Quantifying determinants contributing to plant species richness in mosaic landscapes: a single- and multi-patch perspective

Despite good theoretical knowledge about determinants of plant species richness in mosaic landscapes, validations based on complete surveys are scarce. We conducted a case study in a highly fragmented, traditional agricultural landscape. In 199 patches of 20 representative multi-patch-plots (MPPs, 1 ha) we recorded a total of 371 plant species. In addition to an additive partitioning of species diversity at the (a) patch- and (b) MPP-scale, we adopted the recently proposed ‘specificity’ measure to quantify the contribution of a spatial subunit to landscape species richness (subunit-to-landscape-contribution, SLC). SLC-values were calculated at both scales with respect to various spatial extents. General regression models were used to quantify the relative importance of hypothesis-driven determinants for species richness and SLC-values. At the patch scale, habitat type was the main determinant of species richness, followed by area and elongated shape. For SLC-values, area was more important than habitat type, and its relevance increased with the extent of the considered landscape. Influences of elongated shape and vegetation context were minor. Differences between habitat types were pronounced for species richness and also partly scale-dependent for SLC-values. Relevant predictors at the MPP-scale were nonlinear habitat richness, the gradient from anthropogenic to seminatural vegetation, and the proportions of natural vegetation and rare habitats. Linear elements and habitat configuration did not contribute to species richness and SLC. Results at the MPP-scale were in complete accordance with the predictions of the mosaic concept. Hence, our study represents its first empirical validation for plant species diversity in mosaic landscapes.     

Matrix is important for mammals in landscapes with small amounts of native forest habitat

Acknowledgment that the matrix matters in conserving wildlife in human-modified landscapes is increasing. However, the complex interactions of habitat loss, habitat fragmentation, habitat condition and land use have confounded attempts to disentangle the relative importance of properties of the landscape mosaic, including the matrix. To this end, we controlled for the amount of remnant forest habitat and the level of fragmentation to examine mammal species richness in human-modified landscapes of varying levels of matrix development intensity and patch attributes. We postulated seven alternative models of various patch habitat, landscape and matrix influences on mammal species richness and then tested these models using generalized linear mixed-effects models within an information theoretic framework. Matrix attributes were the most important determinants of terrestrial mammal species richness; matrix development intensity had a strong negative effect and vegetation structural complexity of the matrix had a strong positive effect. Distance to the nearest remnant forest habitat was relatively unimportant. Matrix habitat attributes are potentially a more important indicator of isolation of remnant forest patches than measures of distance to the nearest patch. We conclude that a structurally complex matrix within a human-modified landscape can provide supplementary habitat resources and increase the probability of movement across the landscape, thereby increasing mammal species richness in modified landscapes.     

Scale-dependent importance of environment, land use and landscape structure for species richness and composition of SE Norwegian modern agricultural landscapes

Knowledge of variation in vascular plant species richness and species composition in modern agricultural landscapes is important for appropriate biodiversity management. From species lists for 2201 land-type patches in 16 1-km² plots five data sets differing in sampling-unit size from patch to plot were prepared. Variation in each data set was partitioned into seven sources: patch geometry, patch type, geographic location, plot affiliation, habitat diversity, ecological factors, and land-use intensity. Patch species richness was highly predictable (75% of variance explained) by patch area, within-patch heterogeneity and patch type. Plot species richness was, however, not predictable by any explanatory variable, most likely because all studied landscapes contained all main patch types – ploughed land, woodland, grassland and other open land – and hence had a large core of common species. Patch species composition was explained by variation along major environmental complex gradients but appeared nested to lower degrees in modern than in traditional agricultural landscapes because species-poor parts of the landscape do not contain well-defined subsets of the species pool of species-rich parts. Variation in species composition was scale dependent because the relative importance of specific complex gradients changed with increasing sampling-unit size, and because the amount of randomness in data sets decreased with increasing sampling-unit size. Our results indicate that broad landscape structural changes will have consequences for landscape-scale species richness that are hard or impossible to predict by simple surrogate variables.     

