to understanding how changes in tree species composition interact with ungulate browse preference to determine prey availability for songbirds.     

How to evaluate and reduce sampling effort for ants

Representative sampling of ant communities is time-consuming and laborious. Due to their manifold habitat requirements and sociobiological attributes, a suite of sampling methods is necessary to detect nearly all ant species in a given habitat. We analysed how well different combinations and intensities of sampling methods are suited to assess ant species richness and community patterns. Sampling occurred at 24 sites in mountain and floodplain habitats in Austria using pitfall traps, Winkler litter extraction, hand sampling of foragers, and colony sampling. Of these, pitfall traps delivered the largest species numbers. We drew 19 subsets of sampling approaches and compared these to the complete data. Many subsets that allowed for substantial reduction of work effort nonetheless reflected well the beta diversity patterns of the whole dataset. In contrast, alpha diversity assessment was more sensitive to reduced sampling effort. Pitfall traps turned out to be indispensable to collect ant species for biodiversity studies, but the optimal combination of sampling methods varied between habitat types. In rugged montane habitats pitfalls should be complemented at least by colony sampling. In floodplain forests the number of pitfall trap replicates can be reduced, but inclusion of Winkler samples is advisable.     

Factors affecting species number and density of dabbling duck guilds in North Europe

We addressed how species number and pair density in guilds of co-existing species is related to habitat structure, and to the abundance and diversity of food resources using the assemblage of seven species of dabbling ducks (genus Anas ) breeding in 60 lakes distributed over six regions in temperate north Europe
Partial correlation and multiple regression revealed that species richness was best predicted by habitat structural diversity as indexed by a principal component analysis based on 18 vegetation and lake characteristics, and by the abundance of aquatic and emergent prey We found no effect of lake size or prey size diversity on species richness Pair density was correlated with the percentage of shoreline with horsetails ( Equisetum ), by habitat structural diversity and by the abundance of emergent invertebrate prey Neither prey size diversity nor abundance of aquatic prey correlated with pair density Species richness and pair density in North European duck guilds vary both with habitat structure and prey availability     

Using demographic data to better interpret pitfall trap catches

The results of pitfall trapping are often interpreted as abundance in a particular habitat. At the same time, there are numerous cases of almost unrealistically high catches of ground beetles in seemingly unsuitable sites. The correlation of catches by pitfall trapping with the true distribution and abundance of Carabidae needs corroboration. During a full year survey in 2006/07 in the Lake Elton region (Volgograd Area, Russia), 175 species of ground beetles were trapped. Considering the differences in demographic structure of the local populations, and not their abundances, three groups of species were recognized: residents, migrants and sporadic. In residents, the demographic structure of local populations is complete, and their habitats can be considered "residential". In migrants and sporadic species, the demographic structure of the local populations is incomplete, and their habitats can be considered "transit". Residents interact both with their prey and with each other in a particular habitat. Sporadic species are hardly important to a carabid community because of their low abundances. The contribution of migrants to the structure of carabid communities is not apparent and requires additional research. Migrants and sporadic species represent a "labile" component in ground beetles communities, as opposed to a "stable" component, represented by residents. The variability of the labile component substantially limits our interpretation of species diversity in carabid communities. Thus, the criteria for determining the most abundant, or dominant species inevitably vary because the abundance of migrants in some cases can be one order of magnitude higher than that of residents. The results of pitfall trapping adequately reflect the state of carabid communities only in zonal habitats, while azonal and disturbed habitats are merely transit ones for many species of ground beetles. A study of the demographic structure of local populations and assessment of the migratory/residential status of particular carabid species are potential ways of increasing the reliability of pitfall trap information.     

Effects of habitat complexity on forest beetle diversity: do functional groups respond consistently?

