Spatial scale, abundance and the species-energy relationship in British birds

1. The spatial scale of analysis may influence the nature, strength and underlying drivers of macroecological patterns, one of the most frequently discussed of which is the relationship between species richness and environmental energy availability. 2. It has been suggested that species-energy relationships are hump-shaped at fine spatial grains and consistently positive at larger regional grains. The exact nature of this scale dependency is, however, the subject of much debate as relatively few studies have investigated species-energy relationships for the same assemblage across a range of spatial grains. Here, we contrast species-energy relationships for the British breeding avifauna at spatial grains of 1 km x 1 km, 2 km x 2 km and 10 km x 10 km plots, while maintaining a constant spatial extent. 3. Analyses were principally conducted using data on observed species richness. While survey work may fail to detect some species, observed species richness and that estimated using nonparametric techniques were strongly positively correlated with each other, and thus exhibit very similar spatial patterns. Moreover, the forms of species-energy relationships using observed and estimated species richness were statistically indistinguishable from each other. 4. Positive decelerating species-energy relationships arise at all three spatial grains. There is little evidence that the explanatory power of these relationships varies with spatial scale. However, ratios of regional (large-scale) to local (small-scale) species richness decrease with increasing energy availability, indicating that local richness responds to energy with a steeper gradient than does regional richness. Local assemblages thus sample a greater proportion of regional richness at higher energy levels, suggesting that spatial turnover of species richness is lower in high-energy regions. Similarly, a crude measure of temporal turnover, the ratio of cumulative species richness over a 4-year period to species richness in a single year, is lower in high-energy regions. These negative relationships between turnover and energy appear to be causal as both total and mean occupancy per species increases with energy. 5. While total density in 1 km x 1 km plots correlates positively with energy availability, such relationships are very weak for mean density per species. This suggests that the observed association between total abundance and species richness may not be mediated by population extinction rates, as predicted by the more individuals hypothesis. 6. The sampling mechanism suggests that species-energy relationships arise as high-energy areas support a greater number of individuals, and that random allocation of these individuals to local areas from a regional assemblage will generate species-energy relationships. While randomized local species-energy relationships are linear and positive, predicted richness is consistently greater than that observed. The mismatch between the observed and randomized species-energy relationships probably arises as a consequence of the aggregated nature of species distributions. The sampling mechanism, together with species spatial aggregation driven by limited habitat availability, may thus explain the species-energy relationship observed at this spatial scale.     

Scale dependence of species-energy relationships: evidence from fishes in thousands of lakes

Variation in the shape of relationships between species richness and different measures of energy may be linked to variation in the spatial scale on which such relationships are measured. We examine scale dependence in the relationship between potential evapotranspiration and the species richness of fishes in 7,885 postglacial lakes. The strength of this relationship is weak across lake communities but strong and positive across groups of lakes or regions. In addition, the strength and slope of this relationship increase significantly as the regional scale of analysis is increased. We interpret the observed patterns in terms of a simple model whereby energy influences the linear character of the species-energy relationship through its influence on spatial turnover in the species composition (beta diversity). Our results suggest that if energy is strongly tied to patterns of site occupancy or abundance, the parameters of species-energy relationships will depend, to a considerable extent, on the scale of measurement. Furthermore, the ability of high-energy regions to accommodate relatively large numbers of rare or infrequent species may underlie any general tendency for the strength or shape of species-energy relationships to change with scale.     

Evidence that niche specialization explains species-energy relationships in lake fish communities

1. Interspecific niche differences have long been identified as a major explanation for the occurrence of species-rich communities. However, much fieldwork studying variation in local species richness has focused upon physical habitat attributes or regional factors, such as the size of the regional species pool. 2. We applied indices of functional diversity and niche overlap to data on the species niche to examine the importance of interspecific niche differentiation for species richness in French lake fish communities. We combined this information with environmental data to test generalizations of the physiological tolerance and niche specialization hypotheses for species-energy relationships. 3. We found evidence for a largely non-saturating relationship (relative to random expectation) between species richness and functional evenness (evenness of spacing between species in niche space), while functional richness (volume of niche space occupied) peaked at moderate levels of species richness and niche overlap showed an initial decrease followed by saturation. This suggests that increased niche specialization may have allowed species to coexist in the most species-rich communities. 4. We tested for evidence that increased temperature, local habitat area, local habitat diversity and immigration affected species richness via increased niche specialization. Temperature explained by far the largest amount of variation in species richness, functional diversity and niche overlap. These results, combined with the largely non-saturating species richness-functional evenness relationship, suggest that increased temperature may have permitted increased species richness by allowing increased niche specialization. 5. These results emphasize the importance of niche differences for species coexistence in species-rich communities, and indicate that the conservation of functional diversity may be vital for the maintenance of species diversity in biological communities. Our approach may be applied readily to many types of community, and at any scale, thus providing a flexible means of testing niche-based hypotheses for species richness gradients.     

