Agricultural landscapes with organic crops support higher pollinator diversity

Pollinators are traditionally thought to perceive non-flowering crop fields as hostile landscape matrix. In this study, we show that landscapes composed of higher proportions of organic crop fields support more bee species at greater abundances in fallow strips. An increase in organic cropping in the surrounding landscape from 5% to 20% enhanced bee species richness in fallow strips by 50%, density of solitary bees by 60% and bumble bee density by 150%. Bee species richness and bumble bee density responded strongest to organic cropping in landscape sectors with 500 m radius, solitary bee density in landscape sectors with 250 m radius. The most likely source of these results is that crop and noncrop habitats are strongly connected via bee foraging at the landscape scale. It seems likely that bees depending on nesting sites in fallow strips benefited from the more abundant flower resources provided by broadleaved weeds in organic crop fields. We conclude that the incorporation of organic crop fields into conventionally managed agricultural landscapes can provide food resources needed to sustain greater pollinator species richness in noncrop habitats.     

Quantification of bacterial chemotaxis by measurement of model parameters using the capillary assay

The capillary assay for quantitative characterization of bacterial motility and chemotaxis is analyzed in terms of a mathematical model for cell population migration, in order to determine values for the cell random motility coefficient, mu and the cell chemotaxis coefficient, chi. The analysis involves both analytical perturbation methods and numerical finite-difference techniques. Transient cell density profiles within the capillary tube are determined as they depend upon mu and chi, providing a means for estimating mu and chi from the common protocol measurements of cell accumulation in the tube at specified observation times. The effects of extraneous factors such as assay geometry, stimulus diffusivity, bacterial density, and observation time are thus separated from the intrinsic cell-stimulus interaction and response. This allows independent population measurements of cell chemosensory movement properties to be extrapolated to situations involving growth and competition of populations, for purposes of better understanding microbial population dynamics in systems of biotechnological and microbial ecological importance.     

Stochastic Optimal Foraging: Tuning Intensive and Extensive Dynamics in Random Searches

Recent theoretical developments had laid down the proper mathematical means to understand how the structural complexity of search patterns may improve foraging efficiency. Under information-deprived scenarios and specific landscape configurations, Lévy walks and flights are known to lead to high search efficiencies. Based on a one-dimensional comparative analysis we show a mechanism by which, at random, a searcher can optimize the encounter with close and distant targets. The mechanism consists of combining an optimal diffusivity (optimally enhanced diffusion) with a minimal diffusion constant. In such a way the search dynamics adequately balances the tension between finding close and distant targets, while, at the same time, shifts the optimal balance towards relatively larger close-to-distant target encounter ratios. We find that introducing a multiscale set of reorientations ensures both a thorough local space exploration without oversampling and a fast spreading dynamics at the large scale. Lévy reorientation patterns account for these properties but other reorientation strategies providing similar statistical signatures can mimic or achieve comparable efficiencies. Hence, the present work unveils general mechanisms underlying efficient random search, beyond the Lévy model. Our results suggest that animals could tune key statistical movement properties (e.g. enhanced diffusivity, minimal diffusion constant) to cope with the very general problem of balancing out intensive and extensive random searching. We believe that theoretical developments to mechanistically understand stochastic search strategies, such as the one here proposed, are crucial to develop an empirically verifiable and comprehensive animal foraging theory.     

Time‐to‐kill: measuring attack rates in a heterogenous landscape with multiple prey types

