Impacts of Land Abandonment on Vegetation: Successional Pathways in European Habitats

Changes in traditional agricultural systems in Europe in recent decades have led to widespread abandonment and colonization of various habitats by shrubs and trees. We combined several vegetation databases to test whether patterns of changes in plant diversity after land abandonment in different habitats followed similar pathways. The impacts of land abandonment and subsequent woody colonization on vegetation composition and plant traits were studied in five semi-natural open habitats and two arable habitats in six regions of Europe. For each habitat, vegetation surveys were carried out in different stages of succession using either permanent or non-permanent plots. Consecutive stages of succession were defined on a physiognomic basis from initial open stages to late woody stages. Changes in vegetation composition, species richness, numbers of species on Red Lists, plant strategy types, Ellenberg indicator values of the vegetation, Grime CSR strategy types and seven ecological traits were assessed for each stage of the successional pathway. Abandonment of agro-pastoral land-use and subsequent woody colonization were associated with changes in floristic composition. Plant richness varied according to the different habitats and stages of succession, but semi-natural habitats differed from arable fields in several ecological traits and vegetation responses. Nevertheless, succession occurred along broadly predictable pathways. Vegetation in abandoned arable fields was characterized by a decreasing importance of R-strategists, annuals, seed plants with overwintering green leaves, insect-pollinated plants with hemi-rosette morphology and plants thriving in nutrient-rich conditions, but an increase in species considered as endangered according to the Red Lists. Conversely, changes in plant traits with succession within the initially-open semi-natural habitats showed an increase in plants thriving in nutrient-rich conditions, stress-tolerant plants and plants with sexual and vegetative reproduction, but a sharp decrease in protected species. In conclusion, our study showed a set of similarities in responses of the vegetation in plant traits after land abandonment, but we also highlighted differences between arable fields and semi-natural habitats, emphasizing the importance of land-use legacy.     

Ecosystem engineers maintain a rare species of butterfly and increase plant diversity

We evaluated whether ecosystem engineers can accomplish two conservation goals simultaneously: (1) indirectly maintain populations of an endangered animal through habitat modification and (2) increase riparian plant diversity. We tested for effects of a prominent ecosystem engineer, the beaver Castor canadensis, on populations of St. Francis' satyr butterfly Neonympha mitchellii francisci and plant species richness and composition. We performed our test by surveying riparian vegetation communities in all stages of beaver‐influenced wetland succession. We found that beavers created wetland habitats that supported plant species not found elsewhere in riparian zones and increased plant species diversity across the landscape by creating a novel combination of patch types. Our results confirmed what others have found about engineering effects on plant diversity, but these results further demonstrated a case where ecosystem engineers indirectly maintain populations of rare animals by modifying the composition and diversity of plant communities within wetlands. Our research demonstrates how an ecosystem engineer can influence habitat availability and composition of plant communities important for an endangered insect, and maintain overall plant species diversity by increasing habitat heterogeneity.     

Secondary Succession and Natural Habitat Restoration in Abandoned Rice Fields of Central Korea

Floristic composition and soil characteristics (moisture, pH, nutrient contents) in abandoned upland rice paddies of different ages were analyzed to clarify the regenerative aspects of succession as a tool for habitat restoration. The study sites represented five seral stages: newly abandoned paddy fields; successional paddy fields abandoned for 3, 7, and 10 years; and a 50‐year‐old Alnus japonica forest. A vegetation sere was apparent in changes of dominant plant species in the order Alopecurus aequalis var. amurensis (annual grass), Aneilema keisak (annual forb), Juncus effusus var. decipiens (rush), Salix koriyanagi (willow), and Alnus japonica (alder) communities. These temporal stages resemble the spatial zonation of vegetation in local riparian floodplain ecosystems, indicating a hydrosere, with soil moisture decreasing over time. Age distributions and life forms of the dominant plant species support a “tolerance” model of secondary succession, in which the established species persist into later successional stages. Persistence of earlier colonizers led to a net cumulative increase in species richness and a more even distribution of species cover with increasing field age. Between 10 and 50 years, vegetation stabilizes as an alder community. Soil moisture content decreased steadily with paddy field age after an initial rise immediately after their abandonment, whereas pools of organic matter, N, P, K, Ca, and Mg, increased with field age. The pace and direction of recovery of native vegetation and natural soil properties in these abandoned rice paddies resembled classic old field succession, a form of secondary succession that often serves as a template for guiding restoration efforts. Active intervention, in particular dismantling artificial levees, could accelerate the recovery process, but natural habitat recovery generally appears sufficiently robust to achieve “passive” restoration of this rare community without intervention.     

