Duration: 10/01/2004 to 09/30/2009

Administrative Advisor(s):

NIFA Reps:

Non-Technical Summary

Statement of Issues and Justification

STATEMENT OF THE PROBLEM:

Bark beetles and pathogens interact to cause extensive losses in the forests of North America. The vast areas affected the hidden nature of root diseases, the episodic nature of bark beetle infestations and the declining emphasis on extracting timber contribute to a lack of current, regional estimates of the impacts of these pests. Yet, these losses are significant. For example, bark beetle outbreaks are occurring in the southwestern states and devastating the pinyon pine systems in New Mexico, Arizona and Colorado, a resource not typically considered at risk to bark beetles. In other areas such as Alaska, a spruce beetle outbreak has resulted in thousands of acres of dead timber. In British Columbia in 2002, mountain pine beetle infested over 4 million ha of lodgepole pine. These examples may be directly related to changing environmental conditions and can provide insight into future bark beetle outbreaks. Pitch canker provides another example of the extent of insect-pathogen problems. This introduced pest was found in California in 1986. After 14 years, there is an active infestation zone of over 23 million acres. Movement of potential host materials out of this zone is discouraged. These examples illustrate that we lack the ability to make informed management decisions with regard to these issues.

PROGRESS:

The results from research conducted by W-187 members have provided both fundamental understanding and management tools for pathogenic fungi and bark beetles. For example, members have been actively involved with the testing and development of anti-aggregation pheromones for the management of Douglas-fir beetle and mountain pine beetle in the west. Potential range expansion of mountain pine beetle in response to global climate change is being investigated, as is the role of host nutrition and nitrogen content on the outset of bark beetle outbreaks. The interaction between natural enemies and bark beetle population dynamics, the role of fungal associates in bark beetle population dynamics, and vectoring of tree pathogens are all being cooperatively pursued by many W-187 members. Also, several members are involved in the fire-fire surrogate studies that are occurring throughout the country. In addition to the published research results, the membership of W-187 has made significant contributions in the areas of scientific policy (i.e. the letter submitted to CSREES and included in the evaluation of current funding structure) and the development of science infrastructure (i.e. the training of M.S. and Ph.D. students and collaboration with underrepresented groups in science such as the Native American students being mentored by Dr. Diana Six at the University of Montana and the involvement of Dr. Stephen Cook at the University of Idaho with management of Nez Perce land in Idaho). The critical review appended to this proposal details our more recent publications dealing with this subject matter.

EMERGING ISSUES:

There are several issues that have received increased attention since the W-187 regional research project was formed. In the western states, forest fires have been large and intense during the past several summers. These fires will result in changed landscapes and impact forest management practices throughout the west. The fires and subsequent changes in management strategies will also impact the interactions between bark beetles, pathogens and host trees for years to come. Biological invasions by insects such as the Asian longhorn beetle, the emerald ash borer, and pine shoot beetle, as well as, the causal agent of sudden oak death in California are impacting regional management issues and changing some of the interactions that occur among bark beetles, pathogens and host trees. Changes in global climate conditions (both temperature and precipitation) will alter the distributions of both the tree hosts and the insect/pathogen species. Some bark beetle and pathogen species may exploit the changes and become a threat beyond their traditional geographic or host range. Important examples include increasing activity of mountain pine beetle outbreaks in high elevation 5-needle pine ecosystems and possible movement of this species into jack pine, and the presence of southern pine beetles in Arizona. All of the above issues (fire, biological invasions, and global climate change) will result in forest fragmentation and changes in local landscapes. A cooperative research effort that crosses disciplinary and regional boundaries will be needed to address these issues and others as they emerge.

JUSTIFICATION:

Regional research project W-110, and its successor W-187 have solved problems facing our forests. New scientific technologies enable us to address old problems that were previously intractable as well as emerging problems. As outlined below, regional research project W-187 should continue because of the values at stake, the intrinsic nature of the problems, the need for cooperative work, the benefits that accrue from the proposed research, the relationship to current regional and national priorities, and this project?s impact on science. An important value of the collaborative research proposed here is that the ecological mechanisms associated with interactions among bark beetles, pathogens and conifers are similar among different taxa found throughout North America. The investigators participating in this regional project present strength through diversity in research approaches, individual subject species, and by research projects conducted under different climatic regimes.

