Relationships between species richness, evenness, and abundance in a southwestern savanna

Species richness and evenness are components of biological diversity that may or may not be correlated with one another and with patterns of species abundance. We compared these attributes among flowering plants, grasshoppers, butterflies, lizards, summer birds, winter birds, and rodents across 48 plots in the grasslands and mesquite-oak savannas of southeastern Arizona. Species richness and evenness were uncorrelated or weakly negatively correlated for each taxonomic group, supporting the conclusion that richness alone is an incomplete measure of diversity. In each case, richness was positively correlated with one or more measures of abundance. By contrast, evenness usually was negatively correlated with the abundance variables, reflecting the fact that plots with high evenness generally were those where all species present were about equally uncommon. Therefore richness, but not evenness, usually was a positive predictor of places of conservation value, if these are defined as places where species of interest are especially abundant. Species diversity was more positively correlated with evenness than with richness among grasshoppers and flowering plants, in contrast to the other taxonomic groups, and the positive correlations between richness and abundance were comparatively weak for grasshoppers and plants as well. Both of these differences can be attributed to the fact that assemblages of plants and grasshoppers were numerically dominated by small subsets of common species (grasses and certain spur-throated grasshoppers) whose abundances differed greatly among plots in ways unrelated to species richness of the groups as a whole.     

Connell and Slatyer's models of succession in the biodiversity era

Understanding how species interactions drive succession is a key issue in ecology. In this study we show the utility of combining the concepts and methodologies developed within the biodiversity-ecosystem functioning research program with J. H. Connell and R. O. Slatyer's classic framework to understand succession in assemblages where multiple interactions between early and late colonists may include both inhibitory and facilitative effects. We assessed the net effect of multiple species interactions on successional changes by manipulating the richness, composition, and abundance of early colonists in a low-shore assemblage of algae and invertebrates of the northwestern Mediterranean. Results revealed how concomitant changes in species richness and abundance can strongly alter the net effect of inhibitory vs. facilitative interactions on succession. Increasing richness of early colonists inhibited succession, but only under high levels of initial abundance, probably reflecting the formation of a highly intricate matrix that prevented further colonization. In contrast, increasing initial abundance of early colonists tended to facilitate succession under low richness. Thus, changes in abundance of early colonists mediated the effects of richness on succession.     

Effects of site-selection bias on estimates of biodiversity change

Estimates of biodiversity change are essential for the management and conservation of ecosystems. Accurate estimates rely on selecting representative sites, but monitoring often focuses on sites of special interest. How such site-selection biases influence estimates of biodiversity change is largely unknown. Site-selection bias potentially occurs across four major sources of biodiversity data, decreasing in likelihood from citizen science, museums, national park monitoring, and academic research. We defined site-selection bias as a preference for sites that are either densely populated (i.e., abundance bias) or species rich (i.e., richness bias). We simulated biodiversity change in a virtual landscape and tracked the observed biodiversity at a sampled site. The site was selected either randomly or with a site-selection bias. We used a simple spatially resolved, individual-based model to predict the movement or dispersal of individuals in and out of the chosen sampling site. Site-selection bias exaggerated estimates of biodiversity loss in sites selected with a bias by on average 300–400% compared with randomly selected sites. Based on our simulations, site-selection bias resulted in positive trends being estimated as negative trends: richness increase was estimated as 0.1 in randomly selected sites, whereas sites selected with a bias showed a richness change of −0.1 to −0.2 on average. Thus, site-selection bias may falsely indicate decreases in biodiversity. We varied sampling design and characteristics of the species and found that site-selection biases were strongest in short time series, for small grains, organisms with low dispersal ability, large regional species pools, and strong spatial aggregation. Based on these findings, to minimize site-selection bias, we recommend use of systematic site-selection schemes; maximizing sampling area; calculating biodiversity measures cumulatively across plots; and use of biodiversity measures that are less sensitive to rare species, such as the effective number of species. Awareness of the potential impact of site-selection bias is needed for biodiversity monitoring, the design of new studies on biodiversity change, and the interpretation of existing data.     

