Landscape context and selection for forest edge by breeding Brown-headed Cowbirds

We evaluated support for four alternate hypotheses explaining the distribution of breeding Brown-headed Cowbirds (Molothrus ater) in forests at varying distances from the forest edge in three Midwestern USA landscapes with varying amounts of forest fragmentation (core forest area ranged from 5 to 70%). We focused on breeding cowbirds’ use of forest because of the risk of nest parasitism to forest-dwelling hosts and to identify factors affecting breeding cowbird habitat selection. We compared distances of cowbird locations in the forest from the forest edge (“edge distances”) to distances of random forest locations in the entire landscape or within individual cowbird home ranges. We analyzed 1322 locations of 84 cowbirds across three landscapes. We found support for the landscape context hypothesis that breeding cowbird preference for forest edge varied with landscape context. Ninety percent of cowbird locations were within 150–350 m of forest edge, despite the overall availability of forest at greater distances from edge (as far as 500–1450 m) both within cowbird home ranges and the entire forested landscape. Cowbird preference for edge varied by landscape context largely due to differences in the availability of forest edge. In a highly fragmented forest cowbirds utilized the entire forest and likely viewed it as “all edge.” In less fragmented forests, cowbirds preferred edge. We consider how variation in cowbird edge preference might relate to patterns in host abundance, host diversity, and host quality because cowbird movements indicate they are capable of using forest farther from edges.     

Cumulative effects of roads and logging on landscape structure in the San Juan Mountains,Colorado (USA)

In the southern Rocky Mountains of temperate North America, the effects of Euro-American activities on disturbance regimes and landscape patterns have been less ubiquitous and less straightforward in high-elevation landscapes than in low-elevation landscapes. Despite apparently little change in the natural disturbance regime, there is increasing concern that forest management activities related mainly to timber harvest and to the extensive network of roads constructed to support timber harvest, fire control, and recreation since the late 1800s have altered disturbance regimes and landscape structure. We investigated the magnitude of change in landscape structure resulting from roads and logging since the onset of timber harvest activities in 1950. We found limited evidence for significant impacts in our study area when all lands within the landscape were considered. The relatively minor changes we observed reflected the vast buffering capacity of the large proportion of lands managed for purposes other than timber (e.g., wilderness). Significant changes in landscape structure and fragmentation of mature forest were, however, evident on lands designated as suitable timberlands. Roughly half of the mature coniferous forest was converted to young stands; mean patch size and core area declined by 40% and 25%, respectively, and contrast-weighted edge density increased 2- to 3-fold. Overall, roads had a greater impact on landscape structure than logging in our study area. Indeed, the 3-fold increase in road density between 1950–1993 accounted for most of the changes in landscape configuration associated with mean patch size, edge density, and core area. The extent of area evaluated and the period over which change was evaluated had a large impact on the magnitude of change detected and our conclusions regarding the ecological significance of those changes. Specifically, the cumulative impact on landscape structure was negligible over a 10-year period, but was notable over a 40-year period. In addition, the magnitude of change in landscape structure between 1950–1993 varied as a function of landscape extent. At the scale of the 228000 ha landscape, change in landscape structure was trivial, suggesting that the landscape was capable of fully incorporating the disturbances with minimal impact. However, at intermediate scales of 1000–10000 ha landscapes, change in landscape structure was quite evident, suggesting that there may be an optimal range of scales for detecting changes in landscape structure within the study area.     

The protean relationship between boreal forest landscape structure and red squirrel distribution at multiple spatial scales

This paper investigates two fundamental questions in landscape ecology: what influence does landscape context, or the composition of the matrix, have on an animals’ response to landscape structure, and how does this relationship extrapolate between landscapes? We investigate how the distribution of North American red squirrels (Tamiasciurus hudsonicus) in the boreal mixedwood forest is influenced by anthropogenically (forest harvest) and naturally (forest fire) derived landscape structure. We studied the presence and absence of red squirrels over two years in three landscape types: one managed for timber harvest, one recently burned by wildfire, and a third unburned unmanaged landscape. Landscape composition and configuration, measured at several spatial scales, predicted red squirrel’s distribution in all three landscapes, but the significant landscape variables changed across spatial scales, across time, and across landscapes. These findings emphasize the variability in landscape structure/animal distribution relationships, and enforce the need to link pattern-finding studies, such as this one, with searches for the mechanisms behind the observed pattern.     

