Direct versus indirect effects of habitat fragmentation on community patterns in experimental landscapes.

Habitat area and fragmentation are confounded in many ecological studies investigating fragmentation effects. We thus devised an innovative experiment founded on fractal neutral landscape models to disentangle the relative effects of habitat area and fragmentation on arthropod community patterns in red clover (Trifolium pratense). The conventional approach in experimental fragmentation studies is to adjust patch size and isolation to create different landscape patterns. We instead use fractal distributions to adjust the overall amount and fragmentation of habitat independently at the scale of the entire landscape, producing different patch properties. Although habitat area ultimately had a greater effect on arthropod abundance and diversity in this system, we found that fragmentation had a significant effect in clover landscapes with ≤40 % habitat. Landscapes at these lower habitat levels were dominated by edge cells, which had fewer arthropods and lower richness than interior cells. Fragmentation per se did not have a direct effect on local-scale diversity, however, as demonstrated by the lack of a broader landscape effect (in terms of total habitat area and fragmentation) on arthropods within habitat cells. Fragmentation-through the creation of edge habitat-thus had a strong indirect effect on morphospecies richness and abundance at the local scale. Although it has been suggested that fragmentation should be important at low habitat levels (≤20-30 %), we show that fragmentation per se is significant only at intermediate (40 %) levels of habitat, where edge effects were neither too great (as at lower levels of habitat) nor too weak (as at higher levels of habitat).

Linking snake habitat use to nest predation risk in grassland birds: the dangers of shrub cover.

Extremes in rangeland management, varying from too-frequent fire and intensive grazing to the suppression of both, threaten rangeland ecosystems worldwide. Intensive fire and grazing denude and homogenize vegetation whereas their suppression increases woody cover. Although habitat loss is implicated in grassland bird declines, degradation through intensive management or neglect also decreases breeding habitat and may reduce nesting success through increased rates of nest predation. Snakes are important nest predators, but little is known about how habitat use in snakes relates to predation risk for grassland birds nesting within tallgrass prairie subjected to different grazing and fire frequencies. We evaluated nest survival in the context of habitat used by nesting songbirds and two bird-eating snakes, the eastern yellowbelly racer Coluber constrictor flaviventris and Great Plains ratsnake Pantherophis emoryi. Daily nest survival rates decreased with increasing shrub cover and decreasing vegetation height, which characterize grasslands that have been neglected or intensively managed, respectively. Discriminant function analysis revealed that snake habitats were characterized by higher shrub cover, whereas successful nests were more likely to occur in areas with tall grass and forbs but reduced shrub cover. Because snakes often use shrub habitat, birds nesting in areas with increased shrub cover may be at higher risk of nest predation by snakes in addition to other predators known to use shrub habitat (e.g., mid-sized carnivores and avian predators). Depredated nests also occurred outside the discriminant space of the snakes, indicating that other predators (e.g., ground squirrels Spermophilus spp. and bullsnakes Pituophis catenifer) may be important in areas with denuded cover. Targeted removal of shrubs may increase nest success by minimizing the activity of nest predators attracted to shrub cover.

Grassland bird responses to land management in the largest remaining tallgrass prairie.

Extensive habitat loss and changing agricultural practices have caused widespread declines in grassland birds throughout North America. The Flint Hills of Kansas and Oklahoma--the largest remaining tallgrass prairie--is important for grassland bird conservation despite supporting a major cattle industry. In 2004 and 2005, we assessed the community, population, and demographic responses of grassland birds to the predominant management practices (grazing, burning, and haying) of the region, including grasslands restored under the Conservation Reserve Program (CRP). We targeted 3 species at the core of this avian community: the Dickcissel (Spiza americana), Grasshopper Sparrow (Ammodramus savannarum), and Eastern Meadowlark (Sturnella magna). Bird diversity was higher in native prairie hayfields and grazed pastures than CRP fields, which were dominated by Dickcissels. Although Dickcissel density was highest in CRP, their nest success was highest and nest parasitism by Brown-headed Cowbirds (Moluthrus ater) lowest in unburned hayfields (in 2004). Conversely, Grasshopper Sparrow density was highest in grazed pastures, but their nest success was lowest in these pastures and highest in burned hayfields, where cowbird parasitism was also lowest (in 2004). Management did not influence density and nest survival of Eastern Meadowlarks, which were uniformly low across the region. Nest success was extremely low (5-12%) for all 3 species in 2005, perhaps because of a record spring drought. Although the CRP has benefited grassland birds in agricultural landscapes, these areas may have lower habitat value in the context of native prairie. Hayfields may provide beneficial habitat for some grassland birds in the Flint Hills because they are mowed later in the breeding season than elsewhere in the Midwest. Widespread grazing and annual burning have homogenized habitat-and thus grassland-bird responses-across the Flint Hills. Diversification of management practices could increase habitat heterogeneity and enhance the conservation potential of the Flint Hills for grassland birds.

Pest and disease management: why we shouldn't go against the grain.

