Scales and costs of habitat selection in heterogeneous landscapes

Summary Two scales of habitat selection are likely to influence patterns of animal density in heterogeneous landscapes. At one scale, habitat selection is determined by the differential use of foraging locations within a home range. At a larger scale, habitat selection is determined by dispersal and the ability to relocate the home range. The limits of both scales must be known for accurate assessments of habitat selection and its role in effecting spatial patterns in abundance. Isodars, which specify the relationships between population density in two habitats such that the expected reproductive success of an individual is the same in both, allow us to distinguish the two scales of habitat selection because each scale has different costs. In a two-habitat environment, the cost of rejecting one of the habitats within a home range can be expressed as a devaluation of the other, because, for example, fine-grained foragers must travel through both. At the dispersal scale, the cost of accepting a new home range in a different habitat has the opposite effect of inflating the value of the original habitat to compensate for lost evolutionary potential associated with relocating the home range. These costs produce isodars at the foraging scale with a lower intercept and slope than those at the dispersal scale.Empirical data on deer mice occupying prairie and badland habitats in southern Alberta confirm the ability of isodar analysis to differentiate between foraging and dispersal scales. The data suggest a foraging range of approximately 60 m, and an effective dispersal distance near 140 m. The relatively short dispersal distance implies that recent theories may have over-emphasized the role of habitat selection on local population dynamics. But the exchange of individuals between habitats sharing irregular borders may be substantial. Dispersal distance may thus give a false impression of the inability of habitat selection to help regulate population density.     

Ecosystem Linkages Between Lakes and the Surrounding Terrestrial Landscape in Northeast Iceland

Despite a recent emphasis on understanding cross-habitat interactions, few studies have examined the ecological linkages between lakes and surrounding terrestrial habitats. The current paradigm of land–lake interactions is typically unidirectional: the view is that nutrients and matter are transported downslope from the surrounding watershed to their ultimate lacustrine destination. Emergent aquatic insects, which spend their larval stages in lake sediments and emerge as adults to mate over land, can act as vectors of material, energy and nutrients from aquatic to terrestrial habitats. In this study, we document a gradient of midge (Diptera: Chironomidae) infall rates into terrestrial habitats (measured as g dw midges m^?2 d^?1) surrounding eight lakes in Northern Iceland (≈66°N latitude). Lakes ranged from having virtually no midge infall (for example, Helluvaðstjörn, 0.03 g m^?2 d^?1) to extreme levels (for example, Mývatn, 19 g m^?2 d^?1) with abundances of midges decreasing logarithmically with distance from shore. Annual midge input rates are estimated as high as 1200–2500 kg midges ha^?1 y^?1. As midges are approximately 9.2% total N, this can result in a significant fertilization effect of terrestrial habitats with consequences for plant quality and community structure. In addition, we used naturally-occurring δ¹³C and δ¹⁵N isotopes to examine food web structure and diet sources of terrestrial arthropod consumers surrounding lakes with differing amounts of midge input. Terrestrial arthropods showed increased utilization of aquatic-derived (that is, midge) C relative to terrestrial sources as midge infall increased. This pattern was particularly pronounced for predators, such as spiders and opiliones, and some detritivores (Collembola). These findings suggest that, despite being largely ignored, aquatic-to-terrestrial linkages can be large and midges can fuel terrestrial communities by directly serving as resources for predators and decomposers.     

Variability between Scales: Predictors of Nomadism in Birds of an Australian Mediterranean-climate Ecosystem

Nomadism in animals is a response to resource distributions that are highly variable in time and space. Using the avian fauna of the Mediterranean-climate region of southcentral Australia, we tested a number of variables to determine if they predicted nomadism. These variables were species body mass, the distance in body mass terms to the edge of a body mass aggregation, and diet (for example, seeds, invertebrates, nectar, or plants). We utilized two different classifications of the avifauna that diverged in their definition of nomadic to build two different predictive models. Using both classifications, distance to the edge of a body mass aggregation was found to be a significant predictor of nomadism. There was also evidence that nomads tend to feed on nectar and tend to be large. The significance of the variables body mass and diet (nectar) may reflect the greater energy requirements of large birds and the inherent variability of nectar as a food source. The significance of the variable distance to the edge of a body mass aggregation provides further evidence of inherent variability in resources between domains of scale. The edges of body mass aggregations are hypothesized to be areas of increased resource variability that reflect the transition from one scale of landscape pattern to another. Received 14 November 2000; accepted 27 June 2001.     