Broad-scale heterogeneity influences nest selection by Brown-headed Cowbirds

Increasing habitat heterogeneity is widely considered to improve conditions for biodiversity. Yet benefits for native species depend on scale and the effect of heterogeneity on key processes influencing survival and reproduction. We examined the relationship between habitat heterogeneity and brood parasitism at multiple scales in a region characterized by (1) relatively high cowbird abundance, (2) high abundance of our focal species, the grassland obligate Grasshopper Sparrow (Ammodramus savannarum), (3) variation in the structure and composition of grassland habitats, and (4) a gradient of woodland cover in the landscape matrix. Tree cover at broad scales was found to have the greatest impact on parasitism while factors at finer scales were relatively unimportant. We found that for every 1 % increase in tree cover within 1 km of Grasshopper Sparrow nests, the probability of parasitism decreases by 3 %. Parasitism reduced clutch sizes and the number of Grasshopper Sparrows fledged, but survival rates were similar between non-parasitized and parasitized nests. Furthermore, simple population projection models indicated that parasitism has the greatest impact at moderate survival levels and can inhibit the resiliency of this population. Our results support the hypothesis that cowbirds prefer forest hosts, which may reduce parasitism rates on grassland birds in heterogeneous landscapes. Collectively, our findings suggest that the effect of cowbird parasitism may be greater for Grasshopper Sparrows than was previously thought.     

Simulating the cumulative effects of multiple forest management strategies on landscape measures of forest sustainability

While the cumulative effects of the actions of multiple owners have long been recognized as critically relevant to efforts to maintain sustainable forests at the landscape scale, few studies have addressed these effects. We used the HARVEST timber harvest simulator to predict the cumulative effects of four owner groups (two paper companies, a state forest and non-industrial private owners) with different management objectives on landscape pattern in an upper Michigan landscape managed primarily for timber production. We quantified trends in landscape pattern metrics that were linked to Montreal Process indicators of forest sustainability, and used a simple wildlife habitat model to project habitat trends. Our results showed that most trends were considered favorable for forest sustainability, but that some were not. The proportion of all age classes and some forest types moved closer to presettlement conditions. The trend for the size of uneven-aged patches was essentially flat while the average size of patches of the oldest and youngest age classes increased and the size of patches of the remaining age classes decreased. Forest fragmentation generally declined, but edge density of age classes increased. Late seral forest habitat increased while early successional habitat declined. The owners use different management systems that cumulatively produce a diversity of habitats. Our approach provides a tool to evaluate such cumulative effects on other landscapes owned by multiple owners. The approach holds promise for helping landowner groups develop and evaluate cooperative strategies to improve landscape patterns for forest sustainability.     

Historic range of variability in landscape structure in subalpine forests of the Greater Yellowstone Area,USA

A measure of the historic range of variability (HRV) in landscape structure is essential for evaluating current landscape patterns of Rocky Mountain coniferous forests that have been subjected to intensive timber harvest. We used a geographic information system (GIS) and FRAGSTATS to calculate key landscape metrics on two ∼130,000-ha landscapes in the Greater Yellowstone Area, USA: one in Yellowstone National Park (YNP), which has been primarily shaped by natural fires, and a second in the adjacent Targhee National Forest (TNF), which has undergone intensive clearcutting for nearly 30 years. Digital maps of the current and historical landscape in YNP were developed from earlier stand age maps developed by Romme and Despain. Maps of the TNF landscape were adapted from United States Forest Service Resource Information System (RIS) data. Key landscape metrics were calculated at 20-yr intervals for YNP for the period from 1705-1995. These metrics were used to first evaluate the relative effects of small vs. large fire events on landscape structure and were then compared to similar metrics calculated for both pre- and post-harvest landscapes of the TNF. Large fires, such as those that burned in 1988, produced a structurally different landscape than did previous, smaller fires (1705-1985). The total number of patches of all types was higher after 1988 (694 vs. 340-404 before 1988), and mean patch size was reduced by almost half (186 ha vs. 319-379 ha). The amount of unburned forest was less following the 1988 fires (63% vs. 72-90% prior to 1988), yet the number of unburned patches increased by nearly an order of magnitude (230 vs. a maximum of 41 prior to 1988). Total core area and mean core area per patch decreased after 1988 relative to smaller fires (∼73,700 ha vs. 87,000-110,000 ha, and 320 ha vs. 2,123 ha, respectively). Notably, only edge density was similar (17 m ha⁻¹ after 1988) to earlier landscapes (9.8-14.2 m ha⁻¹).Three decades of timber harvesting dramatically altered landscape structure in the TNF. Total number of patches increased threefold (1,481 after harvest vs. 437 before harvest), and mean patch size decreased by ∼70% (91.3 ha vs. 309 ha). None of the post-harvest landscape metrics calculated for the TNF fell within the HRV as defined in YNP, even when the post-1988 landscape was considered. In contrast, pre-harvest TNF landscape metrics were all within, or very nearly within, the HRV for YNP. While reference conditions such as those identified by this study are useful for local and regional landscape evaluation and planning, additional research is necessary to understand the consequences of changes in landscape structure for population, community, ecosystem, and landscape function. This revised version was published online in July 2006 with corrections to the Cover Date.     