We examined the responses of a beetle assemblage to habitat complexity differences within a single habitat type, Sydney sandstone ridgetop woodland, using pitfall and flight‐intercept trapping. Six habitat characters (tree canopy cover, shrub canopy cover, ground herb cover, soil moisture, amount of leaf litter, and amount of logs, rocks and debris) were scored between 0 and 3 using ordinal scales to reflect habitat complexity at survey sites. Pitfall trapped beetles were more species rich and of different composition in high complexity sites, compared with low complexity sites. Species from the Staphylinidae (Aleocharinae sp. 1 and sp. 2), Carabidae (Pamborus alternans Latreille), Corticariidae (Cartodere Thomson sp. 1) and Anobiidae (Mysticephala Ford sp. 1) were most clearly responsible for the compositional differences, preferring high complexity habitat. Affinities between general functional groupings of pitfall‐trapped beetles and habitat variables were not clear at a low taxonomic resolution (family level). The composition and species richness of flight‐intercept‐trapped beetles were similar in high and low complexity sites. Our study demonstrates that discrete responses of the various functional groups of beetles are strongly associated with their feeding habits, indicated by differing habitat components from within overall composite habitat complexity measures. Although habitat preferences by beetle species may often reflect their foraging habits, clarification of the causal mechanisms underpinning the relationships between habitat complexity and beetles are critical for the development of general principles linking habitat, functional roles and diversity.     

Do spider diversity and assemblages change in different contiguous habitats? A case study in the protected habitats of the humid chaco ecoregion, northeast Argentina

Spiders are a megadiverse group that can be useful indicators of the overall species richness and health of biotic communities. The spider diversity in subtropical forests of the Neotropical region are not yet well known, especially in Argentinean subtropical forests where systematic fieldwork has not been done until recently. The Great Chaco is very important as the unique dry subtropical forest of the earth, but it is suffering increasing degradation by the advance of agriculture. Spider communities have been shown to be more directly influenced by vegetation architecture than vegetation species composition. In this study, we aim to assess whether spider diversity and assemblages change in adjacent habitats with different types of vegetation. We compare the diversity and spider assemblages in two different contiguous protected habitats (hygrophilous woodland and savannah parkland) of the Mburucuyá National Park, (Humid Chaco ecoregion). Seasonal samples were obtained using three types of sampling methods: pitfall trapping, beating, and manual litter extraction. The spider assemblages were different in the studied areas, and the abundance, diversity, evenness, and species richness were higher in the hygrophilous woodland than the savannah parkland. These differences in spider diversity and assemblages indicate that both types of habitats are important if the biodiversity is to be conserved in the Chaco ecoregion, where different types of habitat are shown as a patchy distribution.     

Demographic consequences of prey availability and diet of Red-billed Choughs Pyrrhocorax pyrrhocorax

Capsule Variation in prey availability appears to influence Chough fledging success and juvenile survival.

Aims To determine seasonal and annual variations in Chough prey and how these influence fledging success and juvenile survival.

Methods Chough faeces (n = 437, 3905 invertebrates) were collected year-round and analysed to determine diet composition. Seasonal and annual variation in prey abundance in Chough foraging habitat was assessed using pitfall trapping (n = 747, 27 124 invertebrates) between 1996 and 2003. Fledging success was estimated for a population of 12 breeding pairs; juvenile survival was estimated by year-round resighting (n < 2500) of individual birds that were colour-ringed as nestlings.

Results From April to June a high biomass of prey and a great variety of species were observed. Between July and October, both biomass and species diversity were reduced; prey consisted mainly of ants and a few beetle species. From November to March, biomass availability was intermediate, with the diet consisting mostly of Tipulidae larvae, plants and dung beetles. Chough juvenile monthly survival was low in August, November and December. Annual variation in fledging success was correlated with prey biomass availability in May.

Conclusion Seasonal variation in the availability of prey species and their biomass influences Chough demography.     

Effects of habitat complexity on ant assemblages

We investigated responses of ant communities to habitat complexity, with the aim of assessing complexity as a useful surrogate for ant species diversity. We used pitfall traps to sample ants at twenty-eight sites, fourteen each of low and high habitat complexity, spread over ca 12 km in Sydney sandstone ridge-top woodland in Australia. Ant species richness was higher in low complexity areas, and negatively associated with ground herb cover, tree canopy cover, soil moisture and leaf litter. Ant community composition was affected by habitat complexity, with morphospecies from the genera Monomorium, Rhytidoponera and Meranoplus being the most significant contributors to compositional differences. Functional group responses to anthropogenic disturbance may be facilitated by local changes in habitat complexity. Habitat complexity, measured as a function of differences in multiple strata in forests, may be of great worth as a surrogate for the diversity of a range of arthropod groups including ants.     

