A positive response of mountain pine beetle to pine forest-clearcut edges at the landscape scale in British Columbia,Canada

In British Columbia, large-scale salvage harvesting has been underway to recover timber value from forest stands infested by mountain pine beetle during the current outbreak. Understanding the response of beetles to clearcut edges particularly at the landscape scale is crucial to understanding the impacts of increased habitat fragmentation due to salvage harvesting on the spread of the beetle infestations. A novel proximity analysis approach based on null models of complete spatial randomness with three different spatial extents was developed to examine the spatial patterns of infestations in relation to cutblocks. Inhomogeneous Poisson point process models were fitted to predict how intensities of infestations varied with distances to the nearest cutblocks. Marked Poisson point process models were also fitted to evaluate the effects of the variables associated with the nearest cutblocks and adjacent infested pine stands on the edge response of beetles. The results clearly illustrated a significant positive edge response of beetles at the landscape scale. The intensities of infestations decreased non-linearly with distances to the nearest cutblocks. The results also suggested that the quality and distribution of key habitat resources could not fully explain the fundamental mechanisms underlying the edge response. The behavioural change of beetle dispersal at edges may also be an important factor contributing to a positive edge response. The results from this study may be useful in improving the efficacy of mountain pine beetle management efforts.     

Predictors of bark beetle activity and scale-dependent spatial heterogeneity change during the course of an outbreak in a subalpine forest

Climate conditions and forest structure interact to determine the extent and severity of bark beetle outbreaks, yet the relative importance of each may vary though the course of an outbreak. In 2008, we conducted field surveys and reconstructed forest conditions at multiple stages within a recent mountain pine beetle (MPB) outbreak in Rocky Mountain National Park, Colorado. At each stage in the outbreak, we examined changes in (1) lodgepole pine mortality and surviving stand structure, (2) the influence of topographic versus stand structure variables on mortality rates, and (3) stand complexity and landscape heterogeneity. Lodgepole pine mortality reduced basal area by 71 %, but only 47 % of stems were killed. Relative to pre-outbreak stands, surviving stands had lower mean dbh (11.0 vs. 17.4 cm), lower basal area (8.5 vs. 29.3 m² ha^?1), lower density (915 vs. 1,393 stems ha^?1), and higher proportions of non-host species (23.1 vs. 10.6 % m² ha^?1). Factors predicting mortality rates changed through the course of the outbreak. Tree mortality during the early stage of the outbreak was associated with warm, dry sites and abundant large trees. During the middle and late stages, mortality was associated with stand structure alone. Stand complexity increased, as defined by stand-scale variability in density, basal area, and the proportion of susceptible trees. Landscape heterogeneity decreased according to semi-variograms of tree diameter and basal area. Increased stand complexity may inhibit future MPB population development, but decreased landscape heterogeneity may facilitate outbreak spread across the landscape if a future outbreak were to irrupt.     

Disturbance propagation by bark beetles as an episodic landscape phenomenon

Landscapes are the resultant of ecological processes and events operating on many different space-time scales. Large scale disturbance is recognized as a major influence on landscape patterns, but the impact of small scale events is often overlooked. We develop an hierarchical framework to relate lightning and bark beetle population dynamics to the southern pine forest landscape using the concepts of disturbance propagation and amplification. The low level lightning disturbance can be propagated to the landscape level when weather and forest stand structure facilitate bark beetle epidemics. We identify epidemics as biotically-driven episodes that alter landscape structure. The concept of the landscape as the spatial dimension of these episodes is represented in a conceptual model linking insect-host and landscape mosaic interactions.     

Climate change amplifies the interactions between wind and bark beetle disturbances in forest landscapes

Context

Growing evidence suggests that climate change could substantially alter forest disturbances. Interactions between individual disturbance agents are a major component of disturbance regimes, yet how interactions contribute to their climate sensitivity remains largely unknown.

Objectives

Here, our aim was to assess the climate sensitivity of disturbance interactions, focusing on wind and bark beetle disturbances.

Methods

We developed a process-based model of bark beetle disturbance, integrated into the dynamic forest landscape model iLand (already including a detailed model of wind disturbance). We evaluated the integrated model against observations from three wind events and a subsequent bark beetle outbreak, affecting 530.2 ha (3.8 %) of a mountain forest landscape in Austria between 2007 and 2014. Subsequently, we conducted a factorial experiment determining the effect of changes in climate variables on the area disturbed by wind and bark beetles separately and in combination.