The landscape matrix modifies the effect of habitat fragmentation in grassland butterflies

The landscape matrix is suggested to influence the effect of habitat fragmentation on species richness, but the generality of this prediction has not been tested. Here, we used data from 10 independent studies on butterfly species richness, where the matrix surrounding grassland patches was dominated by either forest or arable land to test if matrix land use influenced the response of species richness to patch area and connectivity. To account for the possibility that some of the observed species use the matrix as their main or complementary habitat, we analysed the effects on total species richness and on the richness of grassland specialist and non-specialist (generalists and specialists on other habitat types) butterflies separately. Specialists and non-specialists were defined separately for each dataset. Total species richness and the richness of grassland specialist butterflies were positively related to patch area and forest cover in the matrix, and negatively to patch isolation. The strength of the species-area relationship was modified by matrix land use and had a slope that decreased with increasing forest cover in the matrix. Potential mechanisms for the weaker effect of grassland fragmentation in forest-dominated landscapes are (1) that the forest matrix is more heterogeneous and contains more resources, (2) that small grassland patches in a matrix dominated by arable land suffer more from negative edge effects or (3) that the arable matrix constitutes a stronger barrier to dispersal between populations. Regardless of the mechanisms, our results show that there are general effects of matrix land use across landscapes and regions, and that landscape management that increases matrix quality can be a complement to habitat restoration and re-creation in fragmented landscapes.     

Successional change in the insect community of a fragmented landscape

Habitat fragmentation strongly affects insect species diversity and community composition, but few studies have examined landscape effects on long term development of insect communities. As mobile consumers, insects should be sensitive to both local plant community and landscape context. We tested this prediction using sweep-net transects to sample insect communities for 8 years at an experimentally fragmented old-field site in northeastern Kansas, USA. The site included habitat patches undergoing secondary succession, surrounded by a low turf matrix. During the first 5 years, plant richness and cover were measured in patches. Insect species richness, total density, and trophic diversity increased over time on all transects. Cover of woody plants and perennial forbs increased each year, adding structural complexity to successional patches and potentially contributing to increased insect diversity. Within years, insect richness was significantly greater on transects through large successional patches (5000 m²) than on transects through fragmented arrays of 6 medium-sized (total area 1728 m²) or 15 small (480 m²) patches. However, plant cover did not differ among patch types and was uncorrelated with insect richness within years. Insect richness was strongly correlated with insect density, but trophic and α diversities did not differ among patch types, indicating that patch insect communities were subsets of a common species pool. We argue that differences in insect richness resulted from landscape effects on the size of these subsets, not patch succession rates. Greater insect richness on large patches can be explained as a community-level consequence of population responses to resource concentration.     

Effects of landscape pattern on bird species distribution in the Mt. Lofty Ranges,South Australia

We assessed how well landscape metrics at 2, 5, and 10 km scales could explain the distribution of woodland bird species in the Mount Lofty Ranges, South Australia. We considered 31 species that have isolated or partially isolated populations in the region and used the Akaike Information Criterion to select a set of candidate logistic regression models. The 2 km distance was the most appropriate scale for a plurality of the species. While the total amount of area of native vegetation around a site was the most important determining factor, the effect of landscape configuration was also important for many species. Most species responded positively to area-independent fragmentation, but the responses to mean patch isolation and mean patch shape were more variable. Considering a set of candidate models for which there is reasonable support (Akaike weights > 0.10), 12 species responded negatively to landscapes with highly linear and isolated patches. No clear patterns emerged in terms of taxonomy or functional group as to how species respond to landscape configuration. Most of the species had models with relatively good discrimination (12 species had ROC values > 0.70), indicating that landscape pattern alone can explain their distributions reasonably well. For six species there were no models that had strong weight of evidence, based on the AIC and ROC criteria. This analysis shows the utility of the Akaike Information Criterion approach to model selection in landscape ecology. Our results indicate that landscape planners in the Mount Lofty Ranges must consider the spatial configuration of vegetation. This revised version was published online in July 2006 with corrections to the Cover Date.     