Although spatial heterogeneity of prey and landscapes are known to contribute to variation around predator‐prey functional response models, few studies have quantified these effects. We illustrate a new approach using data from winter movement paths of GPS‐collared wolves in the Rocky Mountains of Canada and time‐to‐event models with competing risks for measuring the effect of prey and landscape characteristics on the time‐to‐kill, which is the reciprocal of attack rate (aN) in a Holling's functional response. We evaluated 13 a priori models representing hypothesized mechanisms influencing attack rates in a heterogeneous landscape with two prey types. Models ranged from variants on Holling's disc equation, including search rate and prey density, to a full model including prey density and patchiness, search rates, satiation, and landscape features, which were measured along the wolf's movement path. Movement rates of wolves while searching explained more of the variation in time‐to‐kill than prey densities. Wolves did not compensate for low prey density by increasing movement rates and there was little evidence that spatial aggregation of prey influenced attack rates in this multi‐prey system. The top model for predicting time‐to‐kill included only search rate and landscape features. Wolves killed prey more quickly in flat terrain, likely due to increased vulnerability from accumulated snow, whereas attack rates were lower when wolves hunted near human‐made features presumably due to human disturbance. Understanding the sources of variation in attack rates provides refinements to functional response models that can lead to more effective predator–prey management in human‐dominated landscapes.     

Advection–diffusion equations for generalized tactic searching behaviors

 Many organisms search for limiting resources by using repeated responses to local cues, which cumulatively cause movement towards more favorable parts of their environment. This paper presents a general asymptotic expression, derived under the assumption of shallow environmental gradients, for the population-level flux of organisms moving at a constant speed and reorienting at rates determined by the environmental conditions experienced since the last reorientation. The expression takes the form of an advection-diffusion equation, in which the diffusivity and advection velocity are determined by statistics of the turning algorithm that are directly comparable to empirical observations. This work provides a mechanism with which to systematically evaluate a wide variety of tactic and kinetic strategies for determining turning behaviors. The model is applied to searchers on spatially-variable, random distributions of discrete resource patches. Such algorithms are functions of the integrated resource density encountered between turns. It is shown that behaviors in which the turning time distribution is a function of integrated density cannot result in taxis. In contrast, behaviors in which the turning rate is a function of integrated density can result in taxis. These two classes of search algorithm differ in that the latter requires the searcher to “learn” about its local environment, whereas the former requires no such assessment. This suggests neural or physiological mechanisms for remembering previous encounters may be a biological requirement for searchers on discrete resource distributions. Received: 21 September 1995/Revised version: 18 July 1996     

Local and landscape effects on butterfly density in northern Idaho grasslands and forests

Understanding butterfly response to landscape context can inform conservation management and planning. We tested whether local-scale resources (host and nectar plants, canopy cover) or landscape context, measured at two scales, better explained the densities of four butterfly species. The density of Coenonympha tullia, which has host plants strongly associated with grassland habitats, was positively correlated with the amount of grassland in 0.5- and 1-km radius landscapes and only occurred in forests when they bordered grasslands. For the other species, Celastrina ladon, Cupido amyntula, and Vanessa cardui, local-scale resources better explained butterfly densities, emphasizing the importance of local habitat quality for butterflies. These three species also used host plants that were distributed more heterogeneously within and among habitat types. Our findings demonstrate the importance of host plant spatial distributions when determining the scale at which butterfly density relates to resources, and we recommend that both these distributions and landscape context be evaluated when developing butterfly monitoring programs, managing for species of concern, or modeling potential habitat.     

Measurement of leukocyte motility and chemotaxis parameters with a linear under-agarose migration assay

The interpretation of quantitative assays for leukocyte chemotactic migration is usually made in terms of measurements such as leading front distance, total migrating cells, and leukotactic index. These quantities allow comparison of cellular migration behavior under specified conditions. They are not useful; however, for comparisons between systems or for correlation with in vivo performance, because they depend upon specific physical aspects of the assay system, such as the geometry, chemoattractant concentration and diffusivity, and observation time. It would be more helpful to measure intrinsic properties of cell movement that could be used for comparison between systems, for correlation with in vivo studies, and to increase our understanding of the cell physiology. In this paper we demonstrate a means of quantitating leukocyte random motility, chemokinesis, and chemotaxis in terms of parameters that do characterize intrinsic cell properties. These parameters are the random motility coefficient and the chemotaxis coefficient, which appear in theoretical models of cell migration. We examine how well such a model describes the leukocyte density profile data observed in a modified under-agarose assay having a linear geometry. Furthermore, we obtain values for the random motility coefficient (and its dependence upon the concentration of the attractant peptide FNLLP) and for the chemotaxis coefficient for leukocytes responding to FNLLP.     