The influence of forest harvesting on landscape spatial patterns and old-growth-forest fragmentation in southeast British Columbia

Habitat fragmentation is considered one of the major conservation issues of recent decades. We tested predictions of landscape patterns in a 352,253-ha managed forest area in southeast British Columbia. We did this by focussing on forest fragmentation concerns among old-growth, harvest, and wildfire patches in 44 delineated landscapes using patch indices as measures of landscape pattern. We found no significant association between amount of harvesting and 15 old-growth patch indices. Comparisons among patch types revealed that amounts and spatial patterns of harvest patches differed little from amounts and spatial patterns of old-growth patches in control landscapes. Variability indices revealed similar variability between harvest patches and old-growth patches, and more variability between harvest patches and wildfire patches. Little of the evidence gathered in this study supported predictions of fragmentation of old-growth spatial patterns, or predicted differences between harvest spatial patterns and more naturally occurring spatial patterns. We suggest these results could be due to the relatively small amounts of harvesting and old-growth forest in these landscapes, and therefore habitat amount may be a more important factor than spatial configuration of patches in these landscapes.     

Using landscape metrics to model source habitat for Neotropical migrants in the midwestern U.S.

Size of a forest patch is a useful predictor of density and reproductive success of Neotropical migratory birds in much of eastern North America. Within these forested landscapes, large forest tracts appear to be sources – fragments in which surpluses of offspring are produced and can potentially colonize new fragments including woodlot sinks where reproduction fails to balance adult mortality. Within agricultural landscapes of the midwestern U.S., where forests are severely fragmented, high levels of brood parasitism by brown-headed cowbirds (Molothrus ater) and intense predation on nests generally result in low reproductive success for Neotropical migrants regardless of forest size. In some midwestern U.S. landscapes, however, the variation in reproductive success among forest fragments suggests that `source' habitat could still exist for Neotropical migrants. We used vegetation, fragment and landscape metrics to develop multivariate models that attempt to explain the variation in abundance and reproductive success of Neotropical migrants nesting in an agricultural landscape in northern Indiana, USA. We produced models that reasonably described the pattern of species richness of Neotropical migrants and the abundance of wood thrushes (Hylocichla mustelina) and several other Neotropical migrant species within 14 forest fragments. In contrast, we were unable to produce useful models of the reproductive success of wood thrushes breeding in the same forest fragments. Our results suggest that (1) abundance patterns of Neotropical migrants are probably influenced by both landscape- and fragment-scale factors; (2) multivariate analyses of Neotropical migrant abundance are not useful in modeling the corresponding patterns of reproductive success; and (3) the location of any remaining `source' habitat for Neotropical migrants breeding within agricultural landscapes in North America will be difficult to predict with indirect measures such as vegetation composition or landscape context. As a result, the potential for developing conservation strategies for Neotropical migrants will be limited without labor-intensive, direct measurements of demographic parameters.     