Influence of patch and landscape characteristics on the distribution of the subterranean rodent <Emphasis Type="Italic">Ctenomys porteousi</Emphasis>

The understanding and prediction of the responses of animal populations to habitat fragmentation is a central issue in applied ecology. The identification of habitat variables associated to patch occupancy is particularly important when habitat quality is affected by human activities. Here, we analyze the influence of patch and landscape characteristics on patch occupancy by the subterranean herbivorous rodent Ctenomys porteousi. Patch occupancy was monitored in a network of 63 habitat patches identified by satellite imagery analysis which extends along almost the whole distributional range for C. porteousi. Suitable habitat for the occurrence of C. porteousi is highly fragmented and represents <10% of the total area in its distributional range. The distribution of C. porteousi in the patch network is affected not only by characteristics of the habitat patches, but also by those of the surrounding landscape matrix. Significant differences between occupied and empty patches were found in several environmental variables. Overall, occupied patches were larger, less vegetated, more connected, and had larger neighbor patches than empty patches. A stepwise procedure on a generalized linear model selected four habitat variables that explain patch occupancy in C. porteousi; it included the effects of habitat quality in the matrix surrounding the patch, average vegetation cover in the patch, minimum vegetation cover in the matrix surrounding the patch, and the area of the nearest neighbor patch. These results indicate that patch occupancy in C. porteousi is strongly influenced by the availability and quality of habitat both in the patch and in the surrounding landscape matrix.     

Habitat specificity of patches in modern agricultural landscapes

Habitat specificity analysis provides a tool for partitioning landscape species diversity on landscape elements by separating patches with many rare specialist species from patches with the same number of species, all of which are common generalists and thus provide information of relevance to conservation goals at regional and national levels. Our analyses were based upon species data from 2201 patch elements in SE Norwegian modern agricultural landscapes. The context used for measuring habitat specificity strongly influences the results. In general the gamma diversity contribution and core habitat specificity calculated from the patch data set were correlated. High values for both measures were observed for woodland, pastures and road verges whereas midfield islets and boundary transitional types were ranked low, as opposed to findings in traditional, extensively managed agricultural landscapes. This is due to our study area representing intensively used agricultural landscape elements holding a more trivial species composition, in addition to ruderals being favoured by fertility and disturbance, a finding also being supported by the semi-natural affiliation index. Results obtained by use of checklist data from the same study area diverged from patch data. Caution is needed in interpretation of habitat specificity results obtained from checklist data, because modern agricultural landscapes contain several land types which are seldom surveyed by botanists, thus being under-represented in the data set. We propose the use of core habitat specificity and gamma diversity contribution in parallel to obtain a value neutral diversity assessment that addresses patch uniqueness and other properties of conservation interests.     

Quantifying the landscape influence on plant invasions in Mediterranean coastal habitats

Landscape pattern might be an important determinant of non-native plant invasions because it encompasses components influencing the availability of non-native plant propagules and disturbance regimes. We aimed at exploring the relative role of patch and landscape characteristics, compared to those of habitat type and regional human influence on non-native plant species richness. For this purpose, we identified all non-native plant species in 295 patches of four coastal habitat types across three administrative regions in NE Spain differing in the degree of human influence. For each patch, we calculated several variables reflecting habitat patch geometry (size and shape), landscape composition (distribution of land-cover categories) and landscape configuration (arrangement of patches). The last two groups of variables were calculated at five different spatial extents. Landscape composition was by far the most important group of variables associated with non-native species richness. Natural areas close to diverse and urban landscapes had a high number of non-native species while surrounding agricultural areas could buffer this effect. Regional human influence was also strongly associated with non-native species richness while habitat type was the least important factor. Differences in sensitivity of landscape variables across spatial extents proved relevant, with 100 m being the most influential extent for most variables. These results suggest that landscape characteristics should be considered for performing explicit spatial risk analyses of plant invasions. Consequently, the management of invaded habitats should focus not only at the stand scale but also at the highly influential neighbouring landscape. Prior to incorporate landscape characteristics into management decisions, sensitivity analyses should be taken into account to avoid inconsistent variables.     

