Effects of increasing landscape heterogeneity on local plant species richness: how much is enough?

Contemporary landscape ecology continues to explore the causes and consequences of landscape heterogeneity across a range of scales, and demands for the scientific underpinnings of landscape planning and management still remains high. The spatial distribution of resources can be a key element in determining habitat quality, and that in turn is directly related to the level of heterogeneity in the system. In this sense, forest habitat mosaics may be more affected by lack of heterogeneity than by structural fragmentation. Nonetheless, increasing spatial heterogeneity at a given spatial scale can also decrease habitat patch size, with potential negative consequences for specialist species. Such dual effect may lead to hump-backed shape relationships between species diversity and heterogeneity, leading to three related assumptions: (i) at low levels of heterogeneity, an increase in heterogeneity favours local and regional species richness, (ii) there is an optimum heterogeneity level at which a maximum number of species is reached, (iii) further increase in spatial heterogeneity has a negative effect on local and regional species richness, due to increasing adverse effects of habitat fragmentation. In this study, we investigated the existence of a hump-shaped relationship between local plant species richness and increasing forest landscape heterogeneity on a complex mosaic in the French Alps. Forest landscape heterogeneity was quantified with five independent criteria. We found significant quadratic relationships between local forest species richness and two heterogeneity criteria indicators, showing a slight decrease of forest species richness at very high heterogeneity levels. Species richness–landscape heterogeneity relationships varied according to the heterogeneity metrics involved and the type of species richness considered. Our results support the assumption that intermediate levels of heterogeneity may support more species than very high levels of heterogeneity, although we were not able to conclude for a systematic negative effect of very high levels of heterogeneity on local plant species richness.     

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.     

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.     

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.     

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.     

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.     

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.     

Inconsistent effects of landscape heterogeneity and land-use on animal diversity in an agricultural mosaic: a multi-scale and multi-taxon investigation

Context

The landscape heterogeneity hypothesis states that increased heterogeneity in agricultural landscapes will promote biodiversity. However, this hypothesis does not detail which components of landscape heterogeneity (compositional or configurational) most affect biodiversity and how these compare to the effects of surrounding agricultural land-use.

Objectives

Our objectives were to: (1) assess the influence of the components of structural landscape heterogeneity on taxonomic diversity; and (2) compare the effects of landscape heterogeneity to those of different types of agricultural land-use in the same landscape across different taxonomic groups.

Methods

We identified a priori independent gradients of compositional and configurational landscape heterogeneity within an agricultural mosaic of north-eastern Swaziland. We tested how bird, dung beetle, ant and meso-carnivore richness and diversity responded to compositional and configurational heterogeneity and agricultural land-use across five different spatial scales.

Results

Compositional heterogeneity best explained species richness in each taxonomic group. Bird and ant richness were both positively correlated with compositional heterogeneity, whilst dung beetle richness was negatively correlated. Commercial agriculture positively influenced bird species richness and ant diversity, but had a negative influence on dung beetle richness. There was no effect of either component of heterogeneity on the combined taxonomic diversity or richness at any spatial scale.

Conclusions

Our results suggest that increasing landscape compositional heterogeneity and limiting the negative effects of intensive commercial agriculture will foster diversity across a greater number of taxonomic groups in agricultural mosaics. This will require the implementation of different strategies across landscapes to balance the contrasting influences of compositional heterogeneity and land-use. Strategies that couple large patches of core habitat across broader scales with landscape structural heterogeneity at finer scales could best benefit biodiversity.

     

Does the response of bird assemblages to fire mosaic properties vary among spatial scales and foraging guilds?

Context

An increasing number of studies have investigated the impact of environmental heterogeneity on faunal assemblages when measured at multiple spatial scales. Few studies, however, have considered how the effects of heterogeneity on fauna vary with the spatial scale at which the response variable is characterised.

Objectives

We investigated the relationship between landscape properties in a region characterised by diverse fire mosaics, and the structure and composition of avian assemblages measured at both the site- (1 ha) and landscape-scale (100 ha).

Methods

We surveyed birds and calculated spatial landscape properties in sub-tropical woodlands of central Queensland, Australia.

Results

Environmental heterogeneity, as measured by topographic complexity, was consistently important for bird species richness and composition. However, the explanatory power of topographic complexity varied depending on the spatial scale and the component of diversity under investigation. We found different correlates of richness within particular foraging guilds depending on the scale at which richness was measured. Extent of long-unburnt habitat (>10 years since fire) was the most important variable for the landscape-scale richness of frugivores, insectivores and canopy feeders, whereas environmental heterogeneity in the surrounding landscape was more important for site-scale richness of these foraging guilds.