Incorporating geographical and evolutionary rarity into conservation prioritization

Key goals of conservation are to protect both species and the functional and genetic diversity they represent. A strictly species-based approach may underrepresent rare, threatened, or genetically distinct species and overrepresent widespread species. Although reserves are created for a number of reasons, including economic, cultural, and ecological reasons, their efficacy has been measured primarily in terms of how well species richness is protected, and it is useful to compare how well they protect other measures of diversity. We used Proteaceae species-occurrence data in the Cape Floristic Region of South Africa to illustrate differences in the spatial distribution of species and evolutionary diversity estimated from a new maximum-likelihood molecular phylogeny. We calculated species richness, phylogenetic diversity (i.e., summed phylogenetic branch lengths in a site), and a site-aggregated measure of biogeographically weighted evolutionary distinctiveness (i.e., an abundance weighted measure that captures the unique proportion of the phylogenetic tree a species represents) for sites throughout the Cape Floristic Region. Species richness and phylogenetic diversity values were highly correlated for sites in the region, but species richness was concentrated at a few sites that underrepresented the much more spatially extensive distribution of phylogenetic diversity. Biogeographically weighted evolutionary diversity produced a scheme of prioritization distinct from the other 2 metrics and highlighted southern sites as conservation priorities. In these sites, the high values of biogeographically weighted evolutionary distinctiveness were the result of a nonrandom relation between evolutionary distinctiveness and geographical rarity, where rare species also tended to have high levels of evolutionary distinctiveness. Such distinct and rare species are of particular concern, but are not captured by conservation schemes that focus on species richness or phylogenetic diversity alone.     

Using Parasitic Trematode Larvae to Quantify an Elusive Vertebrate Host

Abstract: Digenean trematode parasites require multiple host species to complete their life cycles, and their abundance can often be strongly correlated with the abundance of their host species. Species richness and abundance of parasites in easily sampled host species may yield an accurate estimate of the species richness and abundance of other hosts in a parasite's life cycle that are difficult to survey directly. Accordingly, we investigated whether prevalence and mean abundance of trematodes could be used to estimate the abundance of one of their host species, diamondback terrapins (Malaclemys terrapin), which are difficult to sample and are designated as near threatened (by the International Union for Conservation of Nature [IUCN Red List]) along some U.S. coasts. As an adult the trematode Pleurogonius malaclemys is specific to terrapins. Its larval stages live first inside mud snails (Ilyanassa obsoleta) and are subsequently shed into the environment where they form external metacercarial cysts on hard surfaces such as snail opercula. The life cycle of P. malaclemys is completed when terrapins ingest these cysts. At 12 sites along the coast of Georgia (U.S.A.), we determined the prevalence of internal P. malaclemys larvae in mud snails (proportion of infected snails in a population) and the prevalence and mean abundance of external trematode cysts. We examined whether these data were correlated with terrapin abundance, which we estimated with mark‐recapture methods. The abundance of external cysts and salinity explained ≥59% of the variability in terrapin abundance. We suggest that dependent linkages between the life stages of multihost parasites make them reliable predictors of host species’ abundance, including hosts with abundances that are challenging to quantify directly.     

Climate, habitat, and species interactions at different scales determine the structure of a Neotropical bat community

Climate, habitat, and species interactions are factors that control community properties (e.g., species richness, abundance) across various spatial scales. Usually, researchers study how a few properties are affected by one factor in isolation and at one scale. Hence, there are few multi-scale studies testing how multiple controlling factors simultaneously affect community properties at different scales. We ask whether climate, habitat structure, or insect resources at each of three spatial scales explains most of the variation in six community properties and which theory best explains the distribution of selected community properties across a rainfall gradient. We studied a Neotropical insectivorous bat ensemble in the Isthmus of Panama with acoustic monitoring techniques. Using climatological data, habitat surveys, and insect captures in a hierarchical sampling design we determined how much variation of the community properties was explained by the three factors employing two approaches for variance partitioning. Our results revealed that most of the variation in species richness, total abundance, and feeding activity occurred at the smallest spatial scale and was explained by habitat structure. In contrast, climate at large scales explained most of the variation in individual species' abundances. Although each species had an idiosyncratic response to the gradient, species richness peaked at intermediate levels of precipitation, whereas total abundance was very similar across sites, suggesting density compensation. All community properties responded in a different manner to the factor and scale under consideration.     