Soil seed bank dynamics in alpine wetland succession on the Tibetan Plateau

The primary goal was to address several questions with regard to how soil seed banks change in a successional series. How does the composition of the viable seed bank change, and how does the relationship of the soil seed bank and vegetation change with succession? Can the seed bank be regarded as a potential as a source of seeds for wetland restoration? We collected soil seed bank samples and sampled the vegetation in four different successional stages and used the NMDS (nonmetric multidimensional scaling) to evaluate the relationship of species composition between the seed banks and vegetation. The difference of seed density and species richness in different habitats and soil depths also was compared. Viable seeds of half (37) the species in the early-successional stage were found in all the successional stages. Similarity between seed bank and vegetation increased with succession. Both seed density and species richness in the seed bank increased with successional age and decreased with soil depth. The majority of species from the early-successional stage produced long-lived seeds. Seed density and species richness increased with succession, mainly as a result of increasing seed production, and hypotheses predicting decreasing density of buried seeds and species richness were not confirmed. Seed banks play a minor role in contributing to the regeneration of vegetation, and managers cannot rely on soil-stored seed banks for restoration of wetlands.     

Restore it,and they will come: trap-nesting bee and wasp communities (Hymenoptera: Aculeata) are recovered by restoration of riparian forests

Riparian forests have been greatly affected by anthropogenic actions with formerly continuous riparian forests being slowly converted into small and isolated patches. Riparian forests are extremely important habitats for many groups of insects, including bees and wasps, because they are sources of shelter and food for them and their offspring. There is a growing body of evidence of success in the restoration of riparian forest plant communities; however, little research has been done on the associated invertebrate communities. We test whether restoring plant communities is sufficient for restoring the taxonomic composition of trap-nesting bees and wasps and which functional traits are favored in different sites. We predict that species richness, abundance, and community composition of trap-nesting bees and wasps of riparian sites undergoing restoration will converge on the “target” of a reference site with increasing time, since restoration increases habitat complexity. We also predict that the width of restored patches will also influence the species richness, abundance and community composition of trap-nesting bees and wasps. Bee richness and abundance, and wasp richness, were strongly related to fragment width, but not to age since restoration. Our results indicate that although restored sites are relatively small and scattered in a fragmented landscape, they provide suitable habitat for re-colonization by community assemblages of trap-nesting bees and wasps and the traits selected captured the responses to the habitat restoration. Hence, restored riparian areas can be considered important habitats for invertebrates, thus contributing to an increase in local biodiversity and, possibly, the restoration of some of the ecosystem services they originally provided.     

Diversity and community composition of herbaceous plants in different habitat types in south‐east Cameroon

The composition of herbaceous vegetation was evaluated with the aim of characterizing forests at various ages of stand development. Herb stems were sampled in 250 4‐m² square plots distributed within six habitat types. A total of 36 herb species belonging to 15 families were recorded. Species richness did not significantly differ between habitat types. Most herb species occurred in all habitat types and were therefore generalists. However, a few indicator herb species were detected, and the results roughly suggested that herb species of the families Poaceae and Araceae were indicative of late successional forests; Zingiberaceae are indicative of early successional forests; and Commelinaceae, Costaceae, Cyperaceae and Marantaceae are indicators of flooded habitats. Species diversity and stem density of herbaceous plants did not change with forest succession as a decrease in abundance and frequency of occurrence of pioneer species in late successional forests was counterbalanced by the presence of generalist and late successional species. However, increasing proportions of dwarf stems in late successional forests translated to changes in the vertical structure of herbaceous plant communities. Herbivory pressure by gorillas did not have a notable effect on herbaceous plant community development. This study contributes to the definition of herbaceous ecological indicators of forest succession in different settings.     