Values: The commodity value of western forests has supported the economy of many parts of western North America. However, society also values forests for watershed, wilderness, recreation, and habitat for threatened and endangered species of fish and wildlife. These ecological and aesthetic values can exceed those of forest products such as timber, and their consideration has caused the removal of many forests from the timber-producing base. Traditional methods of pest management often rely on silvicultural options ? most of which involve some form of timber harvest or regeneration techniques to modify the environment. When forest lands are reserved, those silvicultural options are restricted or severely limited. Yet these valuable lands are what must be managed to prevent ecological imbalances that can cause devastating outbreaks of pests and loss of the resource, as well as the quality of life for the largely rural communities dependant on the forest. As forests are removed from the timber-producing base, the value of the remaining timber-producing forests will increase. Pest management in these forests will thus become increasingly important. Also, while addressing the changing landscape dynamics, we need to more fully understand the role insects and pathogens play as disturbance agents in healthy, functioning ecosystems, and the effect of management on these processes.

Bark beetles and pathogens are among the most important agents that destroy forests and result in higher economic cost in U.S. forests than any other disturbance. Recently, in the western states and Alaska severe outbreaks of spruce beetle, mountain pine beetle and Ips beetles have occurred while the southern pine beetle is at outbreak levels in the southeastern United States. In addition, root diseases are responsible for approximately 18% of the total tree mortality in the western United States, causing losses of up to 1.5 billion dollars annually (includes stumpage worth). These losses are greatly underestimated since they include only losses due to tree mortality, while additional losses caused by growth reduction are probably much greater, but are very difficult to measure. Perhaps more importantly, these losses are only in timber value, watershed, wilderness, recreation, and habitat values are not considered.

Importance and Extent of the Problem: Tree losses due to pests are extensive. Reducing these losses can provide a more stable supply of timber and additional social benefits. Reducing these losses will also benefit interregional issues such as management of spotted owl and salmon habitats, level of wildland fuels, and the increase in value of some hardwood species such as aspen in the Great Lakes region. Native insect and disease outbreaks can result in large economic losses and may also lead to additional losses such as the Yellowstone fire, and more recently fires in Southern California, that burned forests that had been heavily infested by bark beetles, dwarf mistletoes, and root diseases. Unfortunately, much of our understanding of insects and diseases comes from studies of pests in outbreak status. We must know much more about the ecological interrelationships among insects, diseases, symbiotic organisms, and trees in non-epidemic situations. These interactions are also now being impacted by introduced species such as pitch canker and the Asian longhorned beetle. As a group, W-187 is focused on the essential components of complex tree-beetle-pathogen interactions so that we can better understand forest and ecosystem level processes.

Need for Cooperative Research: The task facing this group is large and too complex for any one researcher or discipline. Multiple disciplinary cooperative research is needed to assemble the expertise required to attack this problem. Entomologists, pathologists, systematists, and physiologists working under the aegis of W-187 have already made significant progress toward understanding the role of bark beetle-carried fungi in triggering host response and tree decline, the impact of climate on beetle-tree interactions, the role of natural enemies within the systems, and the relationship of forest structure to insect outbreaks. These bark beetle-microbial-tree interactions (including nutrition, dispersion, pheromone signaling and host defense) are complex and thus require a multi-disciplinary approach as described in this proposal. These interactive affects serve as an operational framework for our collaborative research. Work on one process or interaction, without consideration of other processes, has applicability only to that population at that time. For example, studies of bark beetle brood success under various conditions provide little useful information unless they consider the taxonomy and biology of beetle fungal associates. By fitting our work within the context of this conceptual framework, and sharing our techniques, research results can be extrapolated to fill gaps in our current knowledge. Similarly, cooperative research will also help workers overcome the taxonomic barriers that often prevent them from correctly identifying subtle differences among fungal taxa. By fitting our research into this conceptual framework, we eliminate fragmented research and wasteful duplication and broaden the expertise and methods available to solve the problem. Furthermore, this cooperative research identifies new opportunities for sharing new techniques, conducting large-scale applications, and integrating comparisons.