Sunken roads as habitats for forest plant species in a dynamic agricultural landscape: effects of age and isolation

Aim This study tests the hypothesis that linear, woody habitat patches surrounding small, sunken rural roads not only function as an unstable sink but also as a true, sustainable habitat for forest plants. Furthermore, factors affecting the presence of forest plant species in sunken roads are determined. Finally, the implications of these findings for the overall metapopulation dynamics of forest plant species in fragmented agricultural landscapes are assessed. Location The study area, c. 155 km² in size, is situated in a fragmented agricultural landscape within the loamy region of central Belgium. Methods Forest species presence–absence data were collected for 389 sunken roads. The effect of area, depth, age and isolation on sunken road species richness was assessed using linear regression and analysis of variance (anova ). Analysis of covariance was employed to study the interaction between age and isolation. Differences in plant community dispersal spectra in relation to sunken road age and isolation were analysed by means of linear regression and anova . Results Sunken roads proved to function as an important habitat for forest plants. The sink‐hypothesis was falsified by a clear species accumulation in time: sunken road species richness significantly increased with the age of the elements. Sunken road age mainly affected species richness through effects on both area and depth, affecting habitat quality and diversity. Furthermore, sunken road isolation had a significant impact on species richness as well, with the number of forest species decreasing with increasing isolation of the elements, indicating dispersal limitation in sunken road habitats. Moreover, a significant age × isolation interaction effect was demonstrated. Differences in regression slopes for isolation between age classes revealed that the effect of isolation intensified with increasing age of the elements. Differential colonization in relation to forest species dispersal capacities probably account for this, as confirmed by the analysis of sunken road plant community dispersal spectra, with the fraction of species with low dispersal capacities increasing with increasing age and decreasing isolation of the elements. Main conclusions During sunken road development, area and depth increase and, gradually, suitable habitat conditions for forest plant species arise. Depending on their ecological requirements and dispersal capacities, forest species progressively colonize these habitats as a function of the element's isolation. The functioning of sunken roads as a sustainable habitat for forest species enhances the metapopulation viability of forest plants in agricultural landscapes and has important consequences for forest restoration practices. Moreover, the results of this work call for integrating the presence of forest species in small‐scaled linear habitat patches in forest fragmentation studies.     

Frugivores and seed dispersal: mechanisms and consequences for biodiversity of a key ecological interaction

The 5th Symposium on Frugivores and Seed Dispersal, held in Montpellier (France), 13-18 June 2010, brought together more than 220 researchers exemplifying a wide diversity of approaches to the study of frugivory and dispersal of seeds. Following Ted Fleming and Alejandro Estrada's initiative in 1985, this event was a celebration of the 25th anniversary of the first meeting in Veracruz, Mexico. Frugivory and seed dispersal are active research areas that have diversified in multiple directions since 1985 to include evolution (e.g. phylogenetic diversity and dispersal adaptations), physiology (e.g. sensory cues and digestion), landscape ecology (movement patterns), molecular ecology (e.g. gene flow, genetic diversity and structure), community ecology (e.g. mutualistic interaction networks) and conservation biology (effects of hunting, fragmentation, invasion and extinction), among others. This meeting provided an opportunity to assess conceptual and methodological progress, to present ever more sophisticated insights into frugivory in animals and dispersal patterns in plants, and to report the advances made in examining the mechanisms and consequences of seed dispersal for plants and frugivores.     

Laboratory and field-based studies of abundances, small-scale patchiness, and diversity of gymnamoebae in soils of varying porosity and organic content: evidence of microbiocoenoses

Soil samples (varying in granularity) from four natural sites were cultured in microcosms to determine small-scale patchiness in abundance and diversity of gymnamoebae. Eighty grams of the same thoroughly mixed soil, either moistened with distilled water (- nutrients) or supplemented with an equivalent vol. of organically enriched water (+ nutrients), were placed in covered glass jars and incubated for 14 d (25 degrees C). Abundances (number/gram soil) were assessed in each of 3 core samples (5-10 mm apart). Assay precision was estimated to be +/- 4%. Abundances were similar in the 3 closely-spaced samples, but occasional samples had higher abundances, probably representing localized enriched sites ("nutrient hot spots"). Diversity within the triplicate, closely spaced samples varied substantially. Mean abundance and diversity of amoebae were consistently higher in organically enriched soil and in soil of increasing granularity. Field samples collected directly from two of the sites showed similar patterns of abundance and diversity as found in the experimental studies, indicating substantial small-scale compartmentalization of soil protist communities. These data provide evidence of soil eukaryotic microbiocoenoses and indicate that soil microfauna may encounter wide variations in resources and prey communities as they migrate within small distances of several millimeters or less.     