Forest harvest patterns and landscape disturbance processes

A physically-based model of the topographic influence on debris flow initiation and a rule-based model for wind damage were used to assess the influence of forest clearcutting patterns (i.e., location, size, shape and distribution of cut units) on the potential for landscape disturbance by these processes in Charley Creek watershed, Washington State, USA. Simulated clearcutting patterns consisted of 7, 9 or 26 ha square or rectangular harvest units distributed in either an aggregated or dispersed pattern under three stream-buffering scenarios. The slope-stability model predicted that potentially unstable ground is concentrated along steep headwater streams and inner-gorge side-slopes. Areas susceptible to wind damage were determined from the combination of slope, aspect, elevation, soil drainage and primary tree species. Among the variables examined here, the location of harvest units constitutes the most important factor influencing the potential for shallow landsliding. In contrast, the location, size, and shape of clear cuts and the interactions among these three factors significantly influenced the potential for wind damage. Minimal correspondence between areas predicted to be potentially unstable and areas susceptible to wind damage implies that harvest patterns designed to mitigate the potential for shallow landsliding may not necessarily reduce the potential for wind damage. Our results demonstrate that: (1) the location of timber harvesting is more important than the geometry of harvest activity in influencing shallow landsliding; (2) forest harvest patterns strongly influence the potential for disturbance processes; and (3) a single cutting pattern will often fail to meet all landscape management goals. This revised version was published online in July 2006 with corrections to the Cover Date.     

Weak effect of edges on avian nesting success in fragmented and forested landscapes in Ontario,Canada

We studied the effects of anthropogenic edges on predation and parasitism of forest bird nests in an agriculturally fragmented landscape and a continuously forested landscape in Ontario, Canada. Nesting data were collected at 1937 nests across 10 species in the fragmented landscape from 2002–2008, and 464 nests across 4 species in the continuously forested landscape from 2006–2008. Brood parasitism only occurred in the fragmented landscape, and was positively related to the proportion of rural grassland and row crop habitats within 500-m of nests. Daily nest survival was negatively related to the density of roads within 500-m of nests in the fragmented landscape, but was not influenced by distance to anthropogenic edge in either landscape. Predation rates were higher in the fragmented landscape for Ovenbird and Rose-breasted Grosbeak nests, but did not differ between landscapes for Veery and American Redstart nests. Uniformly high predation in the fragmented landscape may be a result of (1) matrix predators that penetrate deep (>300 m) into the forest interior, or (2) the additive effect of forest-dependent and matrix-associated predators that results in high predation pressure in both edge and interior habitats. Further research focused on the identification of nest predators, their population dynamics, and habitat use is required to understand the underlying mechanisms leading to uniformly high nest predation in fragmented landscapes.     

森林公园植物景观改造提升思路

本文以广东省国有林场建立的森林公园植物景观为典型,对森林公园绿化景观中普遍存在的问题进行了归纳,提出了诸多改造思路。森林植物景观要通过合理的选择、配置、改造,才能为游客提供独具特色的森林景观,让人们尊享健康、收获视觉美感,让森林公园有良好的经济收益和社会效益。     

Rates and patterns of landscape change between 1972 and 1988 in the Changbai Mountain area of China and North Korea

Satellite imagery was used to quantify rates and patterns oflandscape change between 1972 and 1988 in the Changbai MountainReserve and its adjacent areas in the Peoples Republic of Chinaand North Korea. The 190,000 ha Reserve was established as anInternational Biosphere Reserve by The United Nations Educational,Scientific and Cultural Organization (UNESCO) in 1979. It is themost important natural landscape remaining in Chinastemperate/boreal climate. The images used in this research cover atotal area of 967,847 ha, about three-fourths of which is in China.Imagery from 1972 and 1988 was classified into 2 broad cover types(forest and non-forest). Overall, forests covered 84.4% of thestudy area in 1972 and 74.5% in 1988. Changes in forest coverwithin the Reserve were minimal. The loss of forest cover outsidethe Reserve appears to be strongly associated with timberharvesting at lower elevations. Landscape patterns in 1988 weremore complex, more irregular, and more fragmented than in 1972.This is one of the few studies to assess landscape changes acrosstwo countries. The rates and patterns of forest-cover loss weredifferent in China and North Korea. In North Korea, extensivecutting appears to have occurred prior to 1972 and this hascontinued through 1988 while in China, most cutting appears to haveoccurred since 1972.     