Benefits: W-187 has benefited forestry throughout the country in many ways. The results from research conducted by W-187 members have provided both fundamental understanding and management tools for pathogenic fungi and bark beetles. These biotic interactions have been studied within the context of the abiotic environment that facilitates extrapolation of the conclusions to different systems in different regions. The ability to extend both the fundamental and applied information across different systems is one of the key benefits of a regional research program of this breadth. Specifically, resource managers already use knowledge provided by W-187 scientists regarding strain variation in the management of root diseases and the differences in host colonization and geographic distinctions in the biology of broadly distributed species. W-187 scientists have also begun to unravel the complex interaction between bark beetles, their semiochemicals, fungal associates, and natural enemies. This understanding will continue to help resource managers to minimize pest populations, especially in areas where management options are complicated by emerging socio-economic pressures.

Related, Current and Previous Work

RELATED CURRENT AND PREVIOUS WORK:

Most of the past work in North America on interactions among bark beetles, pathogens, and conifer hosts has been done by current and past members of W-187. This work was extensively reviewed in a 1993 book entitled Beetle-Pathogen Interactions in Conifer Forests (Schowalter and Filip 1993). This book remains the key reference in the area. A new book covering these topics with an emphasis on emerging issues and methodologies is being proposed by W-187 scientists led by Diana Six (University of Montana) and Steven Seybold (USDA-Forest Service). In addition, the critical review appended to this proposal discusses our more recent publications and presentations dealing with this subject matter.

The interaction between entomologists and pathologists continues to provide a unique perspective and approach for solving problems and synthesizing progress. W-187 members Paine, Raffa, and Harrington, in their Annual Review of Entomology paper Interactions among scolytid bark beetles, their associated fungi, and host conifers (1997) cite three critical areas in bark beetle-fungal-host tree relationships where better understanding is needed:

" Characterization of the multiplicity of potential interactions among organisms
" Description of the dynamic rate of interactions at the biochemical level, and
" Examination of a broader taxonomic range of associated microorganisms.

An additional need related to the research area is the voice provided by such regional research groups when it comes to policy, science and infrastructure for conducting the research (Logan & Bentz 1999, Storer et al. 2001; Cook 2002).. W-187 members are addressing all of these critical areas (Wood 2000). W-187 brings together the scientists working in this area, provides a framework for fitting their research results into a larger conceptual model, and facilitates the collaboration necessary to understand the complex problem of bark beetle-pathogen-tree interactions (Sullivan et al. 2003, Wiker et al. 2003)..

To further the audience and interaction among scientists, we held a joint meeting with scientists contributing to the regional research project W-189 (Biorational Methods for Insect Pest Management: Bioorganic and Molecular Approaches) was held in Taos, NM during 2002. We have also initiated planning with regional research project NC-193 (Insects and diseases of woody plants) for a joint meeting in 2004 or 2005.

Objectives

1. Characterize the role of biotic and abiotic factors in predisposing trees to bark beetle attack and subsequent mortality.
   
2. Characterize the diversity and interactions among tree hosts, bark beetles, their natural enemies and associated fungi.
   
3. Integrate and apply the knowledge gained from objectives 1 and 2 to forest ecosystems as influenced by emerging issues such as invasive species, global climate change, changing land use patterns and multiple and conflicting societal demands.
   