Conclusions

The response of species richness to landscape characteristics varies among scales, and among components of diversity. Thus, depending on the scale at which a biodiversity conservation goal is conceptualised—maximising richness at a site, or across a landscape—different landscape management approaches may be preferred.

     

Species richness of hedgerow habitats in changing agricultural landscapes: are α and γ diversity shaped by the same factors?

Understanding the determinants of hedgerow plant diversity in agricultural landscapes remains a difficult task, because the potential drivers affect the complete range of biodiversity components (alpha to gamma diversity). We surveyed herbaceous plant communities (of a height <1.5 m) in 84 hedgerows in the Seine river floodplain of France. Two types of potential drivers for species richness, accounting for landscape mosaic and hedgerow network, were recorded at both hedgerow and site scale. The distribution of species richness through the components of alpha hedgerow diversity (i.e. the average diversity within a habitat) and gamma hedgerow diversity (i.e. the total diversity across habitats) were assessed using additive partitioning methods, while the relationship between species diversity and its potential landscape drivers at both scales was modeled using Generalized Additive Models. Our results indicated that gamma hedgerow diversity is explained by the heterogeneity of the landscape structure, which is correlated with the mosaic of agricultural land use. At this scale, intrinsic properties of the configuration of the hedgerow networks have a weak influence on species richness. Alpha hedgerow diversity is also explained by landscape variables, accounting for both the configuration of agricultural mosaics and hedgerow networks, but to a lesser extent. Time lags for species responses are shown at both scales, and for the two types of drivers. Extinction or colonization debt may be indicated at both scales, while the remnant effects of former practices may also be responsible for such patterns at a local scale. We suggest that hedgerow management should take the specific parameters of both scales into account. At a local scale, management actions should aim to decrease the influence of adjacent land use when the impact is negative, through the implementation of extended buffer zones, while at the landscape and farm scales, agri-environmental schemes should be dedicated to the conservation of specific agricultural land uses.     

Habitat area trumps fragmentation effects on arthropods in an experimental landscape system

The effects of habitat area and fragmentation are confounded in many studies. Since a reduction in habitat area alone reduces patch size and increases patch isolation, many studies reporting fragmentation effects may really be documenting habitat-area effects. We designed an experimental landscape system in the field, founded on fractal neutral landscape models, to study arthropod community responses to clover habitat in which we adjusted the level of fragmentation independently of habitat area. Overall, habitat area had a greater and more consistent effect on morphospecies richness than fragmentation. Morphospecies richness doubled between 10 and 80% habitat, with the greatest increase occurring up to 40% habitat. Fragmentation had a more subtle and transient effect, exhibiting an interaction at intermediate levels of habitat only at the start of the study or in the early-season (June) survey. In these early surveys, morphospecies richness was higher in clumped 40–50% landscapes but higher in fragmented landscapes at 60–80% habitat. Rare or uncommon species are expected to be most sensitive to fragmentation effects, and we found a significant interaction with fragmentation at intermediate levels of habitat for these types of morphospecies in early surveys. Although the effects of fragmentation are expected to amplify at higher trophic levels, all trophic levels exhibited a significant fragmentation effect at intermediate levels of habitat in these early surveys. Predators/parasitoids were more sensitive to habitat area than herbivores, however. Thus, our results confirm that habitat area is more important than fragmentation for predicting arthropod community responses, at least in this agricultural system.     

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.     

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.

     

Environmental factors at different spatial scales governing soil fauna community patterns in fragmented forests

Spatial and temporal changes in community structure of soil organisms may result from a myriad of processes operating at a hierarchy of spatial scales, from small-scale habitat conditions to species movements among patches and large-sale landscape features. To disentangle the relative importance of spatial and environmental factors at different scales (plot, patch and landscape), we analyzed changes in Collembola community structure along a gradient of forest fragmentation, testing predictions of the Hierarchical Patch Dynamics Paradigm (HPDP) in different European biogeographic regions (Boreal, Continental, Atlantic, Mediterranean, Alpine). Using variance partitioning methods, based on partial CCAs, we observed that the independent effect of environmental processes was significantly explaining Collembola community variance in all regions, while the relative effect of spatial variables was not significant, due to the observed high levels of landscape heterogeneity along the gradient. Environmental factors at the patch and plot scales were generally significant and explained the larger part of community changes. Landscape variables were not significant across all study sites. Yet, at the landscape level, an increase in forest habitat and proximity of forest patches were showed to have an indirect influence on local community changes, by influencing microhabitat heterogeneity at lower spatial scales in all studied regions. In line with HPDP, large-scale landscape features influenced spatio-temporal changes in soil fauna communities by constraining small-scale environmental processes. In turn, these provided mechanistic understanding for diversity patterns operating at the patch scale, via shifts in community weighted mean of Collembola life-forms occurring in local communities along the fragmentation gradient.     