Dissecting the species-energy relationship

Environmental energy availability can explain much of the spatial variation in species richness. Such species-energy relationships encompass a diverse range of forms, and there is intense debate concerning which of these predominate, and the factors promoting this diversity. Despite this there has been relatively little investigation of whether the form, and relative strength, of species-energy relationships varies with (i) the currency of energy availability that is used, and (ii) the ecological characteristics of the constituent species. Such investigations can, however, shed light on the causal mechanisms underlying species-energy relationships. We illustrate this using the British breeding avifauna. The strength of the species-energy relationship is dependent on the energy metric used, with species richness being more closely correlated with temperature than the Normalized Difference Vegetation Index, which is a strong correlate of net primary productivity. We find little evidence, however, for the thermoregulatory load hypothesis that high temperatures enable individuals to invest in growth and reproduction, rather than thermoregulation, increasing population sizes that buffer species from extinction. High levels of productive energy may also elevate population size, which is related to extinction risk by a negative decelerating function. Therefore, the rarest species should exhibit the strongest species-energy relationship. We find evidence to the contrary, together with little support for suggestions that high-energy availability elevates species richness by increasing the numbers of specialists or predators.     

Overexploiting marine ecosystem engineers: potential consequences for biodiversity

Overfishing is a major environmental problem in the oceans. In addition to the direct loss of the exploited species, the very act of fishing, particularly with mobile bottom gear, destroys habitat and ultimately results in the loss of biodiversity. Furthermore, overfishing can create trophic cascades in marine communities that cause similar declines in species richness. These effects are compounded by indirect effects on habitat that occur through removal of ecological or ecosystem engineers. Mass removal of species that restructure the architecture of habitat and thus increase its complexity or influence the biogeochemistry of sediments could have devastating effects on local biodiversity and important water–sediment processes. The possible overexploitation of engineering species requires more attention because the consequences extend beyond their own decline to affect the rest of the ecosystem. This is particularly problematic in the deep ocean, where oil and gas exploration and fishing pressure are likely to increase.     

Relative importance of the land‐use composition and intensity for the bird community composition in anthropogenic landscapes

Humans are changing the biosphere by exerting pressure on land via different land uses with variable intensities. Quantifying the relative importance of the land‐use composition and intensity for communities may provide valuable insights for understanding community dynamics in human‐dominated landscapes. Here, we evaluate the relative importance of the land‐use composition versus land‐use intensity on the bird community structure in the highly human‐dominated region surrounding Paris, France. The land‐use composition was calculated from a land cover map, whereas the land‐use intensity (reverse intensity) was represented by the primary productivity remaining after human appropriation (NPP_remaining), which was estimated using remote sensing imagery. We used variance partitioning to evaluate the relative importance of the land‐use composition versus intensity for explaining bird community species richness, total abundance, trophic levels, and habitat specialization in urban, farmland, and woodland habitats. The land‐use composition and intensity affected specialization and richness more than trophic levels and abundance. The importance of the land‐use intensity was slightly higher than that of the composition for richness, specialization, and trophic levels in farmland and urban areas, while the land‐use composition was a stronger predictor of abundance. The intensity contributed more to the community indices in anthropogenic habitats (farmland and urban areas) than to those in woodlands. Richness, trophic levels, and specialization in woodlands tended to increase with the NPP_remaining value. The heterogeneity of land uses and intensity levels in the landscape consistently promoted species richness but reduced habitat specialization and trophic levels. This study demonstrates the complementarity of NPP_remaining to the land‐use composition for understanding community structure in anthropogenic landscapes. Our results show, for the first time, that the productivity remaining after human appropriation is a determinant driver of animal community patterns, independent of the type of land use.     