Quantifying habitat specificity to assess the contribution of a patch to species richness at a landscape scale

Assessing and predicting the species richness of a complex landscape remains a problem because there is no simple scaling function of species richness in a heterogeneous environment. Furthermore, the potential value of an area for biodiversity conservation may depend on which, rather than how many, species the area contains. This paper shows how we can objectively evaluate the contribution of an area, e.g., a habitat patch, to larger-scale plant species richness, e.g., a landscape composed of patches of several habitat types, and how we can test hypotheses that attempt to explain this contribution. We quantified the concept of habitat specificity to assess the proportion of each observed plant population that is concentrated within a given spatial element. A case study of a biodiversity-monitoring program in the Swiss Canton of Aargau showed that the relative contribution of the three main types of land use to the overall species richness differed strongly between higher taxa (vascular plants and molluscs). However, the type of data, i.e., presence-absence or abundance, was not important. Resampling of the plant data suggested that stratification provided an unbiased estimate of relative specificity, whereas unstratified sampling caused bias even for large samples. In a second case study of vascular plants in an agricultural landscape in central Switzerland, we tested whether the type, size or shape of a landscape element can predict its contribution to the species richness of the landscape. Habitat types that were less frequently disturbed contributed more per m² to landscape species richness than more frequently disturbed ones. Contrary to expectation, patch size was negatively correlated to specificity per m² for arable fields, whereas patch shape appeared to be unrelated to the specificity per m² both for arable fields and for meadows. The specificity approach provides a solution to the problem of scaling species richness and is ideally suited for testing hypotheses on the effect of landscape structure on landscape species richness. Specificity scores can easily be combined with measures of other aspects of rarity to assess the contribution of a spatial element to conservation goals formulated at regional, national or global level.     

The concept of habitat specificity revisited

Habitat specificity indices reflect richness (α) and/or distinctiveness (β) components of diversity. The latter may be defined by α and γ (landscape) diversity in two alternative ways: multiplicatively () and additively (). We demonstrate that the original habitat specificity concept of Wagner and Edwards (Landscape Ecol 16:121–131, 2001) consists of three independent components: core habitat specificity (uniqueness of the species composition), patch area and patch species richness. We describe habitat specificity as a family of indices that may include either area or richness components, or none or both, and open for use of different types of mean in calculation of core habitat specificity. Core habitat specificity is a beta diversity measure: the effective number of completely distinct communities in the landscape. Habitat specificity weighted by species number is a gamma diversity measure: the effective number of species that a patch contributes to landscape richness. We compared 12 habitat specificity indices by theoretical reasoning and by use of field data (vascular plant species in SE Norwegian agricultural landscapes). Habitat specificity indices are strongly influenced by weights for patch area and patch species richness, and the relative contribution of rare vs. common species (type of mean). The relevance of properties emphasized by each habitat specificity index for evaluation of patches in a biodiversity context is discussed. Core habitat specificity is emphasized as an ecologically interpretable measure that specifically addresses patch uniqueness while habitat specificity weighted by species number combines species richness and species composition in ways relevant for conservation biological assessment. Electronic supplementary material  The online version of this article (doi:) contains supplementary material, which is available to authorized users.     

Spatial configuration matters: a test of the habitat amount hypothesis for plants in calcareous grasslands

Context

A recent hypothesis, the habitat amount hypothesis, predicts that the total amount of habitat in the landscape can replace habitat patch size and isolation in studies of species richness in fragmented landscapes.

Objectives

To test the habitat amount hypothesis by first evaluating at which spatial scale the relationship between species richness in equal-sized sample quadrats and habitat amount was the strongest, and then test the importance of spatial configuration of habitat—measured as local patch size and isolation—when habitat amount was taken into account.

