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Theory and application of semiochemicals in nuisance fish control

Controlling unwanted, or nuisance, fishes is becoming an increasingly urgent issue with few obvious solutions. Because fish rely heavily on semiochemicals, or chemical compounds that convey information between and within species, to mediate aspects of their life histories, these compounds are increasingly being considered as an option to help control wild fish. Possible uses of semiochemicals include measuring their presence in water to estimate population size, adding them to traps to count or remove specific species of fish, adding them to waterways to manipulate large-scale movement patterns, and saturating the environment with synthesized semiochemicals to disrupt responses to the natural cue. These applications may be especially appropriate for pheromones, chemical signals that pass between members of same species and which also have extreme specificity and potency. Alarm cues, compounds released by injured fish, and cues released by potential predators also could function as repellents and be especially useful if paired with pheromonal attractants in “push-pull” configurations. Approximately half a dozen attractive pheromones now have been partially identified in fish, and those for the sea lamprey and the common carp have been tested in the field with modest success. Alarm and predator cues for sea lamprey also have been tested in the laboratory and field with some success. Success has been hampered by our incomplete understanding of chemical identity, a lack of synthesized compounds, the fact that laboratory bioassays do not always reflect natural environments, and the relative difficulty of conducting trials on wild fishes because of short field seasons and regulatory requirements. Nevertheless, workers continue efforts to identify pheromones because of the great potential elucidated by insect control and the fact that few tools are available to control nuisance fish. Approaches developed for nuisance fish also could be applied to valued fishes, which suffer from a lack of powerful management tools.

Journal of Chemical Ecology

Movement and feeding ecology of recently emerged steelhead in Lake Ontario tributaries

Steelhead ( Oncorhynchus mykiss ) ascend several Lake Ontario tributaries to spawn and juveniles are often the most abundant salmonid where spawning is successful. Movement and diet of recently emerged subyearling steelhead were examined in three New York tributaries of Lake Ontario. Downstream movement occurred mainly at night and consisted of significantly smaller fry that were feeding at lower levels than resident fry. Fry fed at the highest rate during the day and chironomids and baetids were the main components of their diet. The diet composition of steelhead fry was closely associated with the composition of the benthos in Trout Brook but more similar to the composition of the drift in the other streams. Daily ration was similar among streams, ranging from 10.2 to 14.3%. These findings are consistent with previous findings on the ecology of steelhead fry, as well as fry of other salmonid species

Journal of Applied Ichthyology

A database of natural monthly streamflow estimates from 1950 to 2015 for the conterminous United States

Quantifying and understanding the natural streamflow regime, defined as expected streamflow that would occur in the absence of anthropogenic modification to the hydrologic system, is critically important for the development of management strategies aimed at protecting aquatic ecosystems. Water balance models have been applied frequently to estimate natural flows, but are limited in the number of predictor variables that can be included. Here, a statistical machine learning technique — random forest modeling — was applied to estimate natural flows at a monthly time‐step from 1950 to 2015 for >2.5 million stream reaches in the conterminous United States (U.S.) using 200 potential predictor variables. We describe the development and documentation of this dataset and assess model performance. Model fit statistics (mean Nash–Sutcliffe efficiency = 0.85; observed/expected ratio = 0.94) indicate good correspondence between predicted and observed flows at nearly 2,000 streamgages. As an example application of the dataset, the observed streamflow record at a site prior to and after the construction of an upstream reservoir was compared with estimated natural flows to demonstrate the magnitude of seasonal depletions in streamflow due to the reservoir. This dataset can be applied to quantify natural and anthropogenic processes contributing to streamflow depletion or augmentation, and assess associated ecological effects.

Journal of the American Water Resources Associatio

Computationally efficient statistical differential equation modeling using homogenization

Statistical models using partial differential equations (PDEs) to describe dynamically evolving natural systems are appearing in the scientific literature with some regularity in recent years. Often such studies seek to characterize the dynamics of temporal or spatio-temporal phenomena such as invasive species, consumer-resource interactions, community evolution, and resource selection. Specifically, in the spatial setting, data are often available at varying spatial and temporal scales. Additionally, the necessary numerical integration of a PDE may be computationally infeasible over the spatial support of interest. We present an approach to impose computationally advantageous changes of support in statistical implementations of PDE models and demonstrate its utility through simulation using a form of PDE known as “ecological diffusion.” We also apply a statistical ecological diffusion model to a data set involving the spread of mountain pine beetle (Dendroctonus ponderosae) in Idaho, USA.