Results

iLand was well able to reproduce observations with regard to area, temporal sequence, and spatial pattern of disturbance. The observed disturbance dynamics was strongly driven by interactions, with 64.3 % of the area disturbed attributed to interaction effects. A +4 °C warming increased the disturbed area by +264.7 % and the area-weighted mean patch size by +1794.3 %. Interactions were found to have a ten times higher sensitivity to temperature changes than main effects, considerably amplifying the climate sensitivity of the disturbance regime.

Conclusions

Disturbance interactions are a key component of the forest disturbance regime. Neglecting interaction effects can lead to a substantial underestimation of the climate change sensitivity of disturbance regimes.

     

Interactions of landscape disturbances and climate change dictate ecological pattern and process: spatial modeling of wildfire,insect, and disease dynamics under future climates

Context

Interactions among disturbances, climate, and vegetation influence landscape patterns and ecosystem processes. Climate changes, exotic invasions, beetle outbreaks, altered fire regimes, and human activities may interact to produce landscapes that appear and function beyond historical analogs.

Objectives

We used the mechanistic ecosystem-fire process model FireBGCv2 to model interactions of wildland fire, mountain pine beetle (Dendroctonus ponderosae), and white pine blister rust (Cronartium ribicola) under current and future climates, across three diverse study areas.

Methods

We assessed changes in tree basal area as a measure of landscape response over a 300-year simulation period for the Crown of the Continent in north-central Montana, East Fork of the Bitterroot River in western Montana, and Yellowstone Central Plateau in western Wyoming, USA.

Results

Interacting disturbances reduced overall basal area via increased tree mortality of host species. Wildfire decreased basal area more than beetles or rust, and disturbance interactions modeled under future climate significantly altered landscape basal area as compared with no-disturbance and current climate scenarios. Responses varied among landscapes depending on species composition, sensitivity to fire, and pathogen and beetle suitability and susceptibility.

Conclusions

Understanding disturbance interactions is critical for managing landscapes because forest responses to wildfires, pathogens, and beetle attacks may offset or exacerbate climate influences, with consequences for wildlife, carbon, and biodiversity.

     

遵义红军山松材线虫病预防与对策

遵义红军山和红花岗风景林坐落于遵义市区中心,山上有集中连片的马尾松林570．33hm^2,主要树种为马尾松,与遵义县龙坑镇松材线虫病疫点直线距离不足15km。该片松林所处的政治地位、生态地位、社会地位都具有特别重要的位置,对于如何保护好山上的一草一木,使该片马尾松林免遭松材线虫病的毁灭性危害,笔者提出预防设想与对策。     

Topography-mediated controls on local vegetation phenology estimated from MODIS vegetation index

Forest canopy phenology is an important constraint on annual water and carbon budgets, and responds to regional interannual climate variation. In steep terrain, there are complex spatial variations in phenology due to topographic influences on microclimate, community composition, and available soil moisture. In this study, we investigate spatial patterns of phenology in humid temperate forest as a function of topography. Moderate-resolution imaging spectro-radiometer (MODIS) vegetation indices are used to derive local patterns of topography-mediated vegetation phenology using a simple post-processing analysis and a non-linear model fitting. Elevation has the most explanatory power for all phenological variables with a strong linear relationship with mid-day of greenup period, following temperatures lapse rates. However, all other phenological variables show quadratic associations with elevation, reflecting an interaction between topoclimatic patterns of temperature and water availability. Radiation proxies also have significant explanatory power for all phenological variables. Though hillslope position cannot be adequately resolved at the MODIS spatial resolution (250 m) to discern impacts of local drainage conditions, extended periods of greenup/senescence are found to occur in wet years. These findings are strongly supported by previous field measurements at different topographic positions within the study area. The capability of detecting topography-mediated local phenology offers the potential to detect vegetation responses to climate change in mountainous terrain. In addition, the large, local variability of meteorological and edaphic conditions in steep terrain provides a unique opportunity to develop an understanding of canopy response to the interaction of climate and landscape conditions.     