Local-scale species-energy relationships in fish assemblages of some forested streams of the Bolivian Amazon

Productivity (trophic energy) is one of the most important factors promoting variation in species richness. A variety of species-energy relationships have been reported, including monotonically positive, monotonically negative, or unimodal (i.e. hump-shaped). The exact form of the relationship seems to depend, among other things, on the spatial scale involved. However, the mechanisms behind these patterns are still largely unresolved, although many hypotheses have been suggested. Here we report a case of local-scale positive species-energy relationship. Using 14 local fish assemblages in tropical forested headwater streams (Bolivia), and after controlling for major local abiotic factors usually acting on assemblage richness and structure, we show that rising energy availability through leaf litter decomposition rates allows trophically specialized species to maintain viable populations and thereby to increase assemblage species richness. By deriving predictions from three popular mechanistic explanations, i.e. the 'increased population size', the 'consumer pressure', and the 'specialization' hypotheses, our data provide only equivocal support for the latter.     

Estimated plant richness pattern across northwest South America provides similar support for the species-energy and spatial heterogeneity hypotheses

Explaining geographic variation in plant species richness at broad spatial scales has long been a major challenge. Many hypotheses have been proposed during the last 200 yr, but recent work has focused on a few major alternatives. Among these, two hypotheses contend that plant species richness reflects 1) variation in energy and water availability among sampling units (the species-energy hypothesis) and 2) habitat and topographic heterogeneity within sampling units (the spatial heterogeneity hypothesis). We used a large botanical database and regression models to simultaneously confront the predictions from both hypotheses against an estimate of vascular plant richness across northwest South America. This estimate provided similar support for both hypotheses, a result that may be seen as contrasting with the notion that variation in energy and water availability among sampling units is the main determinant of plant species richness. We discuss potential explanations for this apparent discrepancy. Regression models that incorporated the relative contributions of both hypotheses predicted that the highest plant species richness in northwest South America is found in topographically complex areas. In contrast to several of the most recent mapping efforts, lowland Amazonia was predicted to be a plant richness trough in the study region. We suggest that diverging portrayals of plant richness across northwest South America result from differences in estimates of the relative importance of the species-energy and the spatial heterogeneity hypotheses.     

Relative contribution of abundant and rare species to species-energy relationships

A major goal of ecology is to understand spatial variation in species richness. The latter is markedly influenced by energy availability and appears to be influenced more by common species than rare ones; species-energy relationships should thus be stronger for common species. Species-energy relationships may arise because high-energy areas support more individuals, and these larger populations may buffer species from extinction. As extinction risk is a negative decelerating function of population size, this more-individuals hypothesis (MIH) predicts that rare species should respond more strongly to energy. We investigate these opposing predictions using British breeding bird data and find that, contrary to the MIH, common species contribute more to species-energy relationships than rare ones.     

Marine reserves indirectly affect fine‐scale habitat associations,but not overall densities,of small benthic fishes

Many large, fishery‐targeted predatory species have attained very high relative densities as a direct result of protection by no‐take marine reserves. Indirect effects, via interactions with targeted species, may also occur for species that are not themselves targeted by fishing. In some temperate rocky reef ecosystems, indirect effects have caused profound changes in community structure, notably the restoration of predator–urchin–macroalgae trophic cascades. Yet, indirect effects on small benthic reef fishes remain poorly understood, perhaps because of behavioral associations with complex, refuge‐providing habitats. Few, if any, studies have evaluated any potential effects of marine reserves on habitat associations in small benthic fishes. We surveyed densities of small benthic fishes, including some endemic species of triplefin (Tripterygiidae), along with fine‐scale habitat features in kelp forests on rocky reefs in and around multiple marine reserves in northern New Zealand over 3 years. Bayesian generalized linear mixed models were used to evaluate evidence for (1) main effects of marine reserve protection, (2) associations with habitat gradients, including complexity, and (3) differences in habitat associations inside versus outside reserves. No evidence of overall main effects of marine reserves on species richness or densities of fishes was found. Both richness and densities showed strong associations with gradients in habitat features, particularly habitat complexity. In addition, some species exhibited reserve‐by‐habitat interactions, having different associations with habitat gradients inside versus outside marine reserves. Two species (Ruanoho whero and Forsterygion flavonigrum) showed stronger positive associations with habitat complexity inside reserves. These results are consistent with the presence of a behavioral risk effect, whereby prey fishes are more strongly attracted to habitats that provide refuge from predation in areas where predators are more abundant. This work highlights the importance of habitat structure and the potential for fishing to affect behavioral interactions and the interspecific dynamic attributes of community structure beyond simple predator–prey consumption and archetypal trophic cascades.     