Journal of Agricultural, Biological, and Environme

Elk migration influences the risk of disease spillover in the Greater Yellowstone Ecosystem

Wildlife migrations provide important ecosystem services, but they are declining. Within the Greater Yellowstone Ecosystem (GYE) some elk ( Cervus canadensis ) herds are losing migratory tendencies, which may increase spatiotemporal overlap between elk and livestock (domestic bison [ Bison bison ] and cattle [ Bos taurus ]), potentially exacerbating pathogen transmission risk. We combined disease, movement, demographic, and environmental data from eight elk herds in the GYE to examine the differential risk of brucellosis transmission (through aborted fetuses) from migrant and resident elk to livestock. For both migrants and residents, we found that transmission risk from elk to livestock occurred almost exclusively on private ranchlands as opposed to state or federal grazing allotments. Weather variability affected the estimated distribution of spillover risk from migrant elk to livestock, with a 7‐12% increase in migrant abortions on private ranchlands during years with heavier snowfall. In contrast, weather variability did not affect spillover risk from resident elk. Migrant elk were responsible for the majority (68%) of disease spillover risk to livestock because they occurred in greater numbers than resident elk. On a per‐capita basis, however, our analyses suggested that resident elk disproportionately contributed to spillover risk. In five of seven herds, we estimated that the per‐capita spillover risk was greater from residents than from migrants. Averaged across herds, an individual resident elk was 23% more likely than an individual migrant elk to abort on private ranchlands. Our results demonstrate links between migration behavior, spillover risk, and environmental variability, and highlight the utility of integrating models of pathogen transmission and host movement to generate new insights about the role of migration in disease spillover risk. Further, they add to the accumulating body of evidence across taxa that suggests that migrants and residents should be considered separately during investigations of wildlife disease ecology. Finally, our findings have applied implications for elk and brucellosis in the GYE, and suggest that managers should prioritize actions that maintain spatial separation of elk and livestock on private ranchlands during years when snowpack persists into the risk period.

Wyoming

Methods for control of tick vectors of Lyme Borreliosis

During the IVth International Conference on Lyme Borreliosis in Stockholm, 1990, a workshop on control of Lyme disease vectors briefly reviewed: basic ecological principles for tick control; biocontrol of ticks; chemical control, including the use of repellents and use of permethrin-treated rodent nest material; tick control by habitat modification; and reduction of tick host availability. It was concluded that, although much research work remains, Lyme borreliosis is to a large extent a preventable infection. Avoidance of heavily tick-infested areas, personal protection using proper clothing, and prompt removal of attached ticks remain the most effective protective measures. Many other prophylactic measures are available and could be efficiently integrated into schemes to reduce the abundance of vectors. However, since the ecology of the infection varies greatly between different localities it may be necessary to apply different combinations of control methods in different endemic regions.

Scandinavian Journal of Infectious Diseases

Identifying optimal remotely-sensed variables for ecosystem monitoring in Colorado Plateau drylands

Water-limited ecosystems often recover slowly following anthropogenic or natural disturbance. Multitemporal remote sensing can be used to monitor ecosystem recovery after disturbance; however, dryland vegetation cover can be challenging to accurately measure due to sparse cover and spectral confusion between soils and non-photosynthetic vegetation. With the goal of optimizing a monitoring approach for identifying both abrupt and gradual vegetation changes, we evaluated the ability of Landsat-derived spectral variables to characterize surface variability of vegetation cover and bare ground across a range of vegetation community types. Using three year composites of Landsat data, we modeled relationships between spectral information and field data collected at monitoring sites near Canyonlands National Park, UT. We also developed multiple regression models to assess improvement over single variables. We found that for all vegetation types, percent cover bare ground could be accurately modeled with single indices that included a combination of red and shortwave infrared bands, while near infrared-based vegetation indices like NDVI worked best for quantifying tree cover and total live vegetation cover in woodlands. We applied four models to characterize the spatial distribution of putative grassland ecological states across our study area, illustrating how this approach can be implemented to guide dryland ecosystem management.