Simple predictions of instream habitat model outputs for fish habitat guilds in large streams

1. In the context of a generalised modification of hydraulic conditions in medium to large streams, modelling the impacts of stream regulation on fish communities in multiple streams is an important challenge for basic and applied freshwater ecology. Conventional instream habitat models such as PHABSIM link a hydraulic model with preference curves for various species to estimate habitat value changes with discharge in stream reaches. Despite world‐wide applications, they have been scarcely used in multiple sites with multiple species. 2. We assigned 21 size classes of European fish species to four habitat guilds (cluster analysis grouping size classes with comparable microhabitat preference curves). Then, we ran a conventional instream habitat model on 28 French stream reaches belonging to the `barbel zone', to estimate habitat values versus discharge curves for the 21 size classes. We summarised the outputs as mean habitat values for guilds, and tested if they were predictable from average characteristics of reaches (discharge, depth, width, particle size). 3. As was obtained elsewhere for populations, habitat values for guilds were strongly related to average, dimensionless characteristics of reaches. The Reynolds number of reaches, equivalent to a discharge per width unit, reflected most of the discharge‐dependent changes in habitat values (within reaches). In particular, habitat values of species preferring bank (respectively midstream) microhabitats decreased (respectively increased) with increasing Reynolds number. The Froude number at median discharge was the major predictor of reach‐dependent but discharge‐independent variations in habitat values. Habitat values of species preferring riffle versus pool or bank microhabitats were higher in reaches with high Froude numbers. These relationships were consistent with existing knowledge on the different species. 4. Such results suggest that the input variables required to estimate habitat values for fish communities can be greatly simplified, as illustrated by a general estimation of the sensitivity of species preferring midstream habitats to discharge changes in any reach. Cost‐efficient alternatives to conventional instream habitat models should facilitate their validation in multiple sites, a point that remains critical in instream habitat modelling of fish communities.     

Influence of tree species and soil properties on ground beetle (Coleoptera: Carabidae) communities

Although Carabidae is among the best-studied families of beetles in Europe from the faunistic point of view, there is still a lack of available information on the ecological requirements of the particular carabid species. The habitat preferences that determine the distribution of species are largely influenced by habitat structure and microclimate. In addition to other factors, these habitat parameters are influenced by the nature of the vegetation. Therefore, our study investigated the influence of tree species on carabid beetle communities. We conducted the research at 9 stands in the Borová Hora Arboretum (Zvolen, Central Slovakia). Each studied site represents a monoculture of one of nine tree species. At each site, some soil and leaf litter attributes (pH, conductivity, and content of H, C, N and P) were evaluated. Ground beetles were collected by pitfall trapping during the vegetation periods in 2008–2011. In total, 3012 individuals of 29 species were obtained. Significant differences in the total dynamic activity and species richness of the carabid beetle communities among the compared forest stands were revealed. The results of the research confirmed statistically significant relationships among 1) the soil conductivity and both the richness and Shannon diversity of the ground beetle communities, 2) the litter and soil N content and richness, the Shannon diversity and the species composition of the ground beetle communities. The Shannon diversity and richness were negatively related to the soil conductivity and positively related with the N content. Our research showed that dominant tree species indirectly influence diversity and composition of carabid communities via the soil properties.     

Tree diversity drives abundance and spatiotemporal β‐diversity of true bugs (Heteroptera)

1. Spatiotemporal patterns of canopy true bug diversity in forests of different tree species diversity have not yet been disentangled, although plant diversity has been shown to strongly impact the diversity and distribution of many insect communities. 2. Here we compare species richness of canopy true bugs across a tree diversity gradient ranging from simple beech to mixed forest stands. We analyse changes in community composition by additive partitioning of species diversity, for communities on various tree species, as well as for communities dwelling on beech alone. 3. Total species richness (γ‐diversity) and α‐diversity, and abundance of true bugs increased across the tree diversity gradient, while diversity changes were mediated by increased true bug abundance in the highly diverse forest stands. The same pattern was found for γ‐diversity in most functional guilds (e.g. forest specialists, herbivores, predators). Temporal and even more, spatial turnover (β‐diversity) among trees was closely related to tree diversity and accounted for ～90% of total γ‐diversity. 4. Results for beech alone were similar, but species turnover could not be related to the tree diversity gradient, and monthly turnover was higher compared to turnover among trees. 5. Our findings support the hypothesis that with increasing tree diversity and thereby increasing habitat heterogeneity, enhanced resource availability supports a greater number of individuals and species of true bugs. Tree species identity and the dissimilarity of true bug communities from tree to tree determine community patterns. 6. In conclusion, understanding diversity and distribution of insect communities in deciduous forests needs a perspective on patterns of spatiotemporal turnover. Heterogeneity among sites, tree species, as well as tree individuals contributed greatly to overall bug diversity.     