Density-dependent adaptive resistance allows swimming bacteria to colonize an antibiotic gradient

During antibiotic treatment, antibiotic concentration gradients develop. Little is know regarding the effects of antibiotic gradients on populations of nonresistant bacteria. Using a microfluidic device, we show that high-density motile Escherichia coli populations composed of nonresistant bacteria can, unexpectedly, colonize environments where a lethal concentration of the antibiotic kanamycin is present. Colonizing bacteria establish an adaptively resistant population, which remains viable for over 24 h while exposed to the antibiotic. Quantitative analysis of multiple colonization events shows that collectively swimming bacteria need to exceed a critical population density in order to successfully colonize the antibiotic landscape. After colonization, bacteria are not dormant but show both growth and swimming motility under antibiotic stress. Our results highlight the importance of motility and population density in facilitating adaptive resistance, and indicate that adaptive resistance may be a first step to the emergence of genetically encoded resistance in landscapes of antibiotic gradients.     

Demographic stochasticity and resource autocorrelation control biological invasions in heterogeneous landscapes

Mounting theoretical evidence suggests that demographic stochasticity, environmental heterogeneity and biased movement of organisms individually affect the dynamics of biological invasions and range expansions. Studies of species spread in heterogeneous landscapes have traditionally characterized invasion velocities as functions of the mean resource density throughout the landscape, thus neglecting higher‐order moments of the spatial resource distribution. Here, we show theoretically that different spatial arrangements of resources lead to different spread velocities even if the mean resource density throughout the landscape is kept constant. Specifically, we find that increasing the resource autocorrelation length causes a reduction in the speed of species spread. The model shows that demographic stochasticity plays a key role in the slowdown, which is strengthened when individuals can actively move towards resources. We then experimentally corroborated the theoretically predicted reduction in propagation speed in microcosm experiments with the protist Euglena gracilis by comparing spread in landscapes with different resource autocorrelation lengths. Our work identifies the resource autocorrelation length as a key modulator and a simple measure of landscape susceptibility to biological invasions, which needs to be considered for predicting invasion dynamics within naturally heterogeneous environmental corridors.     

Animal response to nested self-similar patches: a test with woolly bears

To gain insight into how animals respond to resource patchiness at different spatial scales, we envision their responses in environments comprised of nested, self-similar patches. In these environments, all resources reside within the smallest patches, and resource density declines as a constant exponent of patch size. Accordingly, we use simple mathematical formulations to describe a self-similar environment and a null model of how animals should respond to this environment if they do not perceive resource distribution. We then argue that animals that can perceive resource distribution should partition space by reducing the relative time searching between patches as patch size increases. On an experimental landscape, we found that woolly bear caterpillars Grammia geneura could partition space in this manner, but the range of patch sizes over which they did so tended to increase with resource aggregation. Nevertheless, scaling efficiency (i.e. the scaling of search time versus the scaling or resource density) was similar in all distributions when averaged over all patch sizes. These disparate patterns with similar outcomes resulted from differences in caterpillars' abilities to discriminate spatially among patches of different sizes via their movement pathways, and differences in their use of speed to detect resource items. Our work is relevant to the characterization of resource availability from an animal's perspective, and to the linking of optimal foraging theory to the modeling of search behavior.     

Beyond diffusion: Modelling local and long-distance dispersal for organisms exhibiting intensive and extensive search modes

The distribution of foragers on the landscape has important consequences to, for example, the spread rate of an invasive species or the outcrossing levels between neighbouring crops. Since forager distribution can be difficult to measure directly, mathematical models are often used to predict the population density of dispersing foragers on the landscape. We model organism movement using a diffusion framework in which the foraging population is divided into two subpopulations engaged in intensive and extensive search modes respectively. Movement in the intensive search mode (ISM) is modeled by diffusion, and movement in the extensive search mode (ESM) is modeled by advection. We show that our model provides a superior fit to organism movement data than more traditional diffusion or diffusion-advection models in which the forager population is considered homogeneous. Our results have implications for the understanding of dispersal in a wide variety of applications.     

