Habitat connectivity shapes urban arthropod communities: the key role of green roofs.

The installation of green roofs, defined here as rooftops with a shallow soil cover and extensive vegetation, has been proposed as a possible measure to mitigate the loss of green space caused by the steady growth of cities. However, the effectiveness of green roofs in supporting arthropod communities, and the extent to which they facilitate connectivity of these communities within the urban environment is currently largely unknown. We investigated the variation of species community composition (beta diversity) of four arthropod groups with contrasting mobility (Carabidae, Araneae, Curculionidae, and Apidae) on 40 green roofs and 40 extensively managed green sites on the ground in the city of Zurich, Switzerland. With redundancy analysis and variation partitioning, we (1) disentangled the relative importance of local environmental conditions, the surrounding land cover composition, and habitat connectivity on species community composition, (2) searched for specific spatial scales of habitat connectivity for the different arthropod groups, and (3) discussed the ecological and functional value of green roofs in cities. Our study revealed that on green roofs community composition of high-mobility arthropod groups (bees and weevils) were mainly shaped by habitat connectivity, while low-mobility arthropod groups (carabids and spiders) were more influenced by local environmental conditions. A similar but less pronounced pattern was found for ground communities. The high importance of habitat connectivity in shaping high-mobility species community composition indicates that these green roof communities are substantially connected by the frequent exchange of individuals among surrounding green roofs. On the other hand, low-mobility species communities on green roofs are more likely connected to ground sites than to other green roofs. The integration of green roofs in urban spatial planning strategies has great potential to enable higher connectivity among green spaces, so that eventually even communities of low-mobility species become connected. Furthermore, improving the design of green roofs (composition and configuration of vegetation and soil types) could enhance the ecological value, particularly for low-mobility species.

Assessing habitat connectivity for ground-dwelling animals in an urban environment.

To ensure viable species populations in fragmented landscapes, individuals must be able to move between suitable habitat patches. Despite the increased interest in biodiversity assessment in urban environments, the ecological relevance of habitat connectivity in highly fragmented landscapes remains largely unknown. The first step to understanding the role of habitat connectivity in urban ecology is the challenging task of assessing connectivity in the complex patchwork of contrasting habitats that is found in cities. We developed a data-based framework, minimizing the use of subjective assumptions, to assess habitat connectivity that consists of the following sequential steps: (1) identification of habitat preference based on empirical habitat-use data; (2) derivation of habitat resistance surfaces evaluating various transformation functions; (3) modeling of different connectivity maps with electrical circuit theory (Circuitscape), a method considering all possible pathways across the landscape simultaneously; and (4) identification of the best connectivity map with information-theoretic model selection. We applied this analytical framework to assess habitat connectivity for the European hedgehog Erinaceus europaeus, a model species for ground-dwelling animals, in the city of Zurich, Switzerland, using GPS track points from 40 individuals. The best model revealed spatially explicit connectivity "pinch points," as well as multiple habitat connections. Cross-validation indicated the general validity of the selected connectivity model. The results show that both habitat connectivity and habitat quality affect the movement of urban hedgehogs (relative importance of the two variables was 19.2% and 80.8%, respectively), and are thus both relevant for predicting urban animal movements. Our study demonstrates that even in the complex habitat patchwork of cities, habitat connectivity plays a major role for ground-dwelling animal movement. Data-based habitat connectivity maps can thus serve as an important tool for city planners to identify habitat corridors and plan appropriate management and conservation measures for urban animals. The analytical framework we describe to model such connectivity maps is generally applicable to different types of habitat-use data and can be adapted to the movement scale of the focal species. It also allows evaluation of the impact of future landscape changes or management scenarios on habitat connectivity in urban landscapes.

Evaluating contemporary pollen dispersal in two common grassland species Ranunculus bulbosus L. (Ranunculaceae) and Trifolium montanum L. (Fabaceae) using an experimental approach.

Pollen flow is a key biological process that connects plant populations, preventing genetic impoverishment and inbreeding. Pollen-mediated long-distance dispersal (LDD) events are especially important for plant species in increasingly fragmented landscapes. Patterns of pollen dispersal were directly estimated and dispersal kernels modelled in an experimental population of Ranunculus bulbosus and Trifolium montanum to determine the potential for LDD. Eight and 11 microsatellite markers were used for R. bulbosus and T. montanum, respectively, to run a likelihood-based paternity analysis on randomly chosen offspring (Ntotal = 180 per species) from five maternal plants. High rates of selfing were found in R. bulbosus (average 45.7%), while no selfing was observed in T. montanum. The majority (60%) of mating events occurred at very short distances: the median of the observed dispersal distances was 0.8 m in both species, and the average distances were 15.9 and 10.3 m in R. bulbosus and T. montanum, respectively. Modelling the pollen dispersal kernel with four different distribution functions (exponential-power, geometric, 2Dt and Weibull) indicated that the best fit for both species was given by a Weibull function. Yet, the tail of the T. montanum pollen dispersal kernel was thinner than in R. bulbosus, suggesting that the probability for LDD is higher in the latter species. Even though the majority of pollen dispersal occurred across short distances, the detection of several mating events up to 362 m (R. bulbosus) and 324 m (T. montanum) suggests that pollen flow may be sufficient to ensure population connectivity in these herb species across fragmented grasslands in Swiss agricultural landscapes.