Spatiotemporal food web dynamics along a desert riparian–upland transition

Increased awareness of spatiotemporal variation in species interactions has motivated the study of temporally-resolved food web dynamics at the landscape level. Empiricists have focused attention on cross-habitat flows of materials, nutrients, and prey, largely ignoring the movement of predators between habitats that differ in productivity (and how predators integrate pulses in resource availability over time). We set out to study seasonal variation in food web interactions between mammalian carnivores and their rodent prey along a riparian–upland gradient in semi-arid southeastern Arizona which features both spatial and temporal heterogeneity in resource availability. Specifically, we tested whether mammalian carnivores spill over from productive, near-river habitats into adjacent, desert-scrub habitats; and if they do, to document the effects of this spillover on rodent communities. Furthermore, we examined seasonal variation in top-down effects by measuring changes in carnivore diet and distribution patterns and rodent populations over time. The results indicate that carnivores track seasonally-abundant resources across the landscape, varying both their diet and movement patterns. In turn, desert-scrub rodent population dynamics track seasonal shifts in carnivore habitat use but not resource availability, suggesting that predation plays a role in structuring rodent communities along the San Pedro River. Further evidence comes from data on rodent community composition, which differs between desert-scrub habitats near and far from the river, despite similarities in resource availability. Our data also suggest that seasonal omnivory helps predators survive lean times, increasing their effects on prey populations. Taken together, these results underscore the importance of spatiotemporal variation in species interactions, highlighting the complexity of natural systems and the need for further detailed studies of food web dynamics.     

Carabid community succession during the forestry cycle in conifer plantations

Pitfalls and soil heat extraction were used to sample Carabidae in recently felled (1 –4 yr from felling), young (5–22 yr after planting), and old plantations (42–63 yr after planting) of Picea sitchensis in northern England Pitfall catches were useful in recording swift-running species, present at low densities, and a DCA ordination based on the pitfall catches indicated major distribution changes during the plantation cycle Soil extractions allowed estimation of minimum densities of the abundant species and indicated that both diversity and density were highest m young plantations and lowest in the old plantations The flightless Trechus obtusus (3 8 mm) was the most abundant species m the soil samples and its population density did not differ significantly between the three stages in the forestry cycle The winged, highly surface-active species that were early colonisers of clear felled sites and formed a high proportion of the pitfall catch were present at low densities only The peak in carabid density and diversity coincided with the stage at which the ground flora was most diverse (in both species and structure) and the densities of other soil surface macro-invertebrates were highest The high carabid density in the young plantations coincided with a the presence of a relatively high proportion of larger carabids (> 59 mm) and it is suggested that the increased carabid biomass m^-2could reflect increased prey availability The density of winged individuals was higher in the soil samples of carabids from the young plantations than from the clear felled areas and this is attributed to a build up over time of colonising species, adapted to exploiting the short-lived open habitat stage Species that overwinter m the adult stage were more abundant in the young than in the old plantations and it is suggested that these species are favoured by the relatively high summer daytime temperatures of the open habitat The interpretation of the successional sequence in the Carabidae was enhanced by the use of heat extraction sampling in conjunction with pitfall catches     

Subalpine vegetation pattern three decades after stand‐replacing fire: effects of landscape context and topography on plant community composition,tree regeneration,and diversity

Objective: Our purpose was to characterize vegetation compositional patterns, tree regeneration, and plant diversity, and their relationships to landscape context, topography, and light availability across the margins of four stand‐replacing subalpine burns. Location: Four 1977 to 1978 burns east of the Continental Divide in Colorado: the Ouzel burn, a burn near Kenosha Pass, the Badger Mountain burn, and the Maes Creek burn. Methods: Vegetation and environmental factors were sampled in 200 0.01‐ha plots on transects crossing burn edges, and stratified by elevation. We utilized dissimilarity indices, mixed‐effects models, and randomization tests to assess relationships between vegetation and environment. Results: Three decades after wildfire, plant communities exhibited pronounced compositional shifts across burn edges. Tree regeneration decreased with increasing elevation and distance into burn interiors; concomitant increases in forbs and graminoids were linked to greater light availability. Richness was roughly doubled in high‐severity burn interiors due to the persistence of a suite of native species occurring primarily in this habitat. Richness rose with distance into burns, but declined with increasing elevation. Only three of 188 plant species were non‐native; these were widespread, naturalized species that comprised <1% total cover. Conclusions: These subalpine wildfires generated considerable, persistent increases in plant species richness at local and landscape scales, and a diversity of plant communities. The findings suggest that fire suppression in such systems must lead to reduced diversity. Concerns about post‐fire invasion by exotic plants appear unwarranted in high‐elevation wilderness settings.     