Biomedicine on the Spatial Periphery: The (Re)Production of a Metaphorical Landscape by Rural Health Care Practitioners in Northern California

I examine the use of spatial concepts by rural health care practitioners in Northern California and suggest that rural and urban spatial metaphors are important means of expressing and (re)producing problems associated with their search for legitimacy and moral authority within a field of relations defined by biomedicine. I present three broad ways in which spatial metaphors are used by rural health care practitioners to continuously enact a "metaphorical landscape." I situate this landscape in the context of a hierarchical field of relations within biomedicine, which is itself underpinned by a distinct urban bias and the uneven distribution of material and technological resources. I suggest that this landscape is partly the result of the rural health care practitioners' position within this field of relations and partly the result of implicit and historically situated frameworks of spatial meanings derived from capitalism.     

Spatial and temporal variations in floral resource availability affect bumblebee communities in heathlands

Modifications of landscape structure and composition can decrease the availability of floral resources, resulting in the decline of many pollinator species, including bumblebees. These declines may have significant ecological consequences, because bumblebees pollinate a large range of plant species. Our study was carried out in heathlands, open semi-natural habitats that have decreased considerably due to human activities. We analysed how floral resources affect bumblebee communities throughout the colony lifetime at three scales: plot scale, heathland patch scale, and landscape scale. Floral density at the plot scale and spruce plantations at the landscape scale influenced bumblebee communities. The abundance of bumblebees on ericaceous species was higher when the landscape included a substantial proportion of unsuitable foraging habitat (i.e., spruce plantations). Both life history traits and colony life cycle stage influenced bumblebee responses to the availability of floral resources. Bumblebees were more affected by floral resources during the colony development phase than during the nest-foundation or mating phases. Moreover, bumblebees of species that form large colonies needed larger quantities of favourable foraging habitat, compared with small-colony bees, and their proportion decreased in habitats dominated by spruce plantations. In conclusion, the conservation of plant–bumblebee interactions will require management at a larger spatial scale than the restricted protected habitats. Moreover, at the landscape scale, both quantity of favourable foraging patches and their ecological continuity are important to conserve both small- and large- colony species.     

Flight morphology corresponds to both surrounding landscape structure and local patch conditions in a highly specialized peatland butterfly (Lycaena epixanthe)

1. Movement mediates the response of populations and communities to landscape and habitat spatial structure, yet movement capability may itself be modified by selection pressures accompanying landscape change. Insect flight morphology can be affected by both the landscape surrounding habitat patches and the distribution of resources within habitat patches. 2. This study investigated the relative influence of local habitat patch conditions and surrounding landscape structure on variation in morphological traits associated with flight in the bog copper (Lycaena epixanthe), a butterfly endemic to temperate Nearctic peatlands. 3. Eight habitat patches were sampled to assess the influence of the surrounding landscape (connectivity of potential habitat and matrix composition) and patch size (an integrated proxy of resource density and spatial distribution) on investment into flight, measured by thorax and abdomen mass, and wing area. 4. The results revealed an effect of both local habitat conditions and landscape structure on flight‐related morphological traits. Increasing forest cover in the surrounding landscape, indicative of increased habitat patch isolation, corresponded with less mobile phenotypes in both sexes. Surrounding landscapes with more water were also generally associated with less mobile phenotypes. Investment into flight was greater in smaller peatlands in which host plant density is higher and more homogeneously distributed. 5. The present study highlights that morphological traits associated with mobility may be responding to both local habitat patch characteristics and surrounding landscape structure. It also supports the hypothesis that local habitat conditions contribute to morphological variation in butterflies.     