The predatory Chironomidae of an iron-rich stream: feeding ecology and food web structure

Abstract. 1. Three species of Tanypodinae (Chironomidae) were found in an acid and iron-rich stream in southern England. Maximum abundance was achieved in summer and they were sparse at other times. Individuals were aggregated on the stream bed and were overrepresented in accumulations of leaf litter.
2. The diets of all three species consisted of a mixture of prey (prominently detritivorous chironomid larvae) and detritus. More detritus and fewer prey were taken in winter than in summer.
3. When comparing large tanypod species with small and, intraspecifically, late instars with early, the proportion of guts containing prey increased with increasing body size.
4. Stonefly larvae were more prominent in the diet of Zavrelimyia barbatipes (Kieffer) in summer than in winter but for the other two species the reverse was true. A bigger proportion of Trissopelopia longimana (Staeger) guts contained prey in early summer than in August whereas more Macropelopia goetghebueri (Kieffer) guts contained prey in August. This was apparently a consequence of seasonal differences in the distribution of body size among the populations of these two species.
5. The stream contains two further common predators, Plectrocnemia conspersa (Curtis) and Sialis fuliginosa Pict. These are important predators of tanypod larvae but might also compete with them since they severely deplete populations of prey taken in common.
6. Analysis of the food-web in Broadstone Stream reveals remarkably high values of connectance (C and C_max) and of species richness times connectance (SC_max). Such characteristics are theoretically associated with fragile and dynamically unstable food webs, and may be found in 'constant' environments. There is also an apparently unusual prevalence of omnivory in the community.     

Biodiversity and Conservation of Diptera in Heterogeneous Land Mosaics: A Fly's Eye View

Classical landscape ecology views spatial heterogeneity of habitats at relatively large 'human scales', and it is at such scales that most decisions of land management and nature conservation are made. The present paper makes use of a wider range of spatial scales to examine land mosaics from the 'fly point of view'. Taking examples from the Diptera faunas of mountainous land mosaics, it is demonstrated that: (i) large scale, 'patch content' landscape management has a direct bearing on Diptera community structure, (ii) borders between large scale patches are not necessarily perceived by flies (or other insects) in the same way as we perceive them, (iii) border complexity between patches at any scale may be as important as patch content as an axis of habitat definition. In this sense, 'border' is not to be confused with 'edge effects'. It is concluded that attention to both patch content and patch border complexity of land mosaics, viewed at the relevant spatial scales, is necessary for future successful conservation of Diptera biodiversity and for the efficient use of these insects in environmental assessment studies.     

Biodiversity and ecosystem functioning in food webs: the vertical diversity hypothesis

One challenge in merging community and ecosystem ecology is to integrate the complexity of natural multitrophic communities into concepts of ecosystem functioning. Here, we combine food‐web and allometry theories to demonstrate that primary production, as measured by the total nutrient uptake of the multitrophic community, is determined by vertical diversity (i.e. food web's maximum trophic level) and structure (i.e. distributions of species and their abundances and metabolic rates across trophic levels). In natural ecosystems, the community size distribution determines all these vertical patterns and thus the total nutrient uptake. Our model suggests a vertical diversity hypothesis (VDH) for ecosystem functioning in complex food webs. It predicts that, under a given nutrient supply, the total nutrient uptake increases exponentially with the maximum trophic level in the food web and it increases with its maximum body size according to a power law. The VDH highlights the effect of top–down regulation on plant nutrient uptake, which complements traditional paradigms that emphasised the bottom–up effect of nutrient supply on vertical diversity. We conclude that the VDH contributes to a synthetic framework for understanding the relationship between vertical diversity and ecosystem functioning in food webs and predicting the impacts of global changes on multitrophic ecosystems.     

Effects of landscape structure and land-use intensity on similarity of plant and animal communities

Aim Species richness in itself is not always sufficient to evaluate land management strategies for nature conservation. The exchange of species between local communities may be affected by landscape structure and land‐use intensity. Thus, species turnover, and its inverse, community similarity, may be useful measures of landscape integrity from a diversity perspective. Location A European transect from France to Estonia. Methods We measured the similarity of plant, bird, wild bee, true bug, carabid beetle, hoverfly and spider communities sampled along gradients in landscape composition (e.g. total availability of semi‐natural habitat), landscape configuration (e.g. fragmentation) and land‐use intensity (e.g. pesticide loads). Results Total availability of semi‐natural habitats had little effect on community similarity, except for bird communities, which were more homogeneous in more natural landscapes. Bee communities, in contrast, were less similar in landscapes with higher percentages of semi‐natural habitats. Increased landscape fragmentation decreased similarity of true bug communities, while plant communities showed a nonlinear, U‐shaped response. More intense land use, specifically increased pesticide burden, led to a homogenization of bee, bug and spider communities within sites. In these cases, habitat fragmentation interacted with pesticide load. Hoverfly and carabid beetle community similarity was differentially affected by higher pesticide levels: for carabid beetles similarity decreased, while for hoverflies we observed a U‐shaped relationship. Main conclusions Our study demonstrates the effects of landscape composition, configuration and land‐use intensity on the similarity of communities. It indicates reduced exchange of species between communities in landscapes dominated by agricultural activities. Taxonomic groups differed in their responses to environmental drivers and using but one group as an indicator for ‘biodiversity’ as such would thus not be advisable.     

