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.

Selection of multiple umbrella species for functional and taxonomic diversity to represent urban biodiversity.

Surrogates, such as umbrella species, are commonly used to reduce the complexity of quantifying biodiversity for conservation purposes. The presence of umbrella species is often indicative of high taxonomic diversity; however, functional diversity is now recognized as an important metric for biodiversity and thus should be considered when choosing umbrella species. We identified umbrella species associated with high taxonomic and functional biodiversity in urban areas in Switzerland. We analyzed 39,752 individuals of 574 animal species from 96 study plots and 1397 presences of 262 plant species from 58 plots. Thirty-one biodiversity measures of 7 taxonomic groups (plants, spiders, bees, ground beetles, lady bugs, weevils and birds) were included in within- and across-taxa analyses. Sixteen measures were taxonomical (species richness and species diversity), whereas 15 were functional (species traits including mobility, resource use, and reproduction). We used indicator value analysis to identify umbrella species associated with single or multiple biodiversity measures. Many umbrella species were indicators of high biodiversity within their own taxonomic group (from 33.3% in weevils to 93.8% in birds), to a lesser extent they were indicators across taxa. Principal component analysis revealed that umbrella species for multiple measures of biodiversity represented different aspects of biodiversity, especially with respect to measures of taxonomic and functional diversity. Thus, even umbrella species for multiple measures of biodiversity were complementary in the biodiversity aspects they represented. Thus, the choice of umbrella species based solely on taxonomic diversity is questionable and may not represent biodiversity comprehensively. Our results suggest that, depending on conservation priorities, managers should choose multiple and complementary umbrella species to assess the state of biodiversity.

Spider, bee, and bird communities in cities are shaped by environmental control and high stochasticity.

Spatially organized distribution patterns of species and communities are shaped by both autogenic processes (neutral mechanism theory) and exogenous processes (niche theory). In the latter, environmental variables that are themselves spatially organized induce spatial structure in the response variables. The relative importance of these processes has not yet been investigated in urban habitats. We compared the variance explained by purely spatial, spatially structured environmental, and purely environmental components for the community composition of spiders (Araneae), bees (Apidae), and birds (Aves) at 96 locations in three Swiss cities. Environmental variables (topography, climate, land cover, urban green management) were measured on four different radii around sampling points (< 10 m, 50 m, 250 m, 1000 m), while Moran's eigenvector maps (MEMs) acted as spatial variables. All three taxonomic groups showed weak spatial structure. Spider communities reacted to very fine-scaled environmental changes of lawn and meadow management and climate. Bird community composition was determined by woody plants as well as solar radiation at all radii, the scale of the influence varying among species. Bee communities were weakly explained by isolated variables only. Our results suggest that the anthropogenic structuring of urban areas has disrupted the spatial organization of environmental variables and inhibited the development of biotic spatial processes. The near absence of spatial structure may therefore be a feature typical of urban species assemblages, resulting in urban community composition mainly influenced by local environmental variables. Urban environments represent a close-knit mosaic of habitats that are regularly disturbed. Species communities in urban areas are far from equilibrium. Our analysis also suggests that urban communities need to be considered as being in constant change to adapt to disturbances and changes imposed by human activities.

Hearing and hunting in red bats (Lasiurus borealis, Vespertilionidae): audiogram and ear properties.

We examined aspects of hearing in the red bat (Lasiurus borealis) related to its use of biosonar. Evoked potential audiograms, obtained from volume-conducted auditory brainstem responses, were obtained in two bats, and the sound pressure transformation of the pinna was measured in three specimens. Field-recorded echolocation signals were analysed for comparison. The fundamental sonar search calls sweep from 45 to 30 kHz (peak energy at 35 kHz), approach-phase calls sweep from 65 to 35 kHz (peak 40 kHz) and terminal calls sweep from 70 to 30 kHz (peak 45 kHz). The most sensitive region of the audiogram extended from 10 kHz to 45-55 kHz, with maximum sensitivity as low as 20 dB SPL occurring between 25 and 30 kHz. A relative threshold minimum occurred between 40 and 50 kHz. With increasing frequency, the acoustic axis of the pinna moves upwards and medially. The sound pressure transformation was noteworthy near 40-45 kHz; the acoustic axis was closest to the midline, the -3 dB acceptance angles showed local minima, and the pinna gain and interaural intensity difference were maximal. These results are related to the known echolocation and foraging behavior of this species and match the spectral components of approach- and final-phase calls. We conclude that co-evolution with hearing prey has put a higher selective pressure on optimizing localization and tracking of prey than on improving detection performance.

What ears do for bats: a comparative study of pinna sound pressure transformation in chiroptera.

Using a moveable loudspeaker and an implanted microphone, we studied the sound pressure transformation of the external ears of 47 species of bats from 13 families. We compared pinna gain, directionality of hearing and interaural intensity differences (IID) in echolocating and non-echolocating bats, in species using different echolocation strategies and in species that depend upon prey-generated sounds to locate their targets. In the Pteropodidae, two echolocating species had slightly higher directionality than a non-echolocating species. The ears of phyllostomid and vespertilionid species showed moderate directionality. In the Mormoopidae, the ear directionality of Pteronotus parnellii clearly matched the dominant spectral component of its echolocation calls, unlike the situation in three other species. Species in the Emballonuridae, Molossidae, Rhinopomatidae and two vespertilionids that use narrow-band search-phase echolocation calls showed increasingly sharp tuning of the pinna to the main frequency of their signals. Similar tuning was most evident in Hipposideridae and Rhinolophidae, species specialized for flutter detection via Doppler-shifted echoes of high-duty-cycle narrow-band signals. The large pinnae of bats that use prey-generated sounds to find their targets supply high sound pressure gain at lower frequencies. Increasing domination of a narrow spectral band in echolocation is reflected in the passive acoustic properties of the external ears (sharper directionality). The importance of IIDs for lateralization and horizontal localization is discussed by comparing the behavioural directional performance of bats with their bioacoustical features.