Landscape features and spatial distribution of adult northern corn rootworm (Coleoptera: Chrysomelidae) in the South Dakota areawide management site

The northern corn rootworm, Diabrotica barberi Smith & Lawrence (Coleoptera: Chrysomelidae), creates economic and environmental concerns in the Corn Belt. To supplement the population control tactics of the Areawide Pest Management Program in Brookings, SD, geographic information systems were used from 1997 to 2001 to examine the spatial relationships between D. barberi population dynamics and habitat structure, soil texture, and elevation. Using the inverse distance weighted interpolation technique, D. barberi population density maps were created from georeferenced emergence and postemergence traps placed in maize, Zea mays L., fields. For each year, these maps were overlaid with vegetation, soil, and elevation maps to search for quantitative relationships between pest numbers and landscape structure. Through visual interpretation and correlation analysis, shifts in landscape structure, such as size, number, and arrangement of patches were shown to associate with D. barberi population abundance and distribution in varying degrees. D. barberi were found in greater proportions than expected on loam and silty clay loam soils and on elevations between 500 and 509 m. An understanding of the interactions between D. barberi population dynamics and landscape variables provides information to pest managers, which can be used to identify patterns in the landscape that promote high insect population density patches to improve pest management strategies.     

More pests but less pesticide applications: Ambivalent effect of landscape complexity on conservation biological control

In agricultural landscapes, the amount and organization of crops and semi-natural habitats (SNH) have the potential to promote a bundle of ecosystem services due to their influence on ecological community at multiple spatio-temporal scales. SNH are relatively undisturbed and are often source of complementary resources and refuges, therefore supporting more diverse and abundant natural pest enemies. However, the nexus of SNH proportion and organization with pest suppression is not trivial. It is thus crucial to understand how the behavior of pest and natural enemy species, the underlying landscape structure, and their interaction, may influence conservation biological control (CBC). Here, we develop a generative stochastic landscape model to simulate realistic agricultural landscape compositions and configurations of fields and linear elements. Generated landscapes are used as spatial support over which we simulate a spatially explicit predator-prey dynamic model. We find that increased SNH presence boosts predator populations by sustaining high predator density that regulates and keeps pest density below the pesticide application threshold. However, predator presence over all the landscape helps to stabilize the pest population by keeping it under this threshold, which tends to increase pest density at the landscape scale. In addition, the joint effect of SNH presence and predator dispersal ability among hedge and field interface results in a stronger pest regulation, which also limits pest growth. Considering properties of both fields and linear elements, such as local structure and geometric features, provides deeper insights for pest regulation; for example, hedge presence at crop field boundaries clearly strengthens CBC. Our results highlight that the integration of species behaviors and traits with landscape structure at multiple scales is necessary to provide useful insights for CBC.     

Spatio‐temporal dynamics of density‐dependent dispersal during a population colonisation

Predicting population colonisations requires understanding how spatio‐temporal changes in density affect dispersal. Density can inform on fitness prospects, acting as a cue for either habitat quality, or competition over resources. However, when escaping competition, high local density should only increase emigration if lower‐density patches are available elsewhere. Few empirical studies on dispersal have considered the effects of density at the local and landscape scale simultaneously. To explore this, we analyze 5 years of individual‐based data from an experimental introduction of wild guppies Poecilia reticulata. Natal dispersal showed a decrease in local density dependence as density at the landscape level increased. Landscape density did not affect dispersal among adults, but local density‐dependent dispersal switched from negative (conspecific attraction) to positive (conspecific avoidance), as the colonisation progressed. This study demonstrates that densities at various scales interact to determine dispersal, and suggests that dispersal trade‐offs differ across life stages.     