Disturbance influences oyster community richness and evenness, but not diversity

Foundation species in space-limited systems can increase diversity by creating habitat, but they may also reduce diversity by excluding primary space competitors. These contrasting forces of increasing associate diversity and suppressing competitor diversity have rarely been examined experimentally with respect to disturbance. In a benthic marine community in central California, where native oysters are a foundation species, we tested how disturbance influenced overall species richness, evenness, and diversity. Surprisingly, overall diversity did not peak across a disturbance gradient because, as disturbance decreased, decreases in overall species evenness opposed increases in overall species richness. Decreasing disturbance intensity (high oyster abundance) led to increasing species richness of sessile and mobile species combined. This increase was due to the facilitation of secondary sessile and mobile species in the presence of oysters. In contrast, decreasing disturbance intensity and high oyster abundance decreased the evenness of sessile and mobile species. Three factors likely contributed to this decreased evenness: oysters reduced abundances of primary sessile species due to space competition; oysters supported more rare mobile species; and oysters disproportionately increased the relative abundance of a few common mobile species. Our results highlight the need for further studies on how disturbance can differentially affect the evenness and richness of different functional groups, and ultimately how these differences affect the relationship between overall diversity and ecosystem function.     

Experimental tree removal in tallgrass prairie: variable responses of flora and fauna along a woody cover gradient

Woody plant encroachment is a worldwide phenomenon in grassland and savanna systems whose consequence is often the development of an alternate woodland state. Theoretically, an alternate state may be associated with changes in system state variables (e.g., species composition) or abiotic parameter shifts (e.g., nutrient availability). When state-variable changes are cumulative, such as in woody plant encroachment, the probability of parameter shifts increases as system feedbacks intensify over time. Using a Before-After Control-Impact (BACI) design, we studied eight pairs of grassland sites undergoing various levels of eastern redcedar (Juniperus virginiana) encroachment to determine whether responses of flora and fauna to experimental redcedar removal differed according to the level of pretreatment redcedar cover. In the first year after removal, herbaceous plant species diversity and evenness, woody plant evenness, and invertebrate family richness increased linearly with pretreatment redcedar cover, whereas increases in small-mammal diversity and evenness were described by logarithmic trends. In contrast, increases in woody plant diversity and total biomass of terrestrial invertebrates were accentuated at levels of higher pretreatment cover. Tree removal also shifted small-mammal species composition toward a more grassland-associated assemblage. During the second year postremoval, increases in herbaceous plant diversity followed a polynomial trend, but increases in most other metrics did not vary along the pretreatment cover gradient. These changes were accompanied by extremely high growing-season precipitation, which may have homogenized floral and faunal responses to removal. Our results demonstrate that tree removal increases important community metrics among grassland flora and fauna within two years, with some responses to removal being strongly influenced by the stage of initial encroachment and modulated by climatic variability. Our results underscore the importance of decisive management for reversing the effects of woody plant encroachment in imperiled grassland ecosystems.     

Changes in Species Abundance, Distribution, and Diversity in a Central European Bird Community

We analyzed possible causes of changes in species abundance, range size, and diversity as well as extinctions and colonizations in a central European bird community. Using data from the semiquantitative "Lake Constance" breeding bird atlas, we demonstrated that changes in regional abundances from 1980–1981 to 1990–1992 of 151 coexisting bird species were influenced by breeding habitat and migratory status. Significant declines were found in populations of farmland species and long-distance migrants. Farmland species lost parts of their ranges but hardly changed in local abundance in sites where they still occurred. In contrast, declines in long-distance migrants were caused by significant declines in local abundance with only slight loss of occupied range. Regional extinctions and colonizations were predictable from overall population trends. For example, all species that went extinct were either farmland species or long-distance migrants. Avian community composition was influenced by disproportionate declines of abundant species. This led to declines in the total number of breeding pairs and in community biomass and to increases in community evenness, but to only slight declines in species richness. Future conservation efforts in Europe need to focus more on farmland species and on understanding causes for the declines of long-distance migrants.     