Methods

A quasi-experimental setup with 20 habitat patches of dry calcareous grasslands varying in patch size, patch isolation and habitat amount at the landscape scale was established in the inner Oslo fjord, Southern Norway. We recorded species richness of habitat specialists of vascular plants in equal-sized sample quadrats and analysed the relationship between species richness, habitat amount in the landscape and patch size and isolation.

Results

Although the total amount of habitat in a 3 km-radius around the local patch was positively related to species richness in the sample quadrats, local patch size had an additional positive effect, and the effect of patch size was higher when the amount of habitat within the 3 km-radius was high than when it was low.

Conclusions

In our study system of specialist vascular plants in dry calcareous grasslands, we do not find support for the habitat amount hypothesis.

     

Aspen patch and migratory bird relationships in the northern Yellowstone ecosystem

We evaluated the effects of aspen patch area and orientation (relative to North and an elevational gradient) on the early breeding season abundance and species richness of migratory and resident birds in the northern ungulate winter range of the Yellowstone ecosystem, USA. Using an information-theoretic model selection approach, we found patch area and basal area of aspen to be the most important covariates for long distance migrants, and patch orientation relative to elevational gradient the most important covariate for residents/short-distance migrants. Basal area of live aspen and aspen snags was marginally important for both migratory strategies, likely because aspen snags are an important habitat for most cavity-nesting species. Landscape ecological theory postulates passive interception of dispersing or migrating organisms by patches of suitable habitat. Our results suggest that residents/short-distance migrants are intercepted by patches that are oriented perpendicular to the elevational gradient of our study region resulting in greater abundances and species richness in those patches. However, long-distance migrants appear to use aspen patches without regard to orientation, but rather to patch area.     

Spatial dynamics of the knob-tailed gecko Nephrurus stellatus in a fragmented agricultural landscape

In fragmented landscapes, a species?? dispersal ability and response to habitat condition are key determinants of persistence. To understand the relative importance of dispersal and condition for survival of Nephrurus stellatus (Gekkonidae) in southern Australia, we surveyed 92 woodland remnants three times. This gecko favours early post-fire succession conditions so may be at risk of extinction in the long-unburnt agricultural landscape. Using N-mixture models, we compared the influence of four measures of isolation, patch area and two habitat variables on the abundance and occurrence of N. stellatus, while taking into account detection probability. Patch occupancy was high, despite the long-term absence of fire from most remnants. Distance to the nearest occupied site was the most informative measure of patch isolation, exhibiting a negative relationship with occupancy. Distance to a nearby conservation park had little influence, suggesting that mainland?Cisland metapopulation dynamics are not important. Abundance and occurrence were positively related to ?%-cover of spinifex (Triodia), indicating that niche-related factors may also contribute to spatial dynamics. Patterns of patch occupancy imply that N. stellatus has a sequence of spatial dynamics across an isolation gradient, with patchy populations and source-sink dynamics when patches are within 300?m, metapopulations at intermediate isolation, and declining populations when patches are separated by >1?C2?km. Considering the conservation needs of the community, habitat condition and connectivity may need to be improved before fire can be reintroduced to the landscape. We speculate that fire may interact with habitat degradation and isolation, increasing the risk of local extinctions.     

Predicting mammal species richness and distributions: testing the effectiveness of satellite-derived land cover data