Given the wide range of scales and mechanisms by which pest or disease agents disperse, it is unclear whether there might exist a general relationship between scale of host heterogeneity and spatial spread that could be exploited by available management options. In this model-based study, we investigate the interaction between host distributions and the spread of pests and diseases using an array of models that encompass the dispersal and spread of a diverse range of economically important species: a major insect pest of coniferous forests in western North America, the mountain pine beetle (Dendroctonus ponderosae); the bacterium Pseudomonas syringae, one of the most-widespread and best-studied bacterial plant pathogens; the mosquito Culex erraticus, an important vector for many human and animal pathogens, including West Nile Virus; and the oomycete Phytophthora infestans, the causal agent of potato late blight. Our model results reveal an interesting general phenomenon: a unimodal ('humpbacked') relationship in the magnitude of infestation (an index of dispersal or population spread) with increasing grain size (i.e., the finest scale of patchiness) in the host distribution. Pest and disease management strategies targeting different aspects of host pattern (e.g., abundance, aggregation, isolation, quality) modified the shape of this relationship, but not the general unimodal form. This is a previously unreported effect that provides insight into the spatial scale at which management interventions are most likely to be successful, which, notably, do not always match the scale corresponding to maximum infestation. Our findings could provide a new basis for explaining historical outbreak events, and have implications for biosecurity and public health preparedness.

Why dispersal should be maximized at intermediate scales of heterogeneity.

Dispersal is a fundamental biological process that results in the redistribution of organisms due to the interplay between the mode of dispersal, the range of scales over which movement occurs, and the scale of spatial heterogeneity, in which patchiness may occur across a broad range of scales. Despite the diversity of dispersal mechanisms and dispersal length scales in nature, we posit that a fundamental scaling relationship should exist between dispersal and spatial heterogeneity. We present both a conceptual model and mathematical formalization of this expected relationship between the scale of dispersal and the scale of patchiness, which predicts that the magnitude of dispersal (number of individuals) among patches should be maximized when the scale of spatial heterogeneity (defined in terms of patch size and isolation) is neither too fine nor too coarse relative to the gap-crossing abilities of a species. We call this the "dispersal scaling hypothesis" (DSH). We demonstrate congruence in the functional form of this relationship under fundamentally different dispersal assumptions, using well-documented isotropic dispersal kernels and empirically derived dispersal parameters from diverse species, in order to explore the generality of this finding. The DSH generates testable hypotheses as to when and under what landscape scenarios dispersal is most likely to be successful. This provides insights into what management scenarios might be necessary to either restore landscape connectivity, as in certain conservation applications, or disrupt connectivity, as when attempting to manage landscapes to impede the spread of an invasive species, pest, or pathogen.

Movement behavior of red flour beetle: response to habitat cues and patch boundaries.

Movement behavior determines the success or failure of insects in finding important resources such as food, mates, reproductive sites, and shelter. We examined the response of female red flour beetles (Tribolium castaneum Herbst: Coleoptera: Tenebrionidae) to habitat cues by quantifying the number of individuals that located a patch (either with or without flour) in response to the distance released from the patch, air movement over the arena, and food-deprivation status. We also investigated how patch characteristics, such as resource amount and presence of cover, influenced time taken to find a flour patch, the frequency of entering or leaving, and residence time within the patch. Although the proportion of beetles successfully locating the patch decreased as a function of release distance, the probability that beetles reached the patch was ultimately unaffected by whether flour was present or not, suggesting that search behavior in red flour beetles may exhibit a simple distance-decay function. Significantly more beetles reached the patch when they had not been food deprived and air was flowing over the arena, which indicates that walking beetles may orient to airflow, exhibiting anemotaxis. Results of the second experiment showed that, on first encounter, fewer beetles entered patches with a greater amount of flour; but once they had entered, they left them less frequently than patches with less resource. Beetles entered covered patches more quickly than uncovered patches irrespective of resource amount, which indicates that shelter is perhaps more important to red flour beetles than resource levels in determining whether to enter patches.

Assessing the risk of invasive spread in fragmented landscapes.

Little theoretical work has investigated how landscape structure affects invasive spread, even though broad-scale disturbances caused by habitat loss and fragmentation are believed to facilitate the spread of exotic species. Neutral landscape models (NLMs), derived from percolation theory in the field of landscape ecology, provide a tool for assessing the risk of invasive spread in fragmented landscapes. A percolation-based analysis of the potential for invasive spread in fragmented landscapes predicts that invasive spread may be enormously enhanced beyond some threshold level of habitat loss, which depends upon the species' dispersal abilities and the degree of habitat fragmentation. Assuming that invasive species spread primarily through disturbed areas of the landscape, poor dispersers may spread better in landscapes in which disturbances are concentrated in space, whereas good dispersers are predicted to spread better in landscapes where disturbances are small and dispersed (i.e., fragmented landscape). Assessing the risk of invasive spread in fragmented landscapes ultimately requires understanding the relative effects of landscape structure on processes that contribute to invasive spread--dispersal (successful colonization) and demography (successful establishment). Colonization success is predicted to be highest when >20% of the landscape has been disturbed, particularly if disturbances are large or aggregated in space, because propagules are more likely to encounter sites suitable for colonization and establishment. However, landscape pattern becomes less important for predicting colonization success if species are capable of occasional long-distance dispersal events. Invasive species are also more likely to persist and achieve positive population growth rates (successful establishment) in landscapes with clumped disturbance patterns, which can then function as population sources that produce immigrants that invade other landscapes. Finally, the invasibility of communities may be greatest in landscapes with a concentrated pattern of disturbance, especially below some critical threshold of biodiversity. Below the critical biodiversity threshold, the introduction of a single species can trigger a cascade of extinctions among indigenous species. The application of NLMs may thus offer new insights and opportunities for the management and restoration of landscapes so as to slow the spread of invasive species.