Effects of Conversion of Dry Tropical Forest to Agricultural Mosaic on Herpetofaunal Assemblages

Abstract: We explored the impact of forest conversion to agricultural mosaic on anuran, lizard, snake, and turtle assemblages of Neotropical dry forests. Over 2 years, we sampled 6 small watersheds on the west coast of Mexico, 3 conserved and 3 disturbed. The disturbed watersheds were characterized by a mosaic of pastures and cultivated fields (corn, beans, squash) intermingled with patches of different successional stages of dry forest. In each watershed, we conducted 11 diurnal and nocturnal time‐constrained searches in 10 randomly established plots. We considered vulnerability traits of species in relation to habitat modification. Eighteen anuran, 18 lizard, 23 snake, and 3 turtle species were recorded. Thirty‐six species (58%) occurred in both forest conditions, and 14 (22%) and 12 species (19%) occurred only in the conserved and disturbed sites, respectively. Assemblages responded differently to disturbance. Species richness, diversity, and abundance of lizards were higher in disturbed forests. Anuran diversity and species richness were lower in disturbed forest but abundance was similar in both forest conditions. Diversity, richness, and abundance of turtles were lower in disturbed forest. The structure and composition of snake assemblages did not differ between forest conditions. We considered species disturbance sensitive if their abundance was significantly less in disturbed areas. Four anuran (22%), 2 lizard (11%), and 3 turtle (100%) species were sensitive to disturbance. No snake species was sensitive. The decline in abundance of disturbance‐sensitive species was associated with the reduction of forest canopy cover, woody stem cover, roots, and litter‐layer ground cover. Anuran species with small body size and direct embryonic development were especially sensitive to forest disturbance. An important goal for the conservation of herpetofauna should be the determination of species traits associated with extinction or persistence in agricultural mosaics.     

The importance of defining focal assemblages when evaluating amphibian and reptile responses to land use

Habitat loss and degradation are primary threats to amphibians and reptiles, but the relative effects of common land uses on assemblages and the mechanisms that underlie faunal responses are poorly studied. We reviewed the effects of four prevalent types of habitat alteration (urbanization, agriculture, livestock grazing, and silviculture) on amphibian and reptile species richness and abundance by summarizing reported responses in the literature and by estimating effect sizes across studies for species richness in each land‐use type. We then used a multinomial model to classify species as natural habitat specialists, generalists, and disturbed habitat specialists and examined variation in effect sizes for each land‐use type according to habitat specialization categories. There were mixed conclusions from individual studies, some reporting negative, neutral, or positive effects of land use on species richness and total abundance. A large proportion of studies reported species‐specific effects of individual species abundance. However, in our analysis of effect sizes, we found a general trend of negative effects of land use on species richness. We also demonstrate that habitat associations of common species and species turnover can explain variation in the effect of land use on herpetofauna. Our review highlights the pervasive negative effects of common land uses on amphibians and reptiles, the importance of identifying groups vulnerable to land‐use change (e.g., forest‐associated species) in conservation studies, and the potential influence of disturbance‐associated species on whole assemblage analyses.     