Response of medium‐ and large‐sized terrestrial fauna to corridor restoration along the middle Sacramento River

A fundamental challenge in restoration ecology is to understand when species are expected to colonize newly created habitat. Determining this is important for assessing progress toward restoration goals and, more generally, for gaining insight into ecosystem functioning and dynamics. We studied this question as it relates to mid‐ to large‐sized terrestrial fauna in restored riparian habitats at the Sacramento River National Wildlife Refuge, in northern California. We used camera traps to document use of 16 riparian corridor sites of varying restoration age. Comparisons of species richness (diversity) and visitation frequency (activity) were made across different‐aged sites. We found that predator diversity and activity levels tended to be higher in restored forests than in remnant forests, and that they tended to be highest in young restored forests. This trend persisted when data from variable sampling periods were pooled, although significant differences occurred more often in wet and cold sampling periods. The trend did not always hold for the animal community at large (consisting of both predator and non‐predator species). We conclude that restoration age affects predator diversity and activity levels in restored and remnant floodplain forests, and that predator communities can establish soon after restoration. Our results suggest that restoring natural river processes that promote habitat regeneration may benefit mid‐ to large‐sized terrestrial predators that appear to mostly use early successional habitat.     

What is the effect of soil use on ant communities?

Studies on ant communities in agroecosystems have contributed to the knowledge of the effect of agricultural activities on biological communities. The aim of this study is to explain the effect of soil use on ant communities. We tested the hypothesis that there was a decrease in ant species richness and a change in the species composition at habitats with more intense soil use. We collected ants using sardine baits, subterranean traps and direct sampling at four habitats with different soil use (secundary forest, Acacia forestry, initial stage of succession and mixed crops). The ant species richness did not decrease with intensity of soil use. In successional habitat the species numbers collected using sardine baits and subterranean traps were significantly different. Species composition of communities had a pronounced variation, with the epigaeic and hypogaeic ant faunas of the habitat with high intense soil use (mixed crops) had low similarity with ant communities of the three other habitats. The predator species were restricted to habitats with low intensity of soil use. Then, species composition could better reflect the functional changes on ant communities than species richness. Our data can help to choose the component of ant community that better reflect the response of biodiversity to agricultural impacts.     

SPECIES-AREA RELATIONSHIP DURING PRIMARY SUCCESSION IN GRANITE OUTCROP PLANT COMMUNITIES

The species-area relationship and its underlying explanatory mechanisms were investigated in a primary successional sere on a southeastern (United States) granite outcrop. There, plant communities occupy soil-filled depressions separated from one another by areas of bare rock. They have been termed “island communities.” Soil depth and area increase as succession proceeds from Sedum smallii, to lichen-annual, to annual-perennial, and to herb-shrub-tree stages. Although plant species richness is significantly and positively correlated with island area in the system studied (all successional stages considered), the relationship between species richness and island maximum soil depth is stronger. However, island maximum soil depth and area are significantly and positively correlated. The exponential function describes the speciesarea relationship better than the transformed power function. Within successional stage, species richness shows no significant relationship with area or depth, except for late-successional island communities. Processes related to community successional development may explain the species-area relationship that exists when islands of all stages are considered. However, mechanisms related to equilibrium between extinction and immigration may be responsible for the speciesarea relationship for late-successional island communities.     

Long-term responses in population dynamics and diversity of small mammals in riparian and upland habitats within an agricultural landscape

Riparian zones in agricultural landscapes provide linear non-crop habitats for a variety of plant and mammal species, and hence are an important component of biodiversity. To date, variable responses of abundance, species richness, and species diversity of small mammals have been recorded in riparian and upland habitats. To address this variability, we provide a detailed analysis of seasonal changes in abundance and diversity of terrestrial small-mammal communities over a 7-year period within an agricultural landscape in south-central British Columbia, Canada. We tested the hypotheses (H) that abundance, species richness, and species diversity of communities of small mammals (H₁), and demographic parameters of reproduction, recruitment, and survival of the major species: deer mouse (Peromyscus maniculatus) and montane vole (Microtus montanus) (H₂), would be higher in riparian than upland habitats. Mean total abundance of small mammals was higher in summer and winter, and species richness higher in summer, in riparian than hedgerow habitats. Winter population data supported the total and species abundance patterns for small mammals, but species richness was similar, and diversity lower, in riparian than hedgerow sites during winter periods. Deer mice were the dominant species in terms of abundance and reproductive output for pregnancies and recruitment, but not survival, in riparian sites. Montane voles were similar in abundance and demographic parameters in the two habitats. House mice (Mus musculus) preferred hedgerows and wandering shrews (Sorex vagrans) riparian sites. Demographic parameters for deer mice and montane voles indicated that both riparian and hedgerow sites were “source” rather than “sink” habitats, and likely contribute to maintenance of mammal diversity in agricultural landscapes.     