Colorado Plateau

A sea lamprey ( Petromyzon marinus ) sex pheromone mixture increases trap catch relative to a single synthesized component in specific environments

Spermiating male sea lamprey ( Petromyzon marinus ) release a sex pheromone, of which a component, 7α, 12α, 24-trihydoxy-3-one-5α-cholan-24-sulfate (3kPZS), has been identified and shown to induce long distance preference responses in ovulated females. However, other pheromone components exist, and when 3kPZS alone was used to control invasive sea lamprey populations in the Laurentian Great Lakes, trap catch increase was significant, but gains were generally marginal. We hypothesized that free-ranging sea lamprey populations discriminate between a partial and complete pheromone while migrating to spawning grounds and searching for mates at spawning grounds. As a means to test our hypothesis, and to test two possible uses of sex pheromones for sea lamprey control, we asked whether the full sex pheromone mixture released by males (spermiating male washings; SMW) is more effective than 3kPZS in capturing animals in traditional traps (1) en route to spawning grounds and (2) at spawning grounds. At locations where traps target sea lampreys en route to spawning grounds, SMW-baited traps captured significantly more sea lampreys than paired 3kPZS-baited traps (~10 % increase). At spawning grounds, no difference in trap catch was observed between 3kPZS and SMW-baited traps. The lack of an observed difference at spawning grounds may be attributed to increased pheromone competition and possible involvement of other sensory modalities to locate mates. Because fishes often rely on multiple and sometimes redundant sensory modalities for critical life history events, the addition of sex pheromones to traditionally used traps is not likely to work in all circumstances. In the case of the sea lamprey, sex pheromone application may increase catch when applied to specifically designed traps deployed in streams with low adult density and limited spawning habitat.

Journal of Chemical Ecology

Plant phenolics and absorption features in vegetation reflectance spectra near 1.66 μm

Past laboratory and field studies have quantified phenolic substances in vegetative matter from reflectance measurements for understanding plant response to herbivores and insect predation. Past remote sensing studies on phenolics have evaluated crop quality and vegetation patterns caused by bedrock geology and associated variations in soil geochemistry. We examined spectra of pure phenolic compounds, common plant biochemical constituents, dry leaves, fresh leaves, and plant canopies for direct evidence of absorption features attributable to plant phenolics. Using spectral feature analysis with continuum removal, we observed that a narrow feature at 1.66 μm is persistent in spectra of manzanita, sumac, red maple, sugar maple, tea, and other species. This feature was consistent with absorption caused by aromatic C- H bonds in the chemical structure of phenolic compounds and non-hydroxylated aromatics. Because of overlapping absorption by water, the feature was weaker in fresh leaf and canopy spectra compared to dry leaf measurements. Simple linear regressions of feature depth and feature area with polyphenol concentration in tea resulted in high correlations and low errors (% phenol by dry weight) at the dry leaf ( r 2 = 0.95, RMSE = 1.0%, n = 56), fresh leaf ( r 2 = 0.79, RMSE = 2.1%, n = 56), and canopy ( r 2 = 0.78, RMSE = 1.0%, n = 13) levels of measurement. Spectra of leaves, needles, and canopies of big sagebrush and evergreens exhibited a weak absorption feature centered near 1.63 μm, short ward of the phenolic compounds, possibly consistent with terpenes. This study demonstrates that subtle variation in vegetation spectra in the shortwave infrared can directly indicate biochemical constituents and be used to quantify them. Phenolics are of lesser abundance compared to the major plant constituents but, nonetheless, have important plant functions and ecological significance. Additional research is needed to advance our understanding of the spectral influences of plant phenolics and terpenes relative to dominant leaf biochemistry (water, chlorophyll, protein/nitrogen, cellulose, and lignin).

International Journal of Applied Earth Observation

Revealing the extent of sea otter impacts on bivalve prey through multi-trophic monitoring and mechanistic models