Species richness and stand stability in conifer forests of the Sierra Nevada

Theoretical and empirical studies have long suggested that stability and complexity are intimately related, but evidence from long-lived systems at large scales is lacking. Stability can either be driven by complex species interactions, or it can be driven by the presence/absence and abundance of a species best able to perform a specific ecosystem function. We use 64 years of stand productivity measures in forest systems composed of four dominant conifer tree species to contrast the effect of species richness and abundance on three stability measures. To perform this contrast, we measured the annual growth increments of > 900 trees in mixed and pure forest stands to test three hypotheses: increased species richness will (1) decrease stand variance, (2) increase stand resistance to drought events, and (3) increase stand resilience to drought events. In each case, the alternate hypothesis was that species richness had no effect, but that species composition and abundance within a stand drove variance, resistance, and resilience. In pure stands, the four species demonstrated significant differences in productivity, and in their resistance and resilience to drought events. The two pine species were the most drought resistant and resilient, whereas mountain hemlock was the least resistant and resilient, and red fir was intermediate. For community measures we found a moderately significant (P = 0.08) increase in the community coefficient of variation and a significant (P = 0.03) increase in resilience with increased species richness, but no significant relationship between species richness and community resistance, though the variance in community resistance to drought decreased with species richness. Community resistance to drought was significantly (P = 0.001) correlated to the relative abundance of lodgepole pine, the most resistant species. We propose that resistance is driven by competition for a single limiting resource, with negative diversity effects. In contrast resilience measures the capacity of communities to partition resources in the absence of a single limiting resource, demonstrating positive diversity effects.     

Animals on marine flowers: does the presence of flowering shoots affect mobile epifaunal assemblage in an eelgrass meadow?

Eelgrass, Zostera marina, produces two types of shoots: morphologically simple vegetative shoots and highly branched flowering (reproductive) shoots, the latter found only in summer months. We examined whether the abundance and diversity of mobile epifaunal assemblage are affected by the presence of flowering shoots in an eelgrass meadow of Otsuchi Bay, northeastern Japan. Comparisons of epifauna in natural vegetation revealed that density and species richness did not differ significantly between sites consisting of both flowering and vegetative shoots, and those only of vegetative shoots. A transplant experiment, conducted to examine the colonization rates of epifauna to defaunated eelgrass planted with different combination of vegetative and flowering shoots, showed no obvious variation in abundance and species richness. At species level, the density of some species such as a tanaid Zeuxo sp. and a polychaete Platynereis sp. was higher at sites and/or treatments with flowering shoots, whereas that of some gastropods, such as Lirularia iridescens and Siphonacmea oblongata was higher at sites without flowering shoots. The species-specific response led to dissimilarity of epifaunal assemblage between sites and among treatments with different densities of vegetative and flowering shoots. Similar patterns observed for natural vegetation and the transplant experiment suggest that the variation in assemblage structure is caused by habitat selection of each species, for example, the utilization of flowering shoots as feeding ground and nursery by Zeuxo sp.     

Effects of Forest Management on Amphibians and Reptiles in Missouri Ozark Forests

Abstract:  As part of the Missouri Ozark Forest Ecosystem Project (MOFEP), we experimentally evaluated the impacts of forest management on the relative abundance of amphibians and reptiles in Missouri's Ozark forests (U.S.A.). Using large study sites (average size of 400 ha) as the experimental unit, we tested the effects of uneven-aged and even-aged forest management treatments compared with no-harvest management (i.e., control) on the relative abundance of 13 focal amphibian and reptile species. Within even-aged management sites, we also focused on the local-scale effects of clearcutting on these species by comparing relative abundance among plots located within clearcut stands, 50 m away from clearcut stands, and 200 m away from clearcut stands. Pretreatment sampling of species abundance occurred from 1992 through 1995, and post-treatment sampling occurred from 1997 through 2000. At the landscape scale, treatment significantly affected the abundance of Bufo americanus . This species declined less on even-aged management sites than on control sites, but the general decline on all sites suggests that other factors may have contributed to this result. Within even-aged management sites, most amphibian species declined and some reptile species increased relative to pretreatment abundances within clearcut stands. We found significant effects of distance from clearcut for two amphibian species, Ambystoma maculatum and Rana clamitans, and two reptile species, Scincella lateralis and Sceloporus undulatus . In general, we conclude that clearcuts within even-aged management sites locally affected amphibian and reptile species but, at a larger spatial scale, we did not detect significant effects of even-aged and uneven-aged forest management. These findings represent relatively short-term data but suggest that forest management and maintenance of biodiversity may be compatible when relatively small amounts of the landscape are disturbed.     