Does long-distance pollen dispersal preclude inbreeding in tropical trees? Fragmentation genetics of Dysoxylum malabaricum in an agro-forest landscape.

Tropical trees often display long-distance pollen dispersal, even in highly fragmented landscapes. Understanding how patterns of spatial isolation influence pollen dispersal and interact with background patterns of fine-scale spatial genetic structure (FSGS) is critical for evaluating the genetic consequences of habitat fragmentation. In the endangered tropical timber tree Dysoxylum malabaricum (Meliaceae), we apply eleven microsatellite markers with paternity and parentage analysis to directly estimate historic gene flow and contemporary pollen dispersal across a large area (216 km(2)) in a highly fragmented agro-forest landscape. A comparison of genetic diversity and genetic structure in adult and juvenile life stages indicates an increase in differentiation and FSGS over time. Paternity analysis and parentage analysis demonstrate high genetic connectivity across the landscape by pollen dispersal. A comparison between mother trees in forest patches with low and high densities of adult trees shows that the frequency of short-distance mating increases, as does average kinship among mates in low-density stands. This indicates that there are potentially negative genetic consequences of low population density associated with forest fragmentation. Single isolated trees, in contrast, frequently receive heterogeneous pollen from distances exceeding 5 km. We discuss the processes leading to the observed patterns of pollen dispersal and the implications of this for conservation management of D. malabaricum and tropical trees more generally.

Forest fragmentation genetics in a formerly widespread island endemic tree: Vateriopsis seychellarum (Dipterocarpaceae).

Habitat fragmentation and changed land use have seriously reduced population size in many tropical forest tree species. Formerly widespread species with limited gene flow may be particularly vulnerable to the negative genetic effects of forest fragmentation and small population size. Vateriopsis seychellarum (Dipterocarpaceae) is a formerly widespread canopy tree of the Seychelles, but is now reduced to 132 adult individuals distributed in eleven sites. Using ten microsatellite loci, a genetic inventory of all adult trees and a sample of 317 progeny, we demonstrate that despite its restricted range, overall genetic diversity was relatively high (H(E) : 0.56). The juvenile cohort, however, had significantly lower allelic richness (adults R(S) : 3.91; juveniles R(S) : 2.83) and observed heterozygosity than adult trees (adults H(O) : 0.62; juveniles H(O) : 0.48). Rare alleles were fewer and kinship between individuals was stronger in juveniles. Significant fine-scale spatial genetic structure was observed in remnant adults, and parentage analysis indicated that more than 90% of sampled progeny disperse <25 m and pollen dispersed <50 m. The molecular data confirmed that two populations were derived entirely from self-fertilized offspring from a single surviving mother tree. These populations produce viable offspring. Despite this extreme genetic bottleneck, self-compatibility may provide V. seychellarum with some resistance to the genetic consequences of habitat fragmentation, at least in the short term. We discuss our findings in the context of other rare and threatened dipterocarp species which are vulnerable to miss-management of genetic resources and population fragmentation.

Back from the brink: potential for genetic rescue in a critically endangered tree.

Rare plant species are vulnerable to genetic erosion and inbreeding associated with small population size and isolation due to increasing habitat fragmentation. The degree to which these problems undermine population viability remains debated. We explore genetic and reproductive processes in the critically endangered long-lived tropical tree Medusagyne oppositifolia, an endemic to the Seychelles with a naturally patchy distribution. This species is failing to recruit in three of its four populations. We evaluate whether recruitment failure is linked to genetic problems associated with fragmentation, and if genetic rescue can mitigate such problems. Medusagyne oppositifolia comprises 90 extant trees in four populations, with only the largest (78 trees) having successful recruitment. Using 10 microsatellite loci, we demonstrated that genetic diversity is high (H(E) : 0.48-0.63; H(O) : 0.56-0.78) in three populations, with only the smallest population having relatively low diversity (H(E) : 0.26 and H(O) : 0.30). All populations have unique alleles, high genetic differentiation, and significant within population structure. Pollen and seed dispersal distances were mostly less than 100 m. Individuals in small populations were more related than individuals in the large population, thus inbreeding might explain recruitment failure in small populations. Indeed, inter-population pollination crosses from the large donor population to a small recipient population resulted in higher reproductive success relative to within-population crosses. Our study highlights the importance of maintaining gene flow between populations even in species that have naturally patchy distributions. We demonstrate the potential for genetic and ecological rescue to support conservation of plant species with limited gene flow.

Insects in fragmented forests: a functional approach.

Insects are highly susceptible to the adverse effects of forest fragmentation. It is now beyond any doubt that fragmentation-induced changes in abundance and species richness occur in many insect groups. However, the study of insects in fragmented forests is still in its infancy and lacks real direction. Simple empirical studies are not answering the questions we most want to answer about fragmented systems. Are we in the midst of a mass-extinction crisis? What is the functional significance of the immense insect biodiversity? Does biodiversity loss affect ecosystem functioning? A more focused, functional approach to the study of forest fragmentation is required to move beyond the description of pattern and to determine how changes in insect communities affect ecosystem processes in fragmented forests.