Convergence of community composition during secondary succession on Zokor rodent mounds on the Tibetan Plateau

Aims The community succession theory is much debated in ecology. We studied succession on Zokor rodent mounds on the Tibetan Plateau to address several fundamental questions, among them: (i) During secondary succession, does the community composition converge towards one community state or multiple states depending on the initial colonization? (ii) Do mound communities located in different background communities exhibit different assembly trajectories?Methods In a sub-alpine meadow, we investigated a total of 80 mound communities at several successional stages in three different background communities resulting from different management histories and compared their changes in species composition. The distribution of plant communities over time was analyzed with quantitative classification and ordination methods. The co-occurrence patterns of species were evaluated at each successional stage, and the degree of convergence/divergence among communities was obtained by calculating two beta-diversity indices.Important findings During secondary succession, species richness of mound communities changed over time, and this change was dependent on the background community. Five life-form groups exhibited different dynamic patterns in species richness and plant cover. Community composition and the degree of species co-occurrence between communities increased over time since disturbance. There was much variation in species composition at earlier stages of succession, but communities on older mounds became more similar to each other and to their surrounding vegetation over the course of secondary succession. Post-disturbance succession of Zokor mound communities transitioned from 'multiple alternative states' to 'background-based deterministic community assembly' over time. Tradeoffs between competition and colonization, as well as the characteristics of different life-forms and mass effects within a limited species pool are the mechanisms responsible for convergence of mound communities.     

An ecosystem engineer,the beaver,increases species richness at the landscape scale

Ecosystem engineering - the physical modification of habitats by organisms - has been proposed as an important mechanism for maintaining high species richness at the landscape scale by increasing habitat heterogeneity. Dams built by beaver (Castor canadensis) dramatically alter riparian landscapes throughout much of North America. In the central Adirondacks, New York, USA, ecosystem engineering by beaver leads to the formation of extensive wetland habitat capable of supporting herbaceous plant species not found elsewhere in the riparian zone. We show that by increasing habitat heterogeneity, beaver increase the number of species of herbaceous plants in the riparian zone by over 33% at a scale that encompasses both beaver-modified patches and patches with no history of beaver occupation. We suggest that ecosystem engineers will increase species richness at the landscape scale whenever there are species present in a landscape that are restricted to engineered habitats during at least some stages of their life cycle.     

Marine invertebrates in an algal succession. I. Variations in abundance and diversity with succession

This is a study of the effects of successional algal changes on the community of small invertebrate animals living in the algae at a marine intertidal boulder field. At each of 14 sampling times over a 2-yr period, replicate samples were taken from all stages of succession. By sampling all successional stages simultaneously at each date over an extended period, we could analyze spatial and temporal variations within and among stages. Species richness, diversity, and abundance of the whole invertebrate community increased from early to middle successional stages, while remaining similar from middle to late stages. Evenness remained about constant throughout succession. The increases occur primarily through the addition of species to the community. Some taxonomic groups in this study showed different patterns. Unlike patterns seen in birds, insects, and spiders associated with terrestrial plant succession, species replacements are rare, and there are few early and middle successional specialists. Trophic structure changed markedly during succession. Spatial variation did not change with succession but temporal variation declined. Many of these results are different from the predictions of equilibrium-based theory. Succession in this marine animal community is very different from that in terrestrial animal groups, probably because of the higher rate of exchange of matter in this open coast system.     

Marine invertebrates in an algal succession. II. Tests of hypotheses to explain changes in diversity with succession