Methods

Objective 1: Characterize the role of biotic and abiotic factors in predisposing trees to bark beetle attack and subsequent mortality. 1.1. Effects of the abiotic environment on bark beetles. Studies are proposed to determine the effects of abiotic conditions, particularly temperature and host nutrient status, on bark beetle flight period activity, host colonization, and developmental success. We also propose to investigate effects of shifting climatic regimes (e.g. warming temperatures) on bark beetle population dynamics, host selection behavior, and shifts in community interactions within single trees being colonized by a variety of insect species. This information well be used in beetle models linked to climate change models for predicting risk to increased bark beetle population outbreaks given global warming. One of the studies under the national Fire-Fire Surrogate program is located in the Blodgett Research Station in California. Four treatments are: (1) no treatment, (2) thin and masticate only, (3) burn only, and (4) thin and burn. These treatments were conducted in 2002. We will continue to monitor landing rates of beetles on sticky traps and in flight intercept traps in randomly located throughout the treated areas. Analyses are currently underway to determine the effect of these treatments on subcortical insect populations, disease incidence, and subsequent tree mortality. 1.2. Effects of the abiotic environment on host trees. Abiotic factors such as water availability, temperature, fire, and soil characteristics alter tree susceptibility to bark beetle colonization and fungal infection. We propose to study the effects of abiotic factors on biochemical and physiological traits, such as host carbon allocation patterns, nitrogen assimilation across trophic levels, phenolics, and resin flow and composition, that influence host susceptibility to bark beetles and associated microorganisms. 1.3. Effects of the abiotic environment on fungal pathogens. Environmental parameters significantly affect the population biology of fungal pathogens. For example, long distance transport and local dispersal of fungal spores depend upon synoptic weather patterns across continental scales. Models derived from historic climatic records will be used to predict incidence of disease severity. 1.4. Effects of the biotic environment on host trees. We propose to study the effects of biotic factors such as other insects, mistletoes, rusts, and root diseases on host susceptibility to bark beetle attack and fungal infection. We will approach this in two ways: correlative studies will analyze the association between the occurrence of bark beetles and the incidence of diseases; mechanistic studies will address the local and systemic effects of fungal pathogens on host defense mechanisms and resistance responses against bark beetles and their microbial associates. We also propose to characterize the change in symptoms and signs of the Sudden Oak Death pathogen infecting coast live oak and tanoak over a 5 year period. We are also investigating the host selection behavior of ambrosia beetles that have been shown to be attracted to cankers produced by this pathogen. We are collecting volatile compounds on chromatographic substrates for identification using GC-MS. We also will determine the sequence of occurrence of decay fungi that are associated with the ambrosia beetle tunnels in the sapwood that may play a role in the catastrophic failure of infected trees while the foliage is still green. We also propose to determine the long term impacts of infection of Monterey pines by the Pitch Canker pathogen in native, old growth forests and a forest regenerated by fire in 1987 on the central coast of California. Also we will conduct experiments aimed at increasing Monterey pine regeneration in old growth forest without the use of fire. We have conducted 3 treatments: (1) removed all vegetation, (2) mulched the vegetation and left it on the site, and (3) left all vegetation intact. We have girdled large old growth Monterey pine to simulate bark beetle killed trees and will record a subsequent regeneration occurring in these gaps. The goal of these studies is to expose seedlings to the pitch canker pathogen so that survivors may exhibit a higher level of resistance to this pathogen. We are investigating naturally occurring infections in native Douglas-fir stands in central cost of California. We have discovered a new insect association with this pathogen (Lepidoptera: Yponomeutidae). Prior to this discovery, this moth was known only from British Columbia. The first record of pitch canker infection in the mixed conifer forests of the Sierra Nevada was found on Douglas-fir in 2003. Institutions with W187 participants who will be working on Objective 1 include: The University of Wisconsin, Madison; USDA Forest Service, Rocky Mountain Research Station; USDA Forest Service, Southern Research Station; The Ohio State University; University of California, Berkeley; University of California, Davis; University of Montana; University of Idaho; Colorado State University; Oregon State University. See the appendix for a list of participants by institution. Objective 2: Characterize the diversity and interactions among tree hosts, bark beetles, their natural enemies and associated fungi. Extending the characterization and understanding of the complexity of interactions among hosts, bark beetles, natural enemies, and their associated fungi is paramount to accumulating the relevant knowledge that will foster synthesis of ecological concepts and facilitate the development of management approaches. To accomplish this task it is of utmost importance to carefully examine from two-way to multiple interactions simultaneously, thus providing mechanisms for understanding the functionality of these systems at multiple levels. The close examination of the richness of these interactions will prove to be challenging yet crucial as we continue to expand our ecological and practical knowledge base. The methods employed will necessarily vary with specific projects, and with specific bark beetle species. However, a number of broadly applicable tools are available for each of the three sub-objectives, and these are described below. These tools offer significant opportunities because they both provide stronger experimental control and are more amenable to interregional comparisons and collaborations, than past approaches. 