Effects of Agricultural Cultivation on Demographics of Southern High Plains Amphibians

Abstract:  Anthropogenic disturbance of landscapes surrounding wetlands is considered a factor in local and global amphibian declines. Few data exist on the effects of agricultural cultivation of wetland watersheds on amphibians, and results from previous studies are contradictory. Our objective was to test the effects of general anthropogenic land use (cultivation vs. grassland) on the demographics of seven species and three age classes of amphibians in the Southern High Plains of Texas. We partially enclosed 16 playa wetlands (4 per land use per year) with drift fences and pitfall traps and monitored relative daily abundance and diversity from 16 May to 17 October 1999 and 19 April to 18 August 2000. In general, relative abundance (i.e., average daily capture) of New Mexico and plains spadefoots (  Spea multiplicata, S. bombifrons ) was greater at cropland than grassland playas; the abundance of other species and diversity of the amphibian assemblage was not affected by land use. Also, abundance generally was greater in 1999 than 2000 for metamorph spadefoots and barred tiger salamanders ( Ambystoma tigrinum mavortium ). Differences in spadefoot abundance between land-use types may have been related to low species-specific vagility, resulting in increased nestedness within disturbed landscapes and reduced abundance of a potential keystone intraguild predator in cropland playas. The yearly difference in amphibian abundance was likely related to annual precipitation, which influenced wetland hydroperiod. Agricultural cultivation surrounding wetlands is associated with the increased abundance of some amphibian species, but other demographic and fitness parameters—such as temporal demographics, body size, and diet diversity—may be negatively affected.     

Change in avian functional fingerprints of a Neotropical montane forest over 100 years as an indicator of ecosystem integrity

Ecologically relevant traits of organisms in an assemblage determine an ecosystem's functional fingerprint (i.e., the shape, size, and position of multidimensional trait space). Quantifying changes in functional fingerprints can therefore provide information about the effects of diversity loss or gain through time on ecosystem condition and is a promising approach to monitoring ecological integrity. This, however, is seldom possible owing to limitations in historical surveys and a lack of data on organismal traits, particularly in diverse tropical regions. Using data from detailed bird surveys from 4 periods across more than a century, and morphological and ecological traits of 233 species, we quantified changes in the avian functional fingerprint of a tropical montane forest in the Andes of Colombia. We found that 78% of the variation in functional space, regardless of period, was described by 3 major axes summarizing body size, dispersal ability (indexed by wing shape), and habitat breadth. Changes in species composition significantly altered the functional fingerprint of the assemblage and functional richness and dispersion decreased 35–60%. Owing to species extirpations and to novel additions to the assemblage, functional space decreased over time, but at least 11% of its volume in the 2010s extended to areas of functional space that were unoccupied in the 1910s. The assemblage now includes fewer large-sized species, more species with greater dispersal ability, and fewer habitat specialists. Extirpated species had high functional uniqueness and distinctiveness, resulting in large reductions in functional richness and dispersion after their loss, which implies important consequences for ecosystem integrity. Conservation efforts aimed at maintaining ecosystem function must move beyond seeking to sustain species numbers to designing complementary strategies for the maintenance of ecological function by identifying and conserving species with traits conferring high vulnerability such as large body size, poor dispersal ability, and greater habitat specialization. Article impact statement: Changes in functional fingerprints provide a means to quantify the integrity of ecological assemblages affected by diversity loss or gain.     