Human influence on the abundance and connectivity of high-risk fuels in mixed forests of northern Wisconsin,USA

Though fire is considered a natural disturbance, humans heavily influence modern wildfire regimes. Humans influence fires both directly, by igniting and suppressing fires, and indirectly, by either altering vegetation, climate, or both. We used the LANDIS disturbance and succession model to compare the relative importance of a direct human influence (suppression of low intensity surface fires) with an indirect human influence (timber harvest) on the long-term abundance and connectivity of high-risk fuel in a 2791 km² landscape characterized by a mixture of northern hardwood and boreal tree species in northern Wisconsin. High risk fuels were defined as a combination of sites recently disturbed by wind and sites containing conifer species/cohorts that might serve as ladder fuel to carry a surface fire into the canopy. Two levels of surface fire suppression (high/current and low) and three harvest alternatives (no harvest, hardwood emphasis, and pine emphasis) were compared in a 2×3 factorial design using 5 replicated simulations per treatment combination over a 250-year period. Multivariate analysis of variance indicated that the landscape pattern of high-risk fuel (proportion of landscape, mean patch size, nearest neighbor distance, and juxtaposition with non fuel sites) was significantly influenced by both surface fire suppression and by forest harvest (p > 0.0001). However, the two human influences also interacted with each other (p < 0.001), because fire suppression was less likely to influence fuel connectivity when harvest disturbance was simultaneously applied. Temporal patterns observed for each of seven conifer species indicated that disturbances by either fire or harvest encouraged the establishment of moderately shade-tolerant conifer species by disturbing the dominant shade tolerant competitor, sugar maple. Our results conflict with commonly reported relationships between fire suppression and fire risk observed within the interior west of the United States, and illustrate the importance of understanding key interactions between natural disturbance, human disturbance, and successional responses to these disturbance types that will eventually dictate future fire risk.This revised version was published online in May 2005 with corrections to the Cover Date.     

Nesting Success of a Songbird in a Complex Floodplain Forest Landscape in llinois, USA: Local Fragmentation vs. Vegetation Structure

Measuring edge effects in complex landscapes is often confounded by the presence of different kinds of natural and anthropogenic edges, each of which may act differently on organisms inhabiting habitat patches. In such landscapes, proportions of different habitats surrounding nests within patches often vary and may affect nesting success independently of distance to edges. We developed methods to measure and study the effects of multiple edges and varying habitat composition around nests on the breeding success of the Acadian flycatcher (Empidonax virescens), an understory, open-cup nesting songbird. The Kaskaskia River in Southwestern Illinois was our study area and consists of wide (>1000-m) floodplain corridors embedded in an agricultural matrix with a variety of natural (wide rivers, backwater swamps, and oxbow lakes) and anthropogenic (internal openings, and agricultural) habitats. We also measured vegetation structure around each nest. Nest survival increased with increasing nest concealment, and probabilities of brood parasitism increased with increasing distances from anthropogenic and natural water-related openings surrounding nests. The magnitude of these effects was small, probably because the landscape is saturated with nest predators and brood parasites. These results illustrate the importance of considering both larger landscape context and details of natural and anthropogenic disturbances when studying the effects of habitat fragmentation on wildlife.     

Edge effects in tropical dry forests of Madagascar: additivity or synergy?

Context

The study of habitat fragmentation is complex because multiple, potentially synergistic, ecological processes may be acting simultaneously. Further, edge effects themselves may be complex in that additivity from multiple edges can give rise to heterogeneous nearest–edge gradients.

Objectives

We used heat diffusion as a proxy for additive edge effects in two study landscapes in order to test whether two key observations recently attributed to synergy between edge and area effects could be more simply explained by additivity; namely, steeper edge gradients in larger fragments and variation in slopes of species–area relationships as a function of distances to fragment edges.

Methods

We sampled forest structure in northwestern Madagascar at various distances from the edge in fragments and continuous forest and used an inverse modelling approach to parameterize the model. In addition, we applied the model to data from a published study of beetle communities in fragmented forests in New Zealand.

Results

With increasing proximity to edges, woody stem densities decreased and, as predicted, smaller fragments had lower stem densities and less steep edge gradients than larger ones. The model successfully predicted shifts in species–area relationships as a function of nearest–edge distances for beetle species, although observed richness for forest specialists in the smallest fragments was lower than predicted.

Conclusions

Two key observations attributed to synergy between edge and area effects were explained by edge additivity. The model is particularly useful in that it can help to disentangle the complex sets of processes acting in fragmented landscapes.