2.1. Contributing factors leading to the mortality of bark beetle colonized trees 1. Physical roles of bark beetles contributing to host tree mortality. 2. Roles of symbiotic microbes contributing to host tree mortality. a. Direct pathogenicity of microbial associates to the host. b. Alteration of the host defense response due to microbial associates. c. Detoxification of host defensive chemicals. Several approaches are available for quantifying host defenses against bark beetle - microbial associations. It is generally accepted that no single measure can fully characterize resistance, as conifer defenses operate in an integrated fashion. The methods employed are readily applied to all systems studied by our group, regardless of tree or beetle species. Published methods include measurement of constitutive resin flow, challenge inoculation with symbiotic fungi, chemical analysis of constitutive and induced chemistry, and beetle behavioral assays. Resin flow is measured by wounding trees to the cortical tissue and measuring the volume over a defined period. Following challenge inoculation, trees undergo histological changes including necrotic lesion formation, and these can be analyzed by a number of measurements and microscopic examinations. Methods for analyzing tree monoterpenes and phenolics are well developed, and methods for analyzing diterpene acids are being improved. These chemical analyses are conducted primarily by glc and hplc, with the methods published by members of W187 and others. Beetle host entry behavior can be assessed by caging beetles directly onto trees, or exposing them to treated natural or synthetic extracts applied to filter paper, ground phloem agar, etc. Arrival behavior can be assessed using flight traps baited with various host compounds. With these assays in place, we can measure responses to various biotic and abiotic stressors. Published and ongoing approaches for measuring responses to such variables include a) correlative analyses with drought, incidence of root pathogens, and defoliation; b) experimental manipulations of water availability, root infection, and foliage removal; c) induction of induced responses by elicitors such as jasmonic acid. 2.2 Symbiotic microbes 1. Transport mechanisms and capacity to disseminate symbiotic microbes. 2. Characterization of diversity and function of microbial associates. 3. Evolution of bark beetle associated microbes. 4. Effects of microbe-microbe, microbe-beetle, and beetle-beetle competition on the dynamics of microbe-beetle associations. 5. Effects of microbes on bark beetle nutrition. a. Prevalence, effects, and importance of mycophagy. b. Alteration of the nutritional quality of host substrate. 6. Interactions of exotic plant pathogens with native bark beetles. 7. Interactions of exotic bark beetles with native plant pathogens. As in the case of conifer defenses, a variety of published methods are available for evaluating beetle - microbial interactions. These involve a blend of traditional culturing and emerging molecular methods. The major changes since earlier versions of W187 are a) great strides have been made in clarifying the systematics of ophiostomatoid fungi. These contributions have greatly improved our capabilities; b) members of W187 have been closely involved in emerging conceptual development of symbiosis at large, and have infused these emerging concepts into models of beetle - fungal - host interactions; c) members of W187 have developed novel methods for statistically analyzing degrees of association between symbionts, e.g., region, host species, beetle population phase, etc., that then lend themselves to hypothesis based tests. The major methods to be employed in these studies are microscopic examinations of beetles and tissues, isolations of fungi from host tissue, beetles, and phoretic mites, chemical analysis of host tissue in response to fungal associates of bark beetles as well as to root fungi that may either elicit defenses to or compromise resistance against bark beetles, evaluation of beetle bacterial gut symbionts by 16S, performance assays of beetles in host tissue in the presence or absence of various microbial symbionts, correlative analyses of associations with population phase or host type, and cladistic phylogenies of beetle - fungal associations. Specifically, we will compare the spore load on four bark beetle species that have been shown to carry the pitch canker pathogen in nature: Ips paraconfusus, I. mexicanus, Pityophthorus setosus, P. carmeli. The goal of this study is to determine the vector efficiency of a few of the many bark beetle species that carry propagules of the pitch canker pathogen in nature. We will artificially contaminate males (the host selecting sex) of these species with the naturally occurring spore load and 10 times higher than the natural spore load. These beetles will be caged on branches on living trees in the native forest. 2.3 Natural enemies 1. Characterization of the diversity and impact of natural enemies. a. Predators. b. Parasitoids. c. Pathogens. 