The invertebrate community living in algal mats on intertidal boulders was studied for 2 yr. The diversity and abundance of the animals increased between the early and middle stages of algal succession, then remained similar into the later stage. Three possible mechanisms producing this pattern were investigated experimentally by manipulating natural algal mats and plastic algal mimics in the field and laboratory and evaluating the community of colonizing invertebrates. The first, the “ecological time” hypothesis, suggests that there are more species and individuals in later stages because they accumulate slowly with time; this hypothesis was tested experimentally and rejected. A second, “algal toxicity” hypothesis suggests that species richness and abundance are lower in earlier successional stages because the early colonizing green algae are more toxic to animals than are the later red algae. This hypothesis was also tested experimentally and rejected. The third, “habitat complexity” hypothesis suggests that increases in complexity of physical aspects of algal structure (biomass, surface area) cause increases in invertebrate richness and abundance. The fact that this result was found in both living algae and plastic mimics indicates that biological aspects of algal structure apparently have only minor importance. Algal biomass and surface area increase from early to middle successional stages; middle and late successional stages are similar. In general, increases in these physical aspects of algal structure produce concomitant increases in the abundance and diversity of the associated animal community. With higher biomass and surface area, increased numbers of individuals accumulate in algal mats. Because a larger sample of the available pool of individuals is therefore collected, more species are found in a given area of algal mat when the structure is more complex. The successional patterns of increase in species richness of this invertebrate community seem to result from this sampling phenomenon, rather than from increases in numbers of resources (i.e., “niches”).     

Is the productivity of vegetation plots higher or lower when there are more species? Variable predictions from interaction of the ‘sampling effect’ and ‘competitive dominance effect’ on the habitat templet

Using a habitat templet model, we predict that the productivity (total biomass) of plots within a plant community may be positively, negatively or not at all related to variation in the number of species per plot, depending on successional stage (time since major disturbance) and habitat carrying capacity (reflecting the total resource supplying power of the habitat). For plots of a given size, a positive relationship between productivity and species richness is predicted in recently disturbed habitats because local neighbourhoods here will have been assembled largely stochastically, usually from a pool of available species with a right‐skewed size frequency distribution. Hence, in the earliest stages of succession, plots will have relatively high total biomass only if they contain at least some of the relatively uncommon larger species which will, in turn, be more likely in those neighbourhoods that contain more species (the sampling effect). Among these will also be some of the more common smaller species; hence, these high biomass, species‐rich plots should have relatively low species evenness, in contrast to what is predicted under effects involving species complementarity. In late succession, the plots with high total biomass will still be those that contain relatively large species but these plots will now contain relatively few species owing to increased competitive exclusion over time (the competitive dominance effect). In intermediate stages of succession, no relationship between plot productivity and species richness is predicted because the opposing sampling and competitive dominance effects cancel each other out. We predict that the intensity of both the sampling and competitive dominance effects on the productivity/species richness relationship will decrease with decreasing habitat carrying capacity (e.g. decreasing substrate fertility) owing to the inherently lower variance in between‐plot productivity that is predicted for more resource‐impoverished habitats.     

Early succession of butterfly and plant communities on set-aside fields

 Hypotheses on secondary succession of butterfly and plant communities were tested using naturally developed 1- to 4-year-old set-aside fields (n = 16), sown fields (n = 8) and old meadows (n = 4) in 1992 in South Germany. Pioneer successional fields (1st and 2nd year of succession, dominated by annuals) and early successional fields (3rd and 4th year of succession where perennials, especially grasses became dominant) had fewer plant species than mid-successional fields (old meadows). In contrast to established hypotheses, mean number of plant species decreased from 1- to 4-year-old set-aside fields. Species richness of butterfly communities did not change during the first four years of succession, but species composition changed greatly. Pioneer successional fields were characterized by (1) specialized butterflies depending on annual pioneer foodplants (e.g. Issoria lathonia), and (2) species preferring the pioneer successions despite their host plants being more abundant on early and mid-successional fields (e.g. Papilio machaon). The variability in butterfly species richness was best explained by flower abundance which was closely correlated with plant species richness. Species whose abundance was correlated with habitat connectivity were significantly smaller than species which correlated with flower abundance. Numbers of caterpillar species were correlated with numbers of adult butterfly species. Life-history features of butterflies changed significantly from pioneer to early and mid-successional fields. We found decreasing body size and migrational ability, decreasing numbers of species hibernating as imago, decreasing numbers of generations and increasing larval stage duration with age of succession, but, contrary to expectation, host plant specialization, numbers of egg-cluster laying species and egg diameter did not change with successional age. Received 18 September 1995 / Accepted: 17 July 1996     

What predicts the use by brook trout (Salvelinus fontinalis) of terrestrial invertebrate subsidies in headwater streams?