Mapping species richness and distributions is an important aspect of conservation and land use planning, but the time and cost of producing maps from field surveys is prohibitive. It is useful, therefore, if mappable environmental variables, from a readily accessible source, can be used as surrogates for species attributes. We evaluated the power of satellite-derived land cover information, from the Land Cover Map of Great Britain, to predict species richness and occurrences of terrestrial mammals in one hundred 10×10 km quadrats, from four regions of Britain. The predictive power of the land cover data was relatively poor – with a few exceptions, land cover explained less than half of the variation in mammal species richness and occurrence in regression models. Predictive power was considerably stronger when regions were analyzed separately than when analyzed together, and best fitting models varied between regions and between mammal taxa. Predictive power was also affected (positively or negatively depending on taxon) when PCA-ordinated land cover variables were used as predictors. The predictive strength of the land cover data was probably limited mostly by the high proportion of British mammal species with geographic distributions changing rapidly and independently of land cover (and hence the non-saturation of preferred habitats), and to a lesser extent by shortcomings in the mammal and land cover data, and the influence of landscape factors other than land cover on mammal distributions. The results suggest that regional stratification is essential when attempting to predict species richness and distributions, even across relatively limited areas such as Great Britain. We conclude that caution is necessary in using results from environmental information systems such as this as a basis for conservation and land use planning decisions.     

Interception of moving organisms: influences of patch shape,size, and orientation on community structure

Island biogeographers have predicted that in oceanic systems, oblong islands oriented perpendicular to the dispersal paths of organisms should intercept more species and individuals than (1) circular islands of the same size, and (2) oblong islands of equal area oriented parallel to the direction of travel. Landscape ecologists expect similar relations with habitat patches in a terrestrial matrix. Yet in neither situation is there adequate empirical information to permit conclusions about the prevalence of such effects. To test the hypothesis that intercept-related patch variables influence community structure on the landscape scale, we studied relations between the richness and abundance of cavity-nesting birds and patch shape, size, and orientation relative to a northerly migration path. The influences of other patch features on nest abundances were removed analytically. Multiple regression indicated that the mean and total number of nesting species, and nest abundances for migrants were significantly associated with patch orientation or a patch area x orientation interaction, but not patch shape. Nest abundances for permanent residents were not associated with patch shape or orientation, although area effects, possibly reflecting dispersal interception, were evident. These results are consistent with the hypothesis that stochastic interception of migrating or dispersing organisms influences patch community structure. In addition to richness and abundance effects apparent in this analysis, the sex ratio, age structure, growth rate, social structure, and genetic features of patch populations may also be influenced. The interception of moving organisms by patches may thus be a key factor influencing population and community persistence in reserves. If so, landscape structure could be manipulated to maximize the interception of dispersing or migrating organisms, or minimize it if the effects are undesirable.     

Matrix Matters: Effects of Surrounding Land Uses on Forest Birds Near Ottawa,Canada

Matrix quality affects probability of persistence in habitat patches in landscape simulation models while empirical studies show that both urban and agricultural land uses affect forest birds. However, due to the fact that forest bird abundance and species richness can be strongly influenced by local habitat factors, it is difficult to analyze matrix effects without confounding effects from such factors. Given this, our objectives were to (1) relate human-dominated land uses to forest bird abundance and species richness without confounding effects from other factors; (2) determine the scale at which forest birds respond to the matrix; and (3) identify whether certain bird migratory strategies or habitat associations vary in richness or abundance as a function of urban and agriculture land uses. Birds were surveyed at a single point count site 100 m from the edge of 23 deciduous forest patches near Ottawa, Ontario. Land uses surrounding each patch were measured within increasingly large circles from 200 to 5000 m radius around the bird survey site. Regression results suggest that effects of urban and agricultural land uses on forest birds (1) are not uniformly positive or negative, (2) can occur at different spatial scales, and (3) differentially affect certain groups of species. In general, agriculture appeared to affect species at a broad spatial scale (within 5 km), while urban land use had an impact at both a narrower spatial scale (within 1.8 km) and at the broad scale. Neotropical and short distance migrant birds seemed to be the most sensitive to land use intensification within the matrix. Limiting urban land use within approximately 200–1800 m of forest patches would be beneficial for Neotropical migrant birds, which are species of growing conservation concern in temperate North America.     