2. Host location by bark beetle natural enemies. a. Chemical ecology of bark beetle - natural enemy interactions. i. Responses to kairomones. ii. Responses to microbial volatiles. b. Visual cues. c. Auditory cues. 3. Amplification and interference among and within natural enemy species. Bark beetles are particularly well suited for studies of predator  prey interactions because of three properties: a) their pheromones are attractive to both beetles and predators, and these pheromones have been synthesized and are commercially available; b) beetles complete their entire life history within host trees, and so complete life tables can be recovered by removing phloem and recording entrance sites, ovipositional galleries, egg niches, larval mines associated with different instars, pupal chambers, and exit holes; c) at the landscape level, bark beetle activities are conspicuous because of their tree-killing nature, and both bark beetles and predators have been shown to be highly amenable to mark-recapture studies. The methods to be employed in these studies will include behavioral studies conducted in the field and in laboratory olfactometers. Field studies are typically done by trapping insects arriving to various treatments. A wide variety of olfactometer studies have been employed in the laboratory, and these methodologies are published. Other methods to be included will be impact studies using manipulative methods such as excluding predators in the field and introducing defined numbers in the laboratory. Population analyses will be conducted to provide correlations between abundances of bark beetles and various natural enemies. Our ability to interpret such analyses has increased due to improved behavioral insights arising from studies conducted by W187 members. Additional methods will include direct observation, mark-recapture, and fungal manipulations. A major change in current over previous proposals is the emphasis on complex interactions, with most studies involving components such as weather, multiple natural enemies, landscape features, host plant effects, and interactions between predators and competitors Approaches to statistical analysis have changed dramatically since the inception of W187 and its predecessor W110. The major changes reflected in experiments conducted during the current proposal will be a) analysis of long-term data sets by time series analysis; b) stand-level replication for regional analyses and inter-regional comparisons. c) application of new spatial statistical and GIS methods; d) more complete characterization of sources of variation at multiple levels of scale, and e) improved methods for contending with the high levels of multicollinearity inherent in many natural systems. Institutions with W187 members who will be working on Objective 2 include: The University of Wisconsin, Madison; USDA Forest Service, Rocky Mountain Research Station; USDA Forest Service, Southern Research Station; The Ohio State University; University of California, Berkeley; University of California, Davis; University of Montana; University of Idaho; Iowa State University. Objective 3: Integrate and apply the knowledge gained from objectives 1 and 2 to forest ecosystems as influenced by emerging issues such as invasive species, global climate change, changing land use patterns and multiple and conflicting societal demands. The development of basic knowledge on the interaction among trees, bark beetles, natural enemies, associated fungi and pathogenic organisms is a meritorious endeavor by itself. Yet, when we integrate our increased basic knowledge and contribute to the development of practical approaches for land-managers, science reaches an important purpose; to provide services to our society based upon our knowledge. For many years, fiber production was the primary force behind the management of forest ecosystems. That perspective has been replaced by the desire and need to provide and facilitate multiple uses such as watershed, recreation, and the protection of ecosystems among others. This also comes at a time when biologists are facing new challenges such as the expression of global climate change which is influencing the dynamics of insects and hosts, the introduction of exotic invasive species which is second only to habitat destruction as a threat to the sustainability of ecosystems and the recent wave of catastrophic forest fires which are perhaps indicative of ecosystems operating outside of their natural range of variation. These are some of the emerging issues which will influence where biologists focus studies and the final utilization of research products. With the above-mentioned issues as a backdrop, we have established a set of common criteria and goals to use across the participatory groups of scientists for transferring our findings to appropriate extension and forest management personnel. In general, we will: 1. Cooperate with land managers and forest health extension specialists in both development and testing of techniques and concepts for understanding the initiation and expansion of bark beetle infestations. 2. Aid extension specialists and forest managers with the development and use of Geographic Information Systems (GIS), remote sensing and spatial and temporal modeling for landscape level monitoring and management of various bark beetle species and predisposing events that may make a forest stand more susceptible to infestations of bark beetles. 3. Contribute to the development and use of genetic markers (and other molecular techniques) for the monitoring and management of bark beetles, associated fungi and relevant tree pathogens. 