退耕还林对地面蜘蛛种群的影响

为探究退耕还林对地面蜘蛛群落的影响,分别于2014和2015年3月—9月期间,采用陷阱法对北京市顺义区南彩镇退耕还林已2a和3a的林带及相邻原非作物生境林带中地面蜘蛛的活动密度进行了系统调查和分析。结果,2014年在调查区域共采集地面蜘蛛1465头,隶属8科18属32种,2015年共收集地面蜘蛛2186头,隶属10科25属45种;退耕还林以后蜘蛛类群发生改变,地面蜘蛛丰富度增加,而部分农田蜘蛛消失;2014年原非作物生境林带中地面蜘蛛的多样性指数、丰富度都极显著高于退耕还林林带(P0.01),均匀性指数不存在显著差异,2015年两个林带的多样性指数仍存在极显著差异(P0.01),丰富度存在显著差异(P0.05),均匀性指数之间不存在显著差异;退耕还林以后第3年新建林带和原非作物生境林带中地面蜘蛛的活动密度随时间的变化趋势相近。研究结果表明:顺义地区退耕还林工程实施3a以后,新建人工林仍处于生境重建阶段,未完全达到稳定状态,退耕还林以后第3年新建林带中地面蜘蛛群落的稳定性高于第2年。退耕还林通过改变原有农田生态系统的植被种类、结构、人为管理等方式,使原有农田生态系统发生了根本性的改变。持续研究退耕还林对地面蜘蛛的类群变化的影响对生境受到干扰后生物群落变化过程的认识有理论意义。     

Increasing isolation reduces predator:prey species richness ratios in aquatic food webs

The number of species that live in a habitat typically declines as that habitat becomes more isolated. However, the influence of habitat isolation on patterns of food web structure, in particular the ratio of predator to prey species richness, is less well understood. We placed aquatic mesocosms at varying distances from ponds that acted as sources of potential colonists; then we examined how isolation affected the ratio of predator:prey species richness in the communities that assembled. In the final sampling, a total of 21 species (12 prey and 9 predators) of insects, crustaceans, and amphibians had colonized the mesocosms. We found that total species richness, as well as the richness of predators and prey, declined with increasing isolation. However, predator richness declined more rapidly than prey richness with increasing isolation, which lead to decreasing predator:prey ratios. This result conflicts with prior demonstrations of invariant predator:prey ratios in freshwater communities.     

Zebra mussels affect benthic predator foraging success and habitat choice on soft sediments

The introduction of zebra mussels (Dreissena spp.) to North America has resulted in dramatic changes to the complexity of benthic habitats. Changes in habitat complexity may have profound effects on predator-prey interactions in aquatic communities. Increased habitat complexity may affect prey and predator dynamics by reducing encounter rates and foraging success. Zebra mussels form thick contiguous colonies on both hard and soft substrates. While the colonization of substrata by zebra mussels has generally resulted in an increase in both the abundance and diversity of benthic invertebrate communities, it is not well known how these changes affect the foraging efficiencies of predators that prey on benthic invertebrates. We examined the effect of zebra mussels on the foraging success of four benthic predators with diverse prey-detection modalities that commonly forage in soft substrates: slimy sculpin (Cottus cognatus), brown bullhead (Ameirus nebulosus), log perch (Percina caprodes), and crayfish (Orconectes propinquus). We conducted laboratory experiments to assess the impact of zebra mussels on the foraging success of predators using a variety of prey species. We also examined habitat use by each predator over different time periods. Zebra mussel colonization of soft sediments significantly reduced the foraging efficiencies of all predators. However, the effect was dependent upon prey type. All four predators spent more time in zebra mussel habitat than in either gravel or bare sand. The overall effect of zebra mussels on benthic-feeding fishes is likely to involve a trade-off between the advantages of increased density of some prey types balanced against the reduction in foraging success resulting from potential refugia offered in the complex habitat created by zebra mussels.