Biodiversity mediates top–down control in eelgrass ecosystems: a global comparative‐experimental approach

Nutrient pollution and reduced grazing each can stimulate algal blooms as shown by numerous experiments. But because experiments rarely incorporate natural variation in environmental factors and biodiversity, conditions determining the relative strength of bottom–up and top–down forcing remain unresolved. We factorially added nutrients and reduced grazing at 15 sites across the range of the marine foundation species eelgrass (Zostera marina) to quantify how top–down and bottom–up control interact with natural gradients in biodiversity and environmental forcing. Experiments confirmed modest top–down control of algae, whereas fertilisation had no general effect. Unexpectedly, grazer and algal biomass were better predicted by cross‐site variation in grazer and eelgrass diversity than by global environmental gradients. Moreover, these large‐scale patterns corresponded strikingly with prior small‐scale experiments. Our results link global and local evidence that biodiversity and top–down control strongly influence functioning of threatened seagrass ecosystems, and suggest that biodiversity is comparably important to global change stressors.     

Industrial Ecosystems and Food Webs: An Expansion and Update of Existing Data for Eco‐Industrial Parks and Understanding the Ecological Food Webs They Wish to Mimic

Cyclical industrial networks are becoming highly desirable for their efficient use of resources and capital. Progress toward this ideal can be enhanced by mimicking the structure of naturally sustainable ecological food webs (FWs). The structures of cyclic industrial networks, sometimes known as eco‐industrial parks (EIPs), are compared to FWs using a variety of important structural ecological parameters. This comparison uses a comprehensive data set of 144 FWs that provides a more ecologically correct understanding of how FWs are organized than previous efforts. In conjunction, an expanded data set of 48 EIPs gives new insights into similarities and differences between the two network types. The new information shows that, at best, current EIPs are most similar to those FWs that lack the components that create a biologically desirable cyclical structure. We propose that FWs collected from 1993 onward should be used in comparisons with EIPs, given that these networks are much more likely to include important network functions that directly affect the structure. We also propose that the metrics used in an ecological analysis of EIPs be calculated from an FW matrix, as opposed to a community matrix, which, to this point, has been widely used. These new insights into the design of ecologically inspired industrial networks clarify the path toward superior material and energy cycling for environmental and financial success.     

Trade-offs between dispersal and competitive ability: a comparative study of wind-dispersed Asteraceae forbs

An underlying assumption in many models of coexistence and species response to fragmentation is the trade-off between dispersal and competitive abilities among species. Despite a well-founded theoretical ground for this assumption, the mechanism itself has not been as thoroughly explored. Empirical studies of the dispersal/competition trade-off have so far mainly concerned the dispersal distance of single offspring, whereas most models where the trade-off is assumed concern dispersal rate, i.e. the number of offspring that is dispersed outside a local patch per time unit. When species differ in fecundity this should also affect the dispersal rate. We therefore investigated different aspects of dispersal ability and competitive ability in the recruitment phase for 15 wind-dispersed Asteraceae species. A trade-off was found between dispersal ability at the offspring level, i.e. distance travelled by single seeds, and competitive ability in the recruitment phase, but no trade-off was found between dispersal ability at the brood level, i.e. the dispersal ability of single seeds in combination with fecundity, and competitive ability in the recruitment phase. The results were supported both by cross-species analysis and analyses by phylogenetically independent contrast. If this is a general pattern then it is troublesome for models making the assumption that there is a trade-off between dispersal rate and competitive ability.     