Aquatic invertebrate communities in tank bromeliads: how well do classic ecological patterns apply?

Tank bromeliads (Bromeliaceae) often occur in high densities in the Neotropics and represent a key freshwater habitat in montane forests, housing quite complex invertebrate communities. We tested the extent to which there are species richness–altitude, richness–environment, richness–size, richness–habitat complexity and richness–isolation relationships for the aquatic invertebrate communities from 157 bromeliads in Cusuco National Park, Honduras. We found that invertebrate species richness and abundance correlated most strongly, and positively, with habitat size, which accounted for about a third of the variance in both. Apart from bromeliad size (equivalent of the species–area relationship), we found remarkably little evidence of classic biogeographic and ecological relationships with species richness in this system. Community composition correlated with altitude, bromeliad size and position, though less than 20% of the variation was accounted for by the tested variables. The turnover component of dissimilarity between the communities correlated with altitude, while the nestedness-resultant component was related to bromeliad size. The unexplained variance could reflect a large stochastic component in the system, associated with the ephemerality of the habitat patches (both the plants themselves and the fluctuations in their water content) and stochasticity due to the dispersal dynamics in the system. We conclude that there is a small contribution of classic biogeographic factors to the diversity and community composition of aquatic invertebrates communities in bromeliads. This may be due to the highly dynamic nature of this system, with small patch sizes and high emigration rates. The patterns may mostly be driven by factors affecting colonisation success.     

The species-area-energy relationship

Area and available energy are major determinants of species richness. Although scale dependency of the relationship between energy availability and species richness (the species-energy relationship) has been documented, the exact relationship between the species-area and the species-energy relationship has not been studied explicitly. Here we show, using two extensive data sets on avian distributions in different biogeographic regions, that there is a negative interaction between energy availability and area in their effect on species richness. The slope of the species-area relationship is lower in areas with higher levels of available energy, and the slope of the species-energy relationship is lower for larger areas. This three-dimensional species-area-energy relationship can be understood in terms of probabilistic processes affecting the proportions of sites occupied by individual species. According to this theory, high environmental energy elevates species' occupancies, which depress the slope of the species-area curve.     

Eulittoral macroinvertebrate communities of lowland lakes: discrimination among trophic states

1. Nutrient inputs from urban and agricultural land use often result in shifts in species composition of pelagic and profundal invertebrate communities. Here, we test if nutrient enrichment affects the composition of eulittoral macroinvertebrate communities, and, if so, if macroinvertebrate communities of five different habitat types reflect differences in trophic state. 2. Macroinvertebrate community composition of 36 lakes was significantly correlated with total phosphorus (TP) concentration, the proportion of coarse woody debris (CWD) and root habitats and the proportion of grassland. 3. However, macroinvertebrate communities of five major habitat types from eight lakes were more dissimilar among habitats than among trophic states. Community composition of reed and stone habitats was significantly correlated with wind exposure but not TP concentration, while macroinvertebrate composition of sand habitats was related to TP concentration and coarse sediments. In CWD and root habitats, both TP concentration and a predominance of invasive species covaried, which made it difficult to relate the observed compositional differences to either trophic state or to the effects of competition between native and invasive species. 4. Trophic state influenced the composition of eulittoral macroinvertebrate communities but to a lesser extent than has been previously reported for profundal habitats. Moreover, the effects of trophic state were nested within habitat type and were partially superseded by biotic interactions and small‐scaled habitat complexity. Although eulittoral macroinvertebrate communities were not strong indicators of the trophic state of lowland lakes, they may be used to assess other anthropogenic impacts on lakeshores.     

Relative Importance of Coral Cover,Habitat Complexity and Diversity in Determining the Structure of Reef Fish Communities

The structure of coral reef habitat has a pronounced influence on the diversity, composition and abundance of reef-associated fishes. However, the particular features of the habitat that are most critical are not always known. Coral habitats can vary in many characteristics, notably live coral cover, topographic complexity and coral diversity, but the relative effects of these habitat characteristics are often not distinguished. Here, we investigate the strength of the relationships between these habitat features and local fish diversity, abundance and community structure in the lagoon of Lizard Island, Great Barrier Reef. In a spatial comparison using sixty-six 2m² quadrats, fish species richness, total abundance and community structure were examined in relation to a wide range of habitat variables, including topographic complexity, habitat diversity, coral diversity, coral species richness, hard coral cover, branching coral cover and the cover of corymbose corals. Fish species richness and total abundance were strongly associated with coral species richness and cover, but only weakly associated with topographic complexity. Regression tree analysis showed that coral species richness accounted for most of the variation in fish species richness (63.6%), while hard coral cover explained more variation in total fish abundance (17.4%), than any other variable. In contrast, topographic complexity accounted for little spatial variation in reef fish assemblages. In degrading coral reef environments, the potential effects of loss of coral cover and topographic complexity are often emphasized, but these findings suggest that reduced coral biodiversity may ultimately have an equal, or greater, impact on reef-associated fish communities.     