Sea otters are apex predators that can exert considerable influence over the nearshore communities they occupy. Since facing near extinction in the early 1900s, sea otters are making a remarkable recovery in Southeast Alaska, particularly in Glacier Bay, the largest protected tidewater glacier fjord in the world. The expansion of sea otters across Glacier Bay offers both a challenge to monitoring and stewardship and an unprecedented opportunity to study the top-down effect of a novel apex predator across a diverse and productive ecosystem. Our goal was to integrate monitoring data across trophic levels, space, and time to quantify and map the predator–prey interaction between sea otters and butter clams Saxidomus gigantea , one of the dominant large bivalves in Glacier Bay and a favoured prey of sea otters. We developed a spatially-referenced mechanistic differential equation model of butter clam dynamics that combined both environmental drivers of local population growth and estimates of otter abundance from aerial survey data. We embedded this model in a Bayesian statistical framework and fit it to clam survey data from 43 intertidal and subtidal sites across Glacier Bay. Prior to substantial sea otter expansion, we found that butter clam density was structured by an environmental gradient driven by distance from glacier (represented by latitude) and a quadratic effect of current speed. Estimates of sea otter attack rate revealed spatial heterogeneity in sea otter impacts and a negative relationship with local shoreline complexity. Sea otter exploitation of productive butter clam habitat substantially reduced the abundance and altered the distribution of butter clams across Glacier Bay, with potential cascading consequences for nearshore community structure and function. Spatial variation in estimated sea otter predation processes further suggests that community context and local environmental conditions mediate the top-down influence of sea otters on a given prey. Overall, our framework provides high-resolution insights about the interaction among components of this food web and could be applied to a variety of other systems involving invasive species, epidemiology or migration.

Journal of Animal Ecology

Behavioral responses of sea lamprey to varying application rates of a synthesized pheromone in diverse trapping scenarios

Use of the first fish pheromone biopesticide, 3-keto petromyzonol sulfate (3kPZS) in sea lamprey ( Petromyzon marinus ) control requires an understanding of both how the amount 3kPZS applied to a trap relates to catch, and how that relationship varies among stream types. By conducting 3kPZS dose-response experiments over two years and across six varied trapping contexts, we conclude (1) that 3kPZS application is best standardized by how much is emitted from the trap instead of the fully mixed concentration achieved downstream, and (2) that 3kPZS is more effective in wide streams (>30 m). In wide streams, emission of 3kPZS at 50 mg hr. −1 from the trap increased capture rate by 10–15% as sea lamprey were 25–50% more likely to enter the trap after encounter. However, in narrow streams (< 15 m), 50 mg hr. −1 3kPZS generally reduced probabilities of upstream movement, trap encounter, and entrance. While 3kPZS significantly influenced upstream movement, encounter, and capture probabilities, these behaviors were also highly influenced by water temperature, stream width, sea lamprey length, and sex. This study highlights that a pheromone component in a stream environment does not ubiquitously increase trap catch in all contexts, but that where, how, and when the pheromone is applied has major impacts on whether it benefits or hinders trapping efforts.

Journal of Chemical Ecology

Using multi-date satellite imagery to monitor invasive grass species distribution in post-wildfire landscapes: An iterative, adaptable approach that employs open-source data and software

Among the most pressing concerns of land managers in post-wildfire landscapes are the establishment and spread of invasive species. Land managers need accurate maps of invasive species cover for targeted management post-disturbance that are easily transferable across space and time. In this study, we sought to develop an iterative, replicable methodology based on limited invasive species occurrence data, freely available remotely sensed data, and open source software to predict the distribution of Bromus tectorum (cheatgrass) in a post-wildfire landscape. We developed four species distribution models using eight spectral indices derived from five months of Landsat 8 Operational Land Imager (OLI) data in 2014. These months corresponded to both cheatgrass growing period and time of field data collection in the study area. The four models were improved using an iterative approach in which a threshold for cover was established, and all models had high sensitivity values when tested on an independent dataset. We also quantified the area at highest risk for invasion in future seasons given 2014 distribution, topographic covariates, and seed dispersal limitations. These models demonstrate the effectiveness of using derived multi-date spectral indices as proxies for species occurrence on the landscape, the importance of selecting thresholds for invasive species cover to evaluate ecological risk in species distribution models, and the applicability of Landsat 8 OLI and the Software for Assisted Habitat Modeling for targeted invasive species management.