Spatial patterns of primary productivity in the Greater Yellowstone Ecosystem

Landscapes are often heterogeneous in abiotic factors such as topography, climate, and soil, yet little is known about how these factors may influence the spatial distribution of primary productivity. We report estimates of aboveground net primary productivity (ANPP) in 90 sample stands stratified by cover type and elevation class, and use the results to predict ANPP across a portion of the Greater Yellowstone Ecosystem. Tree ANPP was estimated by sampling tree density by species and diameter classes and estimating average annual diameter increment by tree coring. Biomass for current tree diameter and past tree diameter were calculated by species and diameter class for each stand using the dimension analysis software BIOPAK. Shrub ANPP was estimated by calculating current biomass from basal area using BIOPAK and dividing by the assumed average life span of the shrubs. Clipping at the end of the growing season was used to estimate herb ANPP. Differences in ANPP among cover types and elevation classes were examined with analysis of variance. Multiple regression was used to examine relationships between ANPP, and soil parent material, topography, and cover type. The best regression model was used to predict ANPP across the study area.We found ANPP was highest in cottonwood, Douglas-fir, and aspen stands, intermediate in various seral stages of lodgepole pine, and lowest in grassland and sagebrush cover types. Parent material explained significant variation in ANPP in mature and old-growth lodgepole pine stands, with rhyolite ash/loess being the most productive parent material type. ANPP decreased with increasing elevation in most cover types, possibly because low temperatures limit plant growth at higher elevations in the study area. ANPP was not related to elevation in mature and old-growth lodgepole pine stands, due to relatively rapid growth of subalpine fir at higher elevations.A regression model based on cover type and elevation explained 89% of the variation in ANPP among the sample stands. This model was used to generate a spatially continuous surface of predicted ANPP across the study area. The frequency distribution of predicted ANPP was skewed towards lower levels of ANPP, and only 6.3% of the study area had a predicted ANPP level exceeding 4500 kg/ha/yr. Patches high in predicted ANPP were primarily at lower elevations, outside of Yellowstone National Park, and near the national forest/private lands boundary. These patterns of ANPP may influence fire behavior, vertebrate population dynamics, and other ecological processes. The results reinforce the need for coordinated management across ownership boundaries in the Greater Yellowstone Ecosystem.     

Reconstructing Landscape-scale Tree Invasion Using Survey Notes in the Medicine Bow Mountains, Wyoming, USA

We assessed landscape-scale invasions of openings in mountain forests by native tree species since EuroAmerican settlement (ca. 1870–1899). We reconstructed historical openings across a 250,240 ha area in the Medicine Bow Mountains, Wyoming, using notes from the original General Land Office (GLO) surveys, and compared historical openings to modern openings interpreted from digital aerial photography. We constructed logistic regression models to describe and predict tree invasion, based on a set of environmental and land use predictors. Openings have decreased by about 7.3% in the last ca. 110 years. Invasion was more likely to occur on moister sites, indicated by high values for steady-state wetness, low values for evaporative demand, proximity to streams, and topographic settings in basins or channels. More invasion also occurred on unprotected public land, in openings surrounded by lodgepole pine and aspen, and on mesic soils. The relatively slow rates of invasion in the study area may be driven by climate and land use.     

Influence of a shrub corridor on movements of passerine birds to a lake littoral zone

A pine forest was separated from a lake littoral zone by a meadow on one area (discontinuous) while these habitats were separated by a shrub strip in another area (continuous). This shrub strip acted as an ecological corridor enhancing the movements of birds between the forest and the littoral reed zone. The number of individuals of non-littoral species that visited the reed zone was higher (p < .001) on the area with the connecting shrub strip in autumn but the number of species visiting the littoral zone was not significantly higher. Significantly more (p < .001) autumn movements by birds in the continuous area were oriented along paths between the forest and the littoral zone whereas movements in the discontinuous area paralleled the littoral and forest zones (p < .001). Movements of birds were concentrated along the edge of the shrub strip. The spatial configuration of the landscape facilitated access by some forest birds to the littoral habitat.     