Assessing habitat fragmentation on marine epifaunal macroinvertebrate communities: an experimental approach

Habitat fragmentation is considered a major cause of biodiversity loss, both on terrestrial and marine environments. Understanding the effects of habitat fragmentation on the structure and dynamics of natural communities is extremely important to support management actions for biodiversity conservation. However, the effects of habitat fragmentation on marine communities are still poorly understood. Here we evaluated whether habitat fragmentation affects the structure of epifaunal communities in the sublittoral zone, in the northern coast of São Paulo state, Brazil. Five experimental landscapes were constructed, each one forming a large continuous patch. After 4 weeks, each landscape was cut on three patches of different sizes. Epifaunal macroinvertebrate communities were sampled at the edge and interior of experimental landscapes before manipulation to evaluate edge effects. After four more weeks, communities from the three patch sizes were also sampled to evaluate patch size effects. We compared the diversity of communities at different levels of fragmentation by total abundance, rarefied taxon richness, Shannon–Wiener diversity index, Simpson’s dominance index, and abundance of dominant taxa. Higher taxon richness and gastropod abundance were recorded in the patch edges, but no significant differences were found among patch sizes. We found a significant effect of habitat fragmentation, with lower abundances of Gammaridea (the dominant taxon), Ophyuroidea, and Pycnogonida after the experimental fragmentation. Lower abundances of dominant taxa resulted in higher diversity and lower dominance in fragmented landscapes when compared to integral, pre-manipulation landscapes. Our results suggest that fragmentation of landscapes in the system studied can reduce dominance, and that even small patch sizes can be important for the conservation of macroinvertebrate diversity.     

Comparing the role of site disturbance and landscape properties on understory species richness in fragmented periurban Mediterranean forests

We hypothesized that the spatial configuration and dynamics of periurban forest patches in Barcelona (NE of Spain) played a minor role in determining plant species richness and assemblage compared to site conditions, and particularly to both direct (measured at plot level) and potential (inferred from landscape metrics) human-associated site disturbance. The presence of all understory vascular plants was recorded on 252 plots of 100 m² randomly selected within forest patches ranging in size from 0.25 ha to 218 ha. Species were divided into 6 groups, according to their ecology and conservation status. Site condition was assessed at plot level and included physical attributes, human-induced disturbance and Quercus spp. tree cover. Landscape structure and dynamics were assessed from patch metrics and patch history. We also calculated a set of landscape metrics related to potential human accessibility to forests. Results of multiple linear regressions indicated that the variance explained for non-forest species groups was higher than for forest species richness. Most of the main correlates corresponded to site disturbance variables related to direct human alteration, or to landscape variables associated to indirect human effects on forests: Quercus tree cover (a proxy for successional status) was the most important correlate of non-forest species richness, which decreased when Quercus tree cover increased. Human-induced disturbance was an important correlate of synanthropic and total species richness, which were higher in recently managed and in highly frequented forests. Potential human accessibility also affected the richness of most species groups. In contrast, patch size, patch shape and connectivity played a minor role, as did patch history. We conclude that human influence on species richness in periurban forests takes place on a small scale, whereas large-scale effects attributable to landscape structure and fragmentation are comparatively less important. Implications of these results for the conservation of plant species in periurban forests are discussed.     

Interactive effects of area and connectivity on the diversity of tachinid parasitoids in highly fragmented landscapes

Although many empirical and theoretical studies have elucidated the effects of habitat fragmentation on the third trophic level, little attention has been paid to the impacts of this driver on more generalist groups of non-hymenopteran parasitoids. Here, we used the highly-diverse group of tachinid flies as an alternative model to test the effects of landscape fragmentation on insect parasitoids. Our aims were: (i) to evaluate the relative importance of habitat area and connectivity losses and their potential interaction on tachinid diversity, (ii) to test whether the effects of habitat fragmentation changes seasonally, and (iii) to further assess the effect of habitat diversity on tachinid diversity and whether different parasitoid-host associations modify the species richness response to fragmentation. In 2012 a pan-trap sampling was conducted in 18 semi-natural grasslands embedded in intensive agricultural landscapes along statistically orthogonal gradients of habitat area, connectivity and habitat diversity. We found an interaction between habitat area and connectivity indicating that tachinid abundance and species richness were more negatively affected by habitat loss in landscapes with low rather than with relatively large habitat connectivity. Although tachinid communities exhibited large within-year species turnover, we found that the effects of landscape fragmentation did not change seasonally. We found that habitat diversity and host association did not affect tachinid species diversity. Our results have important implications for biodiversity conservation as any attempts to mitigate the negative effects of habitat loss need to take the general level of habitat connectivity in the landscape into account.     