International Journal of Applied Earth Observation

Dynamic population models with temporal preferential sampling to infer phenology

To study population dynamics, ecologists and wildlife biologists typically use relative abundance data, which may be subject to temporal preferential sampling. Temporal preferential sampling occurs when the times at which observations are made and the latent process of interest are conditionally dependent. To account for preferential sampling, we specify a Bayesian hierarchical abundance model that considers the dependence between observation times and the ecological process of interest. The proposed model improves relative abundance estimates during periods of infrequent observation and accounts for temporal preferential sampling in discrete time. Additionally, our model facilitates posterior inference for population growth rates and mechanistic phenometrics. We apply our model to analyze both simulated data and mosquito count data collected by the National Ecological Observatory Network. In the second case study, we characterize the population growth rate and relative abundance of several mosquito species in the Aedes genus. Supplementary materials accompanying this paper appear on-line.

Journal of Agricultural, Biological, and Environme

A ‘how-to’ guide for estimating animal diel activity using hierarchical models

Animal diel activity patterns can aid understanding of (a) how species behaviourally adapt to anthropogenic and natural disturbances, (b) mechanisms of species co-existence through temporal partitioning, and (c) community or ecosystem effects of diel activity shifts. Activity patterns often vary spatially, a feature ignored by the kernel density estimators (KDEs) currently used for estimating diel activity. Ignoring this source of heterogeneity may lead to biased estimates of uncertainty and misleading conclusions regarding the drivers of diel activity. Thus, there is a need for more flexible statistical approaches for estimating activity patterns and testing hypotheses regarding their biotic and abiotic drivers. We illustrate how trigonometric terms and cyclic cubic splines combined with hierarchical models can provide a valuable alternative to KDEs. Like KDEs, these models accommodate circular data, but they can also account for site-to-site and other sources of variability, correlation amongst repeated measures, and variable sampling effort. They can also more readily quantify and test hypotheses related to the effects of covariates on activity patterns. Through empirical case studies, we illustrate how hierarchical models can quantify changes in activity levels due to seasonality and in response to biotic and abiotic factors (e.g. anthropogenic stressors and co-occurrence). We also describe frequentist and Bayesian approaches for quantifying site-specific (conditional) and population-averaged (marginal) activity patterns. We provide guidelines and tutorials with detailed step-by-step instructions for fitting and interpreting hierarchical models applied to time-stamped data, such as those recorded by camera traps and audio recorders. We conclude that this approach offers a viable, flexible, and effective alternative to KDEs when modelling animal activity patterns.

Journal of Animal Ecology

Paired air-water annual temperature patterns reveal hydrogeological controls on stream thermal regimes at watershed to continental scales

Despite decades of research into air and stream temperature dynamics, paired air-water annual temperature signals have been underutilized to characterize watershed processes. Annual stream temperature dynamics are useful in classifying fundamental thermal regimes and can enhance process-based interpretation of stream temperature controls, including deep and shallow groundwater discharge, when paired with air signals. In this study, we investigated multi-scale variability in annual paired air-water temperature patterns using sine-wave linear regressions of multi-year daily temperature data from streams of various sizes. A total of 311 sites from two spatially intensive regional datasets (Shenandoah National Park and Olympic Experimental State Forest) and a spatially extensive national dataset spanning the contiguous United States (U.S. Geological Survey gages) were evaluated. We calculated three annual air-water thermal metrics (mean ratio, phase lag, and amplitude ratio) to deduce the influence of groundwater and other watershed processes on stream thermal regimes at multiple spatial scales. Site-specific values of the three annual air-water thermal metrics ranged from 0.69 to 5.29 (mean ratio), −9 to 40 days (phase lag), and 0.29 to 1.12 (amplitude ratio). Regional patterns in the annual thermal metrics revealed persistent yet spatially variable influences of shallow groundwater discharge and high levels of thermal variability within watersheds, indicating the importance of local hydrogeological controls on stream temperature. Furthermore, annual thermal metric patterns from the regional datasets were generally concordant with the national dataset suggesting the utility of these annual thermal metrics for analysis at multiple scales. Analysis of the national dataset showed that previously defined thermal regimes based on water temperature alone could be further refined using air-water metrics and these metrics were related to physiographic watershed characteristics such as contributing area, elevation, and slope. This research demonstrates the importance of spatial scale and heterogeneity for inferring hydrological process in streams and provides guidance for the interpretation of annual air-water temperature metrics that can be efficiently applied to the growing database of multi-year temperature records. Results from this research can aid in the prediction of future thermal habitat suitability for coldwater-adapted species at ecologically and management-relevant spatial scales with readily available data.