艾比湖区域景观格局空间特征与地表水质的关联分析

为进一步明确区域土地利用/覆被—景观格局对河流水质影响的空间尺度。选择新疆艾比湖区域为研究对象,以25个水质采样点为中心,建立5种尺度的河流缓冲区并提取不同尺度下的土地利用/覆被-景观格局数据。首先,通过主成分分析获得水环境的主要水质变量。其次,利用冗余分析(RDA)方法探讨研究区不同宽度缓冲区土地利用/覆被—景观格局对河流水质的影响,获得水质管理的有效缓冲区。最后,引入突变点分析方法进一步寻找导致水质变量沿景观梯度突变的特定位置。结果表明:(1)土地利用/覆被—景观格局在不同宽度缓冲区内对河流水质的影响不同。4 km缓冲区土地利用/覆被—景观格局对区域水质有较强的分异解释能力,因此4 km缓冲区的景观格局合理配置对河流水质管理尤为重要。(2)通过偏RDA分析发现4 km缓冲区中,影响区域水质的主要环境变量为景观水平斑块密度、类型水平耕地斑块密度和森林聚集度。(3)在偏RDA分析的基础上,对4 km缓冲区内的景观指数进行突变点分析研究,发现区域景观水平斑块密度为90—105 m/hm~2,类型水平耕地的斑块密度ED值在90—110 m/hm~2、林草地的AI值在70%—90%,是艾比湖区域水质保护的最佳突变值,该值为艾比湖区域水质保护的阈值。本研究从多个角度对新疆艾比区域的土地利用/覆被—景观格局对区域水质的影响进行定量分析,揭示该地区景观生态变化的规律,为区域景观格局优化和土地合理规划提供理论依据,并对改善"丝绸之路经济带"生态环境、实现资源可持续利用具有重要的现实意义和理论价值。     

Distribution of recreational boating across lakes: do landscape variables affect recreational use?

1. Because people impact lake ecosystems, it is important to consider factors influencing the human use of freshwater resources. We investigated the influence of the landscape position, as well as lake area, recreational facilities, and distance to highways and urban centres, on lake use by boaters in the Northern Highland Lake District of Wisconsin, U.S.A.
2. In aerial surveys of ninety-nine randomly selected lakes, we did not see boats on over half of the lakes. Of the lakes with boats, we found a strong correlation between the number of boats and lake area. Recreational boats tended to be found on large, accessible lakes with good boating facilities. Boats were not seen on small, stained lakes with few recreational facilities.
3. Regression models showed that lake size and landscape position explained 63% of the variability in the average number of boats per lake and landscape position explained 24% of the variability in boat density on all ninety-nine lakes. Social variables representing the quality of boating facilities and the perception of good fishing explained 70% of the variability in number of boats per lake and 54% of the variation in boat density on all lakes. A combined model using both physical and social variables increased the explanatory power for both number and density. Lake use by boaters was correlated with landscape position, the quality of fishing and the availability of recreational facilities. When the analysis was restricted to the forty-six lakes where boats were found, only the availability of recreational facilities proved a significant predictor of boat density.
4. Our results suggest that lake choice by recreational boaters may be best predicted by a combination of the location of a lake in a regional hydrologic landscape, and considerations of available facilities and perceptions regarding fishing.     

Adaptive Patch Searching Strategies in Fragmented Landscapes

The search strategies dispersers employ to search for new habitat patches affect individuals’ search success and subsequently landscape connectivity and metapopulation viability. Some evidence indicates that individuals within the same species may display a variety of behavioural patch searching strategies rather than one species-specific strategy. This may result from landscape heterogeneity. We modelled the evolution of individual patch searching strategies in different landscapes. Specifically, we analysed whether evolution can favour different, co-existing, behavioural search strategies within one population and to what extent this coexistence of multiple strategies was dependent on landscape configuration. Using an individual-based simulation model, we studied the evolution of patch searching strategies in three different landscape configurations: uniform, random and clumped. We found that landscape configuration strongly influenced the evolved search strategy. In uniform landscapes, one fixed search strategy evolved for the entire spatially structured population, while in random and clumped landscapes, a set of different search strategies emerged. The coexistence of several search strategies also strongly depended on the dispersal mortality. We show that our result can affect landscape connectivity and metapopulation dynamics. Co-ordinating editor: N. Yamamura