Trophic level modulates carabid beetle responses to habitat and landscape structure: a pan‐European study

1. Anthropogenic pressures have produced heterogeneous landscapes expected to influence diversity differently across trophic levels and spatial scales. 2. We tested how activity density and species richness of carabid trophic groups responded to local habitat and landscape structure (forest percentage cover and habitat richness) in 48 landscape parcels (1 km²) across eight European countries. 3. Local habitat affected activity density, but not species richness, of both trophic groups. Activity densities were greater in rotational cropping compared with other habitats; phytophage densities were also greater in grassland than forest habitats. 4. Controlling for country and habitat effects, we found general trophic group responses to landscape structure. Activity densities of phytophages were positively correlated, and zoophages uncorrelated, with increasing habitat richness. This differential functional group response to landscape structure was consistent across Europe, indicated by a lack of a country × habitat richness interaction. Species richness was unaffected by landscape structure. 5. Phytophage sensitivity to landscape structure may arise from relative dependency on seed from ruderal plants. This trophic adaptation, rare in Carabidae, leads to lower phytophage numbers, increasing vulnerability to demographic and stochastic processes that the greater abundance, species richness, and broader diet of the zoophage group may insure against.     

Litter identity mediates predator impacts on the functioning of an aquatic detritus-based food web

During past decades, several mechanisms such as resource quality and habitat complexity have been proposed to explain variations in the strength of trophic cascades across ecosystems. In detritus-based headwater streams, litter accumulations constitute both a habitat and a resource for detritivorous macroinvertebrates. Because litter edibility (which promotes trophic cascades) is usually inversely correlated with its structural complexity (which weakens trophic cascades), there is a great scope for stronger trophic cascades in litter accumulations that are dominated by easily degradable litter species. However, it remains unclear how mixing contrasting litter species (conferring both habitat complexity and high quality resource) may influence top–down controls on communities and processes. In enclosures exposed in a second-order stream, we manipulated litter species composition by using two contrasting litter (alder and oak), and the presence–absence of a macroinvertebrate predator (Cordulegaster boltonii larvae), enabling it to effectively exert predation pressure, or not, on detritivores (consumptive versus non-consumptive predation effects). Leaf mass loss, detritivore biomass and community structure were mostly controlled independently by litter identity and mixing and by predator consumption. However, the strength of predator control was mediated by litter quality (stronger on alder), and to a lesser extent by litter mixing (weaker on mixed litter). Refractory litter such as oak leaves may contribute to the structural complexity of the habitat for stream macroinvertebrates, allowing the maintenance of detritivore communities even when strong predation pressure occurs. We suggest that considering the interaction between top–down and bottom–up factors is important when investigating their influence on natural communities and ecosystem processes in detritus-based ecosystems.     

Landscape complexity differentially benefits generalized fourth,over specialized third,trophic level natural enemies

The differential loss of higher trophic levels in the face of natural habitat loss can result in the disruption of important trophic interactions, such as biological control. Natural enemies of herbivorous pests in cropping systems often benefit from the presence of natural habitats in surrounding landscapes, as they provide key resources such as alternative hosts. However, any benefits from a biological control perspective may be dampened if this also enhances enemies at the fourth trophic level. Remarkably, studies of the influence of landscape structure on diversity and interactions of fourth trophic‐level natural enemies are largely lacking. We carried out a large‐scale sampling study to investigate the effects of landscape complexity (i.e. the proportion of non‐crop habitat in the landscapes surrounding focal study areas) on the parasitoid communities of aphids in wheat and on an abundant extra‐field plant, stinging nettle. Primary parasitoid communities (3rd trophic level) attacking the cereal aphid, Sitobion avenae, had little overlap with the communities attacking the nettle aphid, Microlophium carnosum, while secondary parasitoids (4th trophic level) showed high levels of species overlap across these two aphids (25 vs 73% shared species respectively), resulting in significantly higher linkage density and lower specialization for secondary than primary parasitoid webs. In wheat, parasitoid diversity was not related to landscape complexity for either primary or secondary parasitoids. Rates of primary parasitism were generally low, while secondary parasitism rates were high (37–94%) and increased significantly with increasing landscape complexity, although this pattern was driven by a single secondary parasitoid species. Overall, our results demonstrate that extra‐field habitats and landscape complexity can differentially benefit fourth, over third, trophic level natural enemies, and thereby, could dampen biological control. Our results further suggest that fourth trophic‐level enemies may play an important, yet understudied, role in linking insect population dynamics across habitat types.     