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Succession Influences Wild Bees in a Temperate Forest Landscape: The Value of Early Successional Stages in Naturally Regenerated and Planted Forests

In many temperate terrestrial forest ecosystems, both natural human disturbances drive the reestablishment of forests. Succession in plant communities, in addition to reforestation following the creation of open sites through harvesting or natural disturbances, can affect forest faunal assemblages. Wild bees perform an important ecosystem function in human-altered and natural or seminatural ecosystems, as they are essential pollinators for both crops and wild flowering plants. To maintain high abundance and species richness for pollination services, it is important to conserve and create seminatural and natural land cover with optimal successional stages for wild bees. We examined the effects of forest succession on wild bees. In particular, we evaluated the importance of early successional stages for bees, which has been suspected but not previously demonstrated. A range of successional stages, between 1 and 178 years old, were examined in naturally regenerated and planted forests. In total 4465 wild bee individuals, representing 113 species, were captured. Results for total bees, solitary bees, and cleptoparasitic bees in both naturally regenerated and planted conifer forests indicated a higher abundance and species richness in the early successional stages. However, higher abundance and species richness of social bees in naturally regenerated forest were observed as the successional stages progressed, whereas the abundance of social bees in conifer planted forest showed a concave-shaped relationship when plotted. The results suggest that early successional stages of both naturally regenerated and conifer planted forest maintain a high abundance and species richness of solitary bees and their cleptoparasitic bees, although social bees respond differently in the early successional stages. This may imply that, in some cases, active forest stand management policies, such as the clear-cutting of planted forests for timber production, would create early successional habitats, leading to significant positive effects for bees in general.     

Colonization of dynamic Mediterranean landscapes: where do birds come from after fire?

Aim Two main mechanisms may explain post‐disturbance species colonization patterns of early successional habitats such as those originated by wildfires. First, post‐disturbance colonization is not limited by the dispersal ability of the species to reach the newly created open areas and, secondly, colonization is limited by dispersal. Under the first hypothesis, we expect, at a regional scale, to find similar post‐disturbance communities to develop on recently burned sites. However, colonization limited by dispersal will lead to strong between‐site variations in species composition. Location To test these hypotheses, we studied the post‐fire colonization patterns of nine open‐habitat bird species in eight distantly located wildfires in the north‐eastern Iberian Peninsula. Methods We censused post‐fire bird composition by means of field transects and identified potential colonization sources from species–habitat suitability maps derived from atlas data. Results Our results showed strong significant differences in post‐fire species composition between burnt areas. Burnt areas located in areas with low probability of species presence before the fire event showed lower species occurrence and richness after the fire. Main conclusions These results do not support the idea that early successional stages and open habitats have a homogeneous community structure at regional scales and suggest that dispersal is a key constraint determining bird colonization of post‐fire habitats. Further attention should be paid to landscape heterogeneity as a key factor in determining population dynamics of open‐habitat species in the light of current and future land‐use changes in Mediterranean regions.     

Differences in habitat quality explain nestedness in a land snail meta-community

We set up two alternative hypotheses on how environmental variables could foster nestedness; one of “nested habitats” and another of “nested habitat quality”. The former hypothesis refers to situations where the nestedness of species depends on a nestedness of discrete habitats. The latter considers situations where all species in an assemblage increase in abundance along the same environmental gradient, but differ in specialisation or tolerance. We tested whether litter‐dwelling land snails (terrestrial gastropods) in boreal riparian forest exhibited a nested community structure, whether such a pattern was related to differences in environmental variables among sites, and which of the two hypotheses that best could account for the found pattern. We sampled litter from 100 m² plots in 29 mature riparian forest sites along small streams in the boreal zone of Sweden. The number of snail species varied between 3 and 14 per site. Ranking the species‐by‐site matrix by PCA scores of the first ordination axis revealed a similarly significant nested pattern as when the matrix was sorted by number of species, showing that the species composition in this meta‐community can be properly described as nested. Several environmental variables, most notably pH index, were correlated with the first PCA axis. All but two species had positive eigenvectors in the PCA ordination and the abundance increased considerably along the gradient for most of the species implying that the hypothesis of “nested habitats” was rejected in favour of the “nested habitat quality” hypothesis. Analyses of nestedness have seldom been performed on equal sized plots, and our study shows the importance of understanding that variation in environmental variables among sites can result in nested communities. The conservation implications are different depending on which of our two hypotheses is supported; a conservation focus on species “hotspots” is more appropriate if the communities are nested because of “nested habitat quality”.