Strategic removal of host trees in isolated,satellite infestations of emerald ash borer can reduce population growth

Emerald ash borer is an invasive beetle causing significant mortality of ash trees (Fraxinus spp.) in North America and western Russia. The invasive range has expanded to more than half of the states in the United States since the initial detection in Michigan, USA in 2002. Emerald ash borer is typically managed with a combination of techniques including surveys/trapping, insecticide treatments, host tree removal, biological control, and public education/outreach. The insect’s rapid spread rate and cryptic life history and a lack of resistance among most North American Fraxinus spp. have limited opportunities to gather empirical data on how aggressive tree removal may slow population growth in isolated, satellite infestations if detected early. An early detection of an isolated population of emerald ash borer in 2009 in Minnesota, USA was managed by using a selective host-tree removal program (i.e., sanitation). Trees were preferentially removed based on the assumption that evidence of woodpecker foraging (i.e., pecking) was a good indicator of infestation by emerald ash borer. Extensive sampling and survey data on larval densities and the presence/absence of pecking on ash trees in a 6-km² area for the Twin Cities, Minnesota were used to parameterize a model of population growth over the next four years. We found that removing ∼63% of the total trees across four years reduced the cumulative number of beetles produced in the core infested area by ∼54%. However, we also found that increases in efficacy, i.e., larger decreases in beetle production per removed tree, could be achieved by preferentially removing trees with pecking. The invasive range of emerald ash borer in North America and western Russia continues to expand via natural and human-aided dispersal. While silvicultural control tactics alone will not be an adequate management strategy, tree removal is an important component of both a broader pest management program and the systematic replacement of ash canopies in urban forests. Increasing understanding of the efficacy of different management techniques in slowing population growth of emerald ash borer will be useful to support decision–making by land managers.     

Post-storm surveys reveal large-scale spatial patterns and influences of site factors,forest structure and diversity in endemic bark-beetle populations

The storm that struck France on december 26^th and 28^th 1999 felled 140 million m³ of timber and had a high economic, social and landscape impact. This event offered the opportunity to study large-scale patterns in populations of forest insect pests that would benefit from the abundant breeding material. A large-scale survey was carried out in France in 2000 to sample the most frequently observed species developing on spruce (Ips typographus, Pityogene schalcographus) and pine (Tomicus piniperda, Ips sexdentatus) in 898 locations distributed throughout wind-damaged areas. The local abundance of each species scored on a 0 to 5 scale was analysed using geostatistical estimators to explore the extent and intensity of spatial autocorrelation, and was related to site, stand, and neighbourhood landscape metrics of the forest cover (in particular the interconnection with broadleaf forest patches) found within dispersal distance. All species but I. sexdentatus, which was much less abundant, displayed large-scale spatial dependence and regional variations in abundance. Lower infestation levels per tree (windfalls and standing trees) were observed in stands with a high proportion of wind-damaged trees, which was interpreted as the result of beetles distributing themselves among the available breeding material. More infestations were observed in wind-broken trees as compared to wind-felled trees. More importantly, populations showed significant relationships with the structure of coniferous stands (in particular with the number of coniferous patches). T. piniperda population levels were negatively correlated to the amount of coniferous edge shared with broadleaf forest patches, possibly because of the disruptive effect of non-host volatiles on host-finding processes at the landscape-scale. The differences observed between species regarding patterns and relationships to site, stand, and forest cover characteristics are discussed in relation to the ecological characteristics of each species.     

Incoming! Association of landscape features with dispersing mountain pine beetle populations during a range expansion event in western Canada

Mountain pine beetle, Dendroctonus ponderosae Hopkins (Coleoptera: Curculionidae, Scolytinae), is a forest insect that undergoes intermittent population eruptions, causing landscape-level mortality to mature pines. Currently, an outbreak covers over 16.3 million ha of British Columbia and Alberta in western Canada. Recent incursion into the jack pine (Pinus banksiana Lamb.) of northwestern Alberta threatens further range expansion through the boreal forest to central and eastern Canada. The spread from British Columbia into northwestern Alberta has been facilitated by above-canopy dispersal of the insect by meso-scale atmospheric currents. At these scales, dispersing D. ponderosae may behave like inert particles, causing terrain-induced tropospheric convective and advective currents to influence population dispersal and establishment. We use spatial point process regression models to examine the association of meso-scale variables, including landscape features and their orientations, habitat suitability, elevation and treatment efforts, with occurrence of D. ponderosae infestations in 2004, 2005, and 2006. Infestations of D. ponderosae primarily established in canyons and valleys, before moving into more open-sloped areas. Southwestern slopes of midslope ridges and small hills, southwest facing open slopes, and valleys that run in a northeast–southwest cardinal direction were positively associated with higher intensities of infestation. This study provides insight into the influences of complex terrain on landscape disturbance by a forest insect, and can be used to prioritize areas for potential management.     