Continental United States

Geographic variation in the plumage coloration of willow flycatchers Empidonax traillii

The ability to identify distinct taxonomic groups of birds (species, subspecies, geographic races) can advance ecological research efforts by determining connectivity between the non-breeding and breeding grounds for migrant species, identifying the origin of migrants, and helping to refine boundaries between subspecies or geographic races. Multiple methods are available to identify taxonomic groups (e.g., morphology, genetics), and one that has played an important role for avian taxonomists over the years is plumage coloration. With the advent of electronic devices that can quickly and accurately quantify plumage coloration, the potential of using coloration as an identifier for distinct taxonomic groups, even when differences are subtle, becomes possible. In this study, we evaluated the degree to which plumage coloration differs among the four subspecies of the willow flycatcher Empidonax traillii, evaluated sources of variation, and considered the utility of plumage coloration to assign subspecies membership for individuals of unknown origin. We used a colorimeter to measure plumage coloration of 374 adult willow flycatchers from 29 locations across their breeding range in 2004 and 2005. We found strong statistical differences among the mean plumage coloration values of the four subspecies; however, while individuals tended to group around their respective subspecies' mean color value, the dispersion of individuals around such means overlapped. Mean color values for each breeding site of the three western subspecies clustered together, but the eastern subspecies' color values were dispersed among the other subspecies, rather than distinctly clustered. Additionally, sites along boundaries showed evidence of intergradation and intermediate coloration patterns. We evaluated the predictive power of colorimeter measurements on flycatchers by constructing a canonical discriminant model to predict subspecies origin of migrants passing through the southwestern U.S. Considering only western subspecies, we found that individuals can be assigned with reasonable certainty. Applying the model to migrants sampled along the Colorado River in Mexico and the U.S. suggests different migration patterns for the three western subspecies. We believe that the use of plumage coloration, as measured by electronic devices, can provide a powerful tool to look at ecological questions in a wide range of avian species.

Journal of Avian Biology

Status of White Sturgeon (Acipenser transmontanus Richardson, 1863) throughout the species range, threats to survival, and prognosis for the future

White Sturgeon, Acipenser transmontanus (WS), are distributed throughout three major river basins on the West Coast of North America: the Sacramento-San Joaquin, Columbia, and Fraser River drainages. Considered the largest North American freshwater fish, some WS use estuarine habitat and make limited marine movements between river basins. Some populations are listed by the United States or Canada as threatened or endangered (upper Columbia River above Grand Coulee Dam; Kootenai River; lower, middle and, upper Fraser River and Nechako River), while others do not warrant federal listing at this time (Sacramento-San Joaquin Rivers; Columbia River below Grand Coulee Dam; Snake River). Threats that impact WS throughout the species’ range include fishing effects and habitat alteration and degradation. Several populations suffer from recruitment limitations or collapse due to high early life mortality associated with these threats. Efforts to preserve WS populations include annual monitoring, harvest restrictions, habitat restoration, and conservation aquaculture. This paper provides a review of current knowledge on WS life history, ecology, physiology, behavior, and genetics and presents the status of WS in each drainage. Ongoing management and conservation efforts and additional research needs are identified to address present and future risks to the species.

Journal of Applied Ichthyology

A review and synthesis of recreation ecology research supporting carrying capacity and visitor use management decisionmaking

Resource and experiential impacts associated with visitation to wilderness and other similar backcountry settings have long been addressed by land managers under the context of &ldquo;carrying capacity&rdquo; decisionmaking. Determining a maximum level of allowable use, below which high-quality resource and experiential conditions would be sustained, was an early focus in the 1960s and 1970s. However, decades of recreation ecology research have shown that the severity and areal extent of visitor impact problems are influenced by an interrelated array of use-related, environmental, and managerial factors. This complexity, with similar findings from social science research, prompted scientists and managers to develop more comprehensive carrying capacity frameworks, including a new Visitor Use Management framework. These frameworks rely on a diverse array of management strategies and actions, often termed a &ldquo;management toolbox,&rdquo; for resolving visitor impact problems. This article reviews the most recent and relevant recreation ecology studies that have been applied in wildland settings to avoid or minimize resource impacts. The key findings and their management implications are highlighted to support the professional management of common trail, recreation site, and wildlife impact problems. These studies illustrate the need to select from a more diverse array of impact management strategies and actions based on an evaluation of problems to identify the most influential factors that can be manipulated.

Journal of Forestry