Species richness and composition differ in response to landscape and biogeography

Context

Understanding how landscape patterns affect species diversity is of great importance in the fields of biogeography, landscape ecology and conservation planning, but despite the rapid advance in biodiversity analysis, investigations of spatial effects on biodiversity are still largely focused on species richness.

Objectives

We wanted to know if and how species richness and species composition are differentially driven by the spatial measures dominating studies in landscape ecology and biogeography. As both measures require the same limited presence/absence information, it is important to choose an appropriate diversity measure, as differing results could have important consequences for interpreting ecological processes.

Methods

We recorded plant occurrences on 112 islands in the Baltic archipelago. Species richness and composition were calculated for each island, and the explanatory power of island area and habitat heterogeneity, distance to mainland and structural connectivity at three different landscape sizes were examined.

Results

A total of 354 different plant species were recorded. The influence of landscape variables differed depending on which diversity measure was used. Island area and structural connectivity determined plant species richness, while species composition revealed a more complex pattern, being influenced by island area, habitat heterogeneity and structural connectivity.

Conclusions

Although both measures require the same basic input data, species composition can reveal more about the ecological processes affecting plant communities in fragmented landscapes than species richness alone. Therefore, we recommend that species community composition should be used as an additional standard measure of diversity for biogeography, landscape ecology and conservation planning.

     

Trait-dependent responses of flower-visiting insects to distance to semi-natural grasslands and landscape heterogeneity

Protecting semi-natural grasslands may through spill-over benefit species richness and abundance of flower-visiting insects in linear habitats, such as uncultivated field boundaries, in agricultural landscapes. However, whether local diversity increases both with decreasing distance from potential source habitats and increasing landscape heterogeneity is poorly known due to a general lack of studies replicated at the landscape scale. We analysed if local assemblages of bumblebees, butterflies and hoverflies in linear uncultivated habitats increased with increasing distance to the nearest semi-natural grassland in 12 replicated landscapes along a gradient of landscape heterogeneity in Scania, Southern Sweden. Species richness and abundance of bumblebees and butterflies, but not hoverflies, decreased with increasing distance to semi-natural grasslands, but none of these groups were related to increasing landscape heterogeneity. Further analyses on trait-specific groups revealed significant decreases in the abundance of sedentary and grassland specialist butterflies with increasing distance to assumed source populations, whereas this was not the case concerning mobile species and grassland generalists. The abundance of all bumblebee trait groups decreased with increasing distance to semi-natural grasslands, but only some species (those nesting above ground, with long colony cycles and with small colony sizes) also increased with increasing landscape heterogeneity. We conclude that local species assemblages of flower-visiting insects in linear habitat elements were mainly affected by the occurrence of nearby semi-natural grasslands. In order to conserve diverse assemblages of flower-visiting insects, including the ecological services they provide, it is important to conserve semi-natural grasslands dispersed throughout agricultural landscapes.     

Factors Associated with Grassland Bird Species Richness: The Relative Roles of Grassland Area, Landscape Structure, and Prey

The factors responsible for widespread declines of grassland birds in the United States are not well understood. This study, conducted in the short-grass prairie of eastern Wyoming, was designed to investigate the relationship between variation in habitat amount, landscape heterogeneity, prey resources, and spatial variation in grassland bird species richness. We estimated bird richness over a 5-year period (1994–1998) from 29 Breeding Bird Survey locations. Estimated bird richness was modeled as a function of landscape structure surrounding survey routes using satellite-based imagery (1996) and grasshopper density and richness, a potentially important prey of grassland birds. Model specification progressed from simple to complex explanations for spatial variation in bird richness. An information-theoretic approach was used to rank and select candidate models. Our best model included measurements of habitat amount, habitat arrangement, landscape matrix, and prey diversity. Grassland bird richness was positively associated with grassland habitat; was negatively associated with habitat dispersion; positively associated with edge habitats; negatively associated with landscape matrix attributes that may restrict movement of grassland bird; and positively related to grasshopper richness. Collectively, 62% of the spatial variation in grassland bird richness was accounted for by the model (adj-R² = 0.514). These results suggest that the distribution of grassland bird species is influenced by a complex mixture of factors that include habitat area affects, landscape pattern and composition, and the availability of prey.