Predicting grey squirrel expansion in North Italy: a spatially explicit modelling approach

There is growing concern about the spread of the North American grey squirrel (Sciurus carolinensis) in northern Italy which were introduced into Piedmont in 1948. They have since spread across the Po-plain covering an area of approximately 450 km² and continue to expand their range. In parallel to what has been observed in Britain and Ireland, grey squirrels replace the native red squirrel (S. vulgaris) and damage poplar (Populus) plantations through bark-stripping. Spatially explicit population dynamics models have been successfully used to predict the spread of grey squirrels in East Anglia, England. We extended a previous approach employing a sensitivity analysis where life history and other demographic inputs are generated using Latin Hypercube Sampling from the known ranges of each input parameter, and applied it to Italy using field data collected in Piedmont. The analysis indicated that reproductive output was the most important factor determining total population size present in Piedmont. The structure and composition of woodland habitats around the introduction site suggested that initial grey squirrel expansion would have been slow and subject to emigration rates from the available habitat blocks.A comparison of the 1996 survey results with model predictions indicated that a mean litter size of three young gave the best fit with the observed distribution and we use this to predict future grey squirrel spread. We also present a worst case scenario in which grey squirrels experience improved reproductive success due to the availability of high quality habitats beyond the Po plain. In both cases they could disperse along existing continuous woodland corridors into France between 2039–2048. The case of the grey squirrel highlights the problems of implementing conservation conventions and the resulting conflicts between wildlife management, public perception and local political support and the narrow time frame that is available to control alien species effectively before it is too late. If allowed to spread, grey squirrels have the potential of becoming a European forest pest species and are likely to replace the native red squirrel in large parts of its range.     

X-ray Imaging of Stored Dates to Detect Infestation by Saw-Toothed Beetles

The objective of this study was to investigate the capability of X-ray imaging in detecting internal infestations caused by the saw-toothed beetle (Oryzaephilus surinamensis L.) in stored dates. Un-infested and infested dates were x-rayed for different life stages (egg, larvae, pupae, and adult) of O. surinamensis. An algorithm was developed to extract 44 features from the X-ray images. Linear discriminant analysis (LDA) was used to discriminate the un-infested and infested dates using extracted features. Overall accuracy of X-ray imaging system was 80% for un-infested and beetle-infested dates. X-ray imaging system yielded around 97% accuracy in detecting internal infestation of dates with an adult beetle while using a pairwise classification method. Further studies are required to investigate the effects of moisture content and hardness of dates in the discrimination between the un-infested and infested dates.     

Effects of habitat loss on populations of white-footed mice: testing matrix model predictions with landscape-scale perturbation experiments

Habitat loss is the leading cause of decline in wildlife diversity and abundance throughout the world, but its effects on wildlife are not always predictable. Matrix population modeling is an increasingly common tool used to predict the effects of habitat loss. In spite of the growing number of studies using this approach, and its wide use in conservation practice, the predictions generated by matrix population models are rarely explicitly tested in the field. We compared the ability of a suite of spatially explicit demographic matrix models to predict the response of white-footed mice to loss of high quality habitat at mosaic sites in northeast Connecticut, USA. We tested short-term model predictions with landscape scale habitat perturbation experiments, including clear-cut logging or prescribed burning of high quality habitat at two study sites. Comparison of each model’s predictions with the observed responses at both sites qualitatively supported predictions that perturbation of high quality habitat would have negative effects extending into the surrounding landscape. The best-supported model assumed that evicted residents of the perturbed habitat would successfully resettle in nearby intact habitats, and allowed for gradual population recovery in the perturbed habitat. Similarly, long-term simulations (20 years) revealed how loss of a single habitat could trigger population declines throughout a mosaic site. This study shows that careful consideration of model assumptions such as those pertaining to resettlement behavior is crucial if predictions are to be reliable, and highlights the role of experiments for comparing alternative model predictions.     