Landscape fragmentation,land-use legacy and propagule pressure promote plant invasion on coastal dunes: a patch-based approach

Coastal dunes and sand areas are reported to be among the habitats most invaded by alien species in Europe. Landscape pattern could be a significant driver in invasion processes in parallel with land-use legacy. Fragmentation of natural habitats combined with the availability of propagules from the surrounding matrix may enhance the invisibility of ecological communities. Based on multitemporal land cover maps (1954–2008) and a floristic database, we analyzed how habitat fragmentation, propagule pressure and land-use legacy have affected alien plants’ presence and richness on natural dune patches along the Lazio Coast (Central Italy). Floristic data were derived from an existing geo-database of random vegetation plots (64 m²). A set of landscape patch-based metrics, considered to be adequate proxies of the main processes affecting alien invasion and richness, was calculated. First, we fit a generalized linear model (GLM) with binomial errors to assess which landscape metrics are influencing patch invasion. Second, we extracted invaded patches and, with GLMs, we investigated how landscape metrics affect average alien species richness. Alien invasion and alien richness seem to be affected by different processes: although alien invasion of each patch is strongly associated with its land-use legacy, the richness of aliens is more affected by landscape fragmentation and by the propagule pressure to which patch is exposed. By integrating spatial and temporal landscape metrics with floristic data, we were able to disentangle the relations of landscape fragmentation, propagule pressure and land-use legacy with the presence and richness of alien plants. The methodological approach here adopted could be easily extended to other alien species and ecosystems, offering scientifically sound support to prevent the high economic costs derived from both the control and the eradication of aliens.     

Additive partitioning of plant species diversity in an agricultural mosaic landscape

In this paper, we quantify the effects of habitat variability and habitat heterogeneity based on the partitioning of landscape species diversity into additive components and link them to patch-specific diversity. The approach is illustrated with a case study from central Switzerland, where we recorded the presence of vascular plant species in a stratified random sample of 1'280 quadrats of 1 m² within a total area of 0.23 km². We derived components of within- and between-community diversity at four scale levels (quadrat, patch, habitat type, and landscape) for three diversity measures (species richness, Shannon index, and Simpson diversity). The model implies that what we measure as within-community diversity at a higher scale level is the combined effect of heterogeneity at various lower levels. The results suggest that the proportions of the individual diversity components depend on the habitat type and on the chosen diversity aspect. One habitat type may be more diverse than another at patch level, but less diverse at the level of habitat type. Landscape composition apparently is a key factor for explaining landscape species richness, but affects evenness only little. Before we can test the effect of landscape structure on landscape species richness, several problems will have to be solved. These include the incorporation of neighbourhood effects, the unbiased estimation of species richness components, and the quantification of the contribution of a landscape element to landscape species richness.     

Combined effects of area,connectivity, history and structural heterogeneity of woodlands on the species richness of hoverflies (Diptera: Syrphidae)

Context

Hoverflies are often used as bio-indicators for ecosystem conservation, but only few studies have actually investigated the key factors explaining their richness in woodlands.

Objectives

In a fragmented landscape in southwest France, we investigated the joint effects of woodland area, structural heterogeneity, connectivity and history on the species richness of forest-specialist hoverflies, and whether there was a time lag in the response of hoverflies to habitat changes, and tested the effect of spatiotemporal changes.

Methods

Current species richness was sampled in 48 woodlands using 99 Malaise traps. Structural variables were derived from a rapid habitat assessment protocol. Old maps and aerial photographs were used to extract past and present spatial patterns of the woodlands since 1850. Relationships between species richness and explanatory variables were explored using generalized linear models.

Results

We show that current habitat area, connectivity, historical continuity and the average density of tree-microhabitats explained 35 % of variation in species richness. Species richness was affected differently by changes in patch area between 1979 and 2010, depending on woodland connectivity. In isolated woodlands, extinction debt and colonization credit were revealed, showing that even several decades are not sufficient for hoverflies to adapt to landscape-scale habitat conditions.

Conclusions

These findings emphasise the importance of maintaining connectedness between woodlands, which facilitates the dispersion in a changing landscape. Our results also highlight the benefits of using a change-oriented approach to explain the current distribution patterns of species, especially when several spatial processes act jointly.