     

Spatial dynamics of coastal forest bird assemblages: the influence of landscape context,forest type,and structural connectivity

Context

Complex structural connectivity patterns can influence the distribution of animals in coastal landscapes, particularly those with relatively large home ranges, such as birds. To understand the nuanced nature of coastal forest avifauna, where there may be considerable overlap in assemblages of adjacent forest types, the concerted influence of regional landscape context and vegetative structural connectivity at multiple spatial scales warrants investigation.

Objectives

This study determined whether species compositions of coastal forest bird assemblages differ with regional landscape context or with forest type, and if this is influenced by structural connectivity patterns measured at multiple spatial scales.

Methods

Three replicate bird surveys were conducted in four coastal forest types at ten survey locations across two regional landscape contexts in northeast Australia. Structural connectivity patterns of 11 vegetation types were quantified at 3, 6, and 12 km spatial scales surrounding each survey location, and differences in bird species composition were evaluated using multivariate ordination analysis.

Results

Bird assemblages differed between regional landscape contexts and most coastal forest types, although Melaleuca woodland bird assemblages were similar to those of eucalypt woodlands and rainforests. Structural connectivity was primarily correlated with differences in bird species composition between regional landscape contexts, and correlation depended on vegetation type and spatial scale.

Conclusions

Spatial scale, landscape context, and structural connectivity have a combined influence on bird species composition. This suggests that effective management of coastal landscapes requires a holistic strategy that considers the size, shape, and configuration of all vegetative components at multiple spatial scales.

     

Limiting factors and landscape connectivity: the American marten in the Rocky Mountains

In mobile animals, movement behavior can maximize fitness by optimizing access to critical resources and minimizing risk of predation. We sought to evaluate several hypotheses regarding the effects of landscape structure on American marten foraging path selection in a landscape experiencing forest perforation by patchcut logging. We hypothesized that in the uncut pre-treatment landscape marten would choose foraging paths to maximize access to cover types that support the highest density of prey. In contrast, in the post-treatment landscapes we hypothesized marten would choose paths primarily to avoid crossing openings, and that this would limit their ability to optimally select paths to maximize foraging success. Our limiting factor analysis shows that different resistant models may be supported under changing landscape conditions due to threshold effects, even when a species’ response to landscape variables is constant. Our results support previous work showing forest harvest strongly affects marten movement behavior. The most important result of our study, however, is that the influence of these features changes dramatically depending on the degree to which timber harvest limits available movement paths. Marten choose foraging paths in uncut landscapes to maximize time spent in cover types providing the highest density of prey species. In contrast, following landscape perforation by patchcuts, marten strongly select paths to avoid crossing unforested areas. This strong response to patch cutting reduces their ability to optimize foraging paths to vegetation type. Marten likely avoid non-forested areas in fragmented landscapes to reduce risk of predation and to benefit thermoregulation in winter, but in doing so they may suffer a secondary cost of decreased foraging efficiency.     

Divergent effects of tropical forest fragmentation and conversion on leaf litter decomposition

Context

In tropical landscapes, dominant land-use changes involve conversion of intact forest to an agricultural matrix with embedded fragments of remnant forest. However, most research to date has focused on how these land-use changes affect species within the fragmented ecosystem, rather than the flux of energy and nutrients within these different landscape elements.

Objectives

We examined how forest fragmentation and conversion to orange fields impact the potential for litter decomposition in a Costa Rican landscape, in particular via effects on macroinvertebrates (MIs) and microclimate.

Methods

We measured mass losses of a standard leaf litter in four habitats: orange fields, small forest fragments, large forest fragments and intact forest. Litter bags were constructed of mesh that either excluded or allowed MIs. Decomposition rates were measured in wet and dry seasons, and at different distances from the forest edge.

Results

Forest fragmentation and forest conversion had divergent effects on decomposition rates. Decomposition rates were 7 % slower in forest fragments during the dry season than in intact forest, and this result was mediated by forest fragmentation effects on MIs. Decomposition rates were 9 % higher in orange fields during the wet season, relative to intact forest, and this pattern was explained by effects of the litter microenvironment on leaching rates or smaller invertebrates. Fragment area and distance from forest edge had minor or undetectable effects on decomposition in fragments.

Conclusions

We conclude that land-use changes affect decomposition processes in both forest and agroecosystems, and these effects can vary in mechanism and direction across disturbed landscapes.