Tree diversity drives diversity of arthropod herbivores,but successional stage mediates detritivores

The high tree diversity of subtropical forests is linked to the biodiversity of other trophic levels. Disentangling the effects of tree species richness and composition, forest age, and stand structure on higher trophic levels in a forest landscape is important for understanding the factors that promote biodiversity and ecosystem functioning. Using a plot network spanning gradients of tree diversity and secondary succession in subtropical forest, we tested the effects of tree community characteristics (species richness and composition) and forest succession (stand age) on arthropod community characteristics (morphotype diversity, abundance and composition) of four arthropod functional groups. We posit that these gradients differentially affect the arthropod functional groups, which mediates the diversity, composition, and abundance of arthropods in subtropical forests. We found that herbivore richness was positively related to tree species richness. Furthermore, the composition of herbivore communities was associated with tree species composition. In contrast, detritivore richness and composition was associated with stand age instead of tree diversity. Predator and pollinator richness and abundance were not strongly related to either gradient, although positive trends with tree species richness were found for predators. The weaker effect of tree diversity on predators suggests a cascading diversity effect from trees to herbivores to predators. Our results suggest that arthropod diversity in a subtropical forest reflects the net outcome of complex interactions among variables associated with tree diversity and stand age. Despite this complexity, there are clear linkages between the overall richness and composition of tree and arthropod communities, in particular herbivores, demonstrating that these trophic levels directly impact each other.     

Ambient and substrate energy influence decomposer diversity differentially across trophic levels

The species-energy hypothesis predicts increasing biodiversity with increasing energy in ecosystems. Proxies for energy availability are often grouped into ambient energy (i.e., solar radiation) and substrate energy (i.e., non-structural carbohydrates or nutritional content). The relative importance of substrate energy is thought to decrease with increasing trophic level from primary consumers to predators, with reciprocal effects of ambient energy. Yet, empirical tests are lacking. We compiled data on 332,557 deadwood-inhabiting beetles of 901 species reared from wood of 49 tree species across Europe. Using host-phylogeny-controlled models, we show that the relative importance of substrate energy versus ambient energy decreases with increasing trophic levels: the diversity of zoophagous and mycetophagous beetles was determined by ambient energy, while non-structural carbohydrate content in woody tissues determined that of xylophagous beetles. Our study thus overall supports the species-energy hypothesis and specifies that the relative importance of ambient temperature increases with increasing trophic level with opposite effects for substrate energy.     

Fragmentation in calcareous grasslands: species specialization matters

Habitat fragmentation resulting from anthropogenic land-use change may negatively affect both biodiversity and ecosystem structure and function. However, susceptibility to fragmentation varies between species and may be influenced by for instance specialization, functional traits and trophic level. We examined how total and specialist species richness, species composition and functional trait composition at two trophic levels (vascular plants and sap-feeding hoppers) vary with habitat fragmentation (patch size and connectivity) in dry calcareous grasslands in southeast Norway. We found that fragmentation affected plant and hopper species composition both totally and of habitat specialists, but with a net species loss only for the specialists, indicating greater susceptibility of specialized species. Reductions in patch size and increasing isolation negatively affected plant specialists with different sets of traits, effectively reducing the number of species with trait combinations suitable to persist in small and isolated patches. Fragmentation influenced trait composition of the total hopper community, but not of habitat specialists. A lesser degree of habitat association could explain why hoppers, despite belonging to a higher tropic level, seemed to be less susceptible to fragmentation than plants. Nonetheless, our study shows that habitat fragmentation affects both species richness, species composition and trait composition of plants and hoppers, indicating that fragmentation leads not only to a loss of species, but also alters dominance hierarchies and the functionality of grassland communities.