Relating Kirtland's warbler population to changing landscape composition and structure

The population of male Kirtland's warbler (Dendroica kirtlandii) in the breeding season has averaged 206 from 1971 to 1987. The Kirtland's warbler occupies dense jack pine (Pinus banksiana) barrens from 5 to 23 years old and from 1.4 to 5.0 m high, formerly of wildfire origin. In 1984, 73% of the males censused were found in habitat naturally regenerated from wildfire or prescribed burning. The rest were in plantations (11%) or in harvested, unburned jack pine stands stocked by natural regeneration (16%). Twenty-two percent (630 of 2,886) of the Kirtland's warbler males counted in the annual censuses from 1971 through 1984 were found in 26 stands that were unburned and naturally regenerated following harvest. From 1982 to 1987, suitable regenerating areas were barely sufficient to replace currently occupied maturing stands, so population growth was impeded. Ecosystems of suitable size and regeneration characteristics (wildfire and plantation) doubled in area by 1989. In response, the population of Kirtland's warblers increased from 167 to 398 males between 1987 and 1992, but they withdrew almost entirely from the unburned, unplanted barrens by 1989 when the area of more suitable regeneration types increased. Minimum (368 males) and maximum (542 males) population estimates for 1996 were calculated based on 1984 average density (1.9 males per 40 ha) and peak population in burns (2.8 males per 40 ha).     

Fire severity and seed source influence lodgepole pine (<Emphasis Type="Italic">Pinus contorta</Emphasis> var. <Emphasis Type="Italic">murrayana</Emphasis>) regeneration in the southern cascades,Lassen volcanic National Park,California

Rocky Mountain lodgepole pine, (Pinus contorta var. latifolia) regenerates quickly after high severity fire because seeds from serotinous cones are released immediately post-fire. Sierra lodgepole pine (P. contorta var. murrayana) forests burn with variable intensity resulting in different levels of severity and because this variety of lodgepole pine does not have serotinous cones, little is known about what factors influence post-fire regeneration. This study quantifies tree regeneration in a low, moderate, and high severity burn patch in a Sierra lodgepole forest 24 years after fire. Regeneration was measured in ten plots in each severity type. In each plot, we quantified pre- and post-fire forest structure (basal area, density), counted and aged tree seedlings and saplings of all species, and measured distance to the nearest seed bearing tree. There was no difference in the density of seedlings and saplings among severity classes. Distance and direction to the nearest seed bearing lodgepole pine were the best predictors of lodgepole seedling and sapling density in high severity plots. In contrast to Rocky Mountain lodgepole pine, regeneration of Sierra lodgepole pine appears to rely on in-seeding from surviving trees in low or moderate severity burn patches or live trees next to high severity burn patches. Our data demonstrate that Sierra lodgepole pine follows stand development pathways hypothesized for non-serotinous stands of Rocky Mountain lodgepole pine.     

低温逆境下温室茄子专用品种的筛选

以河西地区当地主栽品种紫长茄为对照,从国内外引进4个不同的茄子品种.幼苗在低温逆境下处理2 d,处理后1、3、5、7 d观察幼苗的形态变化和生长情况,测定其株高、茎粗、叶面积、根系活力、叶绿素含量.处理后的幼苗按常规方法定植,分别观察各参试品种的物候期、早期产量和总产量.结果表明:兰杂二号在低温逆境后,形态指标和生理指标均优于其他参试品种,结果产量最高.     

Simulating the reciprocal interaction of forest landscape structure and southern pine beetle herbivory using LANDIS

The reciprocal interaction of landscape structure and ecological processes is a cornerstone of modern landscape ecology. We use a simulation model to show how landscape structure and herbivory interact to influence outbreaks of southern pine beetle (Dendroctonus frontalis Zimmermann) in a landscape representative of the southern Appalachian Mountains, USA. We use LANDIS and its biological disturbance agent module to simulate the effects of landscape composition (proportion of landscape in host area) and host aggregation on the size and severity of insect outbreaks and the persistence of the host species, Table Mountain Pine (Pinus pungens Lamb.). We find that landscape composition is less important in the modeled landscapes than host aggregation in structuring the severity of insect outbreaks. Also, simulated southern pine beetle outbreaks over time tend to decrease the aggregation of host species on the landscape by fragmenting large patches into smaller ones, thereby reducing the severity of future outbreaks. Persistence of Table Mountain pine decreases throughout all simulations regardless of landscape structure. The results of this study indicate that when considering alternative restoration strategies for insect-affected landscapes, it is necessary to consider the patterns of hosts on the landscape as well as the landscape composition.