Using community‐level metrics to monitor the effects of marine protected areas on biodiversity

Marine protected areas (MPAs) are used to protect species, communities, and their associated habitats, among other goals. Measuring MPA efficacy can be challenging, however, particularly when considering responses at the community level. We gathered 36 abundance and 14 biomass data sets on fish assemblages and used meta‐analysis to evaluate the ability of 22 distinct community diversity metrics to detect differences in community structure between MPAs and nearby control sites. We also considered the effects of 6 covariates—MPA size and age, MPA size and age interaction, latitude, total species richness, and level of protection—on each metric. Some common metrics, such as species richness and Shannon diversity, did not differ consistently between MPA and control sites, whereas other metrics, such as total abundance and biomass, were consistently different across studies. Metric responses derived from the biomass data sets were more consistent than those based on the abundance data sets, suggesting that community‐level biomass differs more predictably than abundance between MPA and control sites. Covariate analyses indicated that level of protection, latitude, MPA size, and the interaction between MPA size and age affect metric performance. These results highlight a handful of metrics, several of which are little known, that could be used to meet the increasing demand for community‐level indicators of MPA effectiveness.     

Offsets and Conservation of the Species of the EU Habitats and Birds Directives

Biodiversity offsets are intended to achieve no net loss of biodiversity due to economic and human development. A variety of biodiversity components are addressed by offset policies. It is required that loss of protected species due to development be offset under the EU Habitats and Birds Directives in Europe. We call this type of offset a species‐equality offset because the offset pertains to the same species affected by the development project. Whether species equality can be achieved by offset design is unknown. We addressed this gap by reviewing derogation files (i.e., specific files that describe mitigation measures to ensure no net loss under the EU Habitats and Birds Directives) from 85 development projects in France (2009–2010). We collected information on type of effect (reversible vs. irreversible) and characteristics of affected and offset sites (i.e., types of species, total area). We analyzed how the type of effect and the affected‐site characteristics influenced the occurrence of offset measures. The proportion of species targeted by offset measures (i.e., offset species) increased with the irreversibility of the effect of development and the conservation status of the species affected by development (i.e., affected species). Not all effects on endangered species (International Union for Conservation of Nature Red List) were offset; on average, 82% of affected species would be offset. Twenty‐six percent of species of least concern were offset species. Thirty‐five percent of development projects considered all affected species in their offset measures. Species richness was much lower in offset sites than in developed sites even after offset proposals. For developed areas where species richness was relatively high before development, species richness at offset sites was 5–10 times lower. The species‐equality principle appears to have been applied only partially in offset policies, as in the EU directives. We suggest the application of this principle through offsets is highly important for the long‐term conservation of biodiversity in Europe. Compensaciones y Conservación de las Especies de las Directivas de Hábitats y Aves de la UE     

Spatial scale and species identity influence the indigenous-alien diversity relationship in springtails

Although theory underlying the invasion paradox, or the change in the relationship between the richness of alien and indigenous species from negative to positive with increasing spatial scale, is well developed and much empirical work on the subject has been undertaken, most of the latter has concerned plants and to a lesser extent marine invertebrates. Here we therefore examine the extent to which the relationships between indigenous and alien species richness change from the local metacommunity to the interaction neighborhood scales, and the influences of abundance, species identity, and environmental favorability thereon, in springtails, a significant component of the soil fauna. Using a suite of modeling techniques, including generalized least squares and geographically weighted regressions to account for spatial autocorrelation or nonstationarity of the data, we show that the abundance and species richness of both indigenous and alien species at the metacommunity scale respond strongly to declining environmental favorability, represented here by altitude. Consequently, alien and indigenous diversity covary positively at this scale. By contrast, relationships are more complex at the interaction neighborhood scale, with the relationship among alien species richness and/or density and the density of indigenous species varying between habitats, being negative in some, but positive in others. Additional analyses demonstrated a strong influence of species identity, with negative relationships identified at the interaction neighborhood scale involving alien hypogastrurid springtails, a group known from elsewhere to have negative effects on indigenous species in areas where they have been introduced. By contrast, diversity relationships were positive with the other alien species. These results are consistent with both theory and previous empirical findings for other taxa, that interactions among indigenous and alien species change substantially with spatial scale and that environmental favorability may play a key role in explaining the larger scale patterns. However, they also suggest that the interactions may be affected by the identity of the species concerned, especially at the interaction neighborhood scale.