Geographical and genetic distances among zooplankton populations in a set of interconnected ponds: a plea for using GIS modelling of the effective geographical distance

In systems of interconnected ponds or lakes, the dispersal of zooplankton may be mediated by the active population component, with rivulets and overflows functioning as dispersal pathways. Using a landscape-based approach, we modelled the effective geographical distance among a set of interconnected ponds (De Maten, Genk, Belgium) in a Geographic Information System (GIS) environment. The first model (the Landscape Model; LM) corrects for the presence of direct connections among ponds and was based on the existing landscape structure (i.e. network of connecting elements among ponds, travelling distance and direction of the current). A second model (the Flow Rate Model; FRM) also incorporated field data on flow rates in the connecting elements as the driving force for the passive dispersal of the active zooplankton population component. Finally, the third model (the Dispersal Rate Model; DRM) incorporated field data on zooplankton dispersal rates. An analysis of the pattern of genetic differentiation among Daphnia ambigua populations inhabiting 10 ponds in the pond complex reveals that the effective geographical distance as modelled by the flow rate and the dispersal rate model provide a better approximation of the true rates of genetic exchange among populations than mere Euclidean geographical distances or the landscape model that takes into account solely the presence of physical connections.     

Joint effects of species traits and environmental preferences on range edge shifts of British birds

Aim

Despite the strong evidence of species range shifts as a response to environmental change, attempts to identify species traits that modulate those shifts have been equivocal. We investigate the role of species traits and environmental preferences on birds' range shifts in Great Britain, an island where dispersal is limited by the English Channel and the North Sea.

Location

Great Britain (England, Scotland and Wales).

Taxa

Birds (Aves).

Time Period

1968–2011.

Methods

Using 404,949 occurrence records from two time periods, we investigated the potential drivers of leading and rear range edge shifts of breeding birds using phylogenetic linear mixed models. We hypothesized that shifts are influenced by species' trophic and morphological traits, dispersal abilities and environmental preferences, but also by the geographical boundaries of Great Britain.

Results

Geographical boundaries—the distance from the northern or southern boundaries of Britain—accounted for most of the variability in range edge shifts. Species traits and environmental preferences emerged as relevant drivers of range shifts only for northern and Passeriform species. Northern habitat specialist, those with more predators and those sensitive to precipitation were more likely to shift their rear edge poleward. For Passeriformes, habitat generalists, species with smaller dispersal capabilities, under higher predatory pressure or associated with forest and grassland were more likely to shift their rear edge poleward.

Main Conclusions

While geographical boundaries impose constraints on range shifts in British birds, the subtle effects of species traits and environmental preferences emerge as relevant predictors for Northern and passeriform species' rear edge shifts. This highlights the importance of accounting for geographical boundaries when predicting species responses to global change. Differential range shifts of species across different trophic levels could result in the reorganization of biotic interactions, with consequences for ecosystem structure and stability.     

Tangled webs: reciprocal flows of invertebrate prey link streams and riparian zones

1. Streams and their adjacent riparian zones are closely linked by reciprocal flows of invertebrate prey. We review characteristics of these prey subsidies and their strong direct and indirect effects on consumers and recipient food webs. 2. Fluxes of terrestrial invertebrates to streams can provide up to half the annual energy budget for drift‐feeding fishes such as salmonids, despite the fact that input occurs principally in summer. Inputs appear highest from closed‐canopy riparian zones with deciduous vegetation and vary markedly with invertebrate phenology and weather. Two field experiments that manipulated this prey subsidy showed that it affected both foraging and local abundance of stream fishes. 3. Emergence of adult insects from streams can constitute a substantial export of benthic production to riparian consumers such as birds, bats, lizards, and spiders, and contributes 25–100% of the energy or carbon to such species. Emergence typically peaks in early summer in the temperate zone, but also provides a low‐level flux from autumn to spring in ice‐free streams. This flux varies with in‐stream productivity, and declines exponentially with distance from the stream edge. Some predators aggregate near streams and forage on these prey during periods of peak emergence, whereas others rely on the lower subsidy from autumn through spring when terrestrial prey are scarce. Several field experiments that manipulated this subsidy showed that it affected the short‐term behaviour, growth, and abundance of terrestrial consumers. 4. Reciprocal prey subsidies also have important indirect effects on both stream and riparian food webs. Theory predicts that allochthonous prey should increase density of subsidised predators, thereby increasing predation on in situ prey and causing a negative indirect effect via apparent competition. However, short‐term experiments have produced either positive or negative indirect effects. These contrasting results may be due to characteristics of the subsidies and individual consumers, but could also result from differences in experimental designs. 5. New study approaches are needed to better determine the direct and indirect effects of reciprocal prey subsidies. Experiments coupled with comparative research will be required to measure their effects on individual consumer fitness and population demographics. Future work should investigate whether reciprocal prey fluxes stabilise linked stream–riparian ecosystems, explore how landscape context affects the magnitude and importance of subsidies, and determine how impacts of human disturbance can propagate between streams and riparian zones via these trophic linkages. Study of these reciprocal connections is helping to define a more holistic perspective of catchments, and has the potential to shape new directions for ecology in general.