4. Continue developing the use of tree and beetle-produced compounds for use in monitoring and management of bark beetles and their associates through further understanding and transfer of the chemical ecology of bark beetle-tree interactions. Our ability to integrate and apply the knowledge gained from the first two research objectives will necessarily vary with individual management objectives within various agencies. However, by developing the scientific basis for understanding beetle-pathogen-tree interactions our group will be in a position to offer extension personnel and forest managers assistance in dealing with new and emerging issues such as beetle interaction with invasive species (i.e. Sudden Oak Death), global climate change, changing land use patterns and multiple and conflicting societal demands. Some examples of how the member scientists are currently contributing (or planned future contributions) to the integration of our research findings with forest management follow. Some members of the project are currently participating in the nation-wide fire, fire surrogate study. The results of these projects should aid forest managers in a number of ways including the development of knowledge for analyzing the influence of bark beetles and pathogens on fire behavior and when bark beetles and fungi should be considered in salvage following fire. Some members of the project are involved with the development of predictive risk and hazard maps/rating systems for specific bark beetles and pathogens for providing estimates of infestation development and continued tree mortality. These risk rating systems will be applied at both the stand and landscape levels. New technology such as the use of remotely sensed data are being incorporated with more traditional methods to refine our ability to detect and delineate new infestations as they occur. Global climate change and invasive species are two of the greatest threats to our native forest ecosystems. Some of our project members are directly involved with examining the potential influence of climate change on the behavior and interactions of bark beetles, their associated fungi and tree pathogens. These investigations include studies related to the persistence and range expansion of bark beetles as influenced by climate variables. Such investigations should yield information on the long-term consequences of management decisions (including the option of doing nothing) on bark beetle, their associated fungi and pathogen populations. Invasive, exotic species are becoming more and more of a threat to our forest systems and extension personnel and forest managers are looking for new methods for addressing these new threats. Some members are involved with direct investigations of exotic bark beetle species (i.e. the banded elm bark beetle), or are investigating how bark beetles may be interacting with these new organisms within our forests. For example, we propose to compare the spore load on four bark beetle species that have been shown to carry the pitch canker pathogen in nature: Ips paraconfusus, I. mexicanus, Pityophthorus setosus, P. carmeli. The goal of this study is to determine the vector efficiency of a few of the many bark beetle species that carry propagules of the pitch canker pathogen in nature. Questions that will be addressed by project members with regard to these biological invaders include: how do our management practices exacerbate exotic invasive species as well as native invasive populations and how will exotic and native invasive species affect existing communities of bark beetles, fungi and tree pathogens and their hosts? Some of the project members are continuing the development and use of semiochemical attractants and repellents for use in managing various bark beetle species. These projects can directly benefit forest managers by providing tools for use in managing beetle populations as well as providing potential information for incorporation into tree breeding programs. The investigations should also yield information on how genetic information can be used to inform land management decisions. In addition, such investigations also increase our basic knowledge for using genetics for monitoring population movement and change through time. Technology transfer is always an issue when scientists attempt to apply and incorporate the results of their research into practical management objectives. Several outlets for dissemination of information are available to project members. Many of the project members are active participants in the Western Forest Insect Work Conference and/or the Western International Forest Disease Work Conference. In addition, members plan to be involved with various demonstration projects to demonstrate the consequences of alternative management strategies on bark beetle and fungal pathogens (i.e. the fire, fire surrogate study). Some of these programs will begin to address such questions as what are the potential effects of forest fragmentation and restoration on bark beetles, fungi and tree pathogens and what are the effects of anthropogenic invasions into the urban-wildland interface and the need for accepted short term suppression techniques of bark beetle and tree pathogen populations? All W187 members will contribute to Objective 3 through the integration and application of knowledge gained through research for sustainable management of forest ecosystems.

Measurement of Progress and Results

Outputs

• Fundamental understanding and management tools for pathogenic fungi and bark beetles. For example, increased understanding of the interaction between natural enemies and bark beetle population dynamics, the role of fungal associates in bark beetle population dynamics, and vectoring of tree pathogens, especially exotic invasives.
• Significant contributions to scientific policy (i.e. comments to CSREES including an evaluation of current funding structure for forest entomology research).
• Development of science infrastructure (i.e. the training of M.S. and Ph.D. students and collaboration with underrepresented groups in science such as the Native American students being mentored by Dr. Diana Six at the University of Montana and the involvement of Dr. Stephen Cook at the University of Idaho with management of Nez Perce land in Idaho).

Outcomes or Projected Impacts

• The outcome of this information will be an increased knowledge base upon which to base forest management decisions. Specifically, forest managers will have improved data on which to predict outbreaks, based on a) weather, b) root insects and pathogens, and c) defoliators.
• This objective will help bridge the gap between new research findings and implementation, which is typically conducted by state and federal agencies, and by private timber companies. The members of this working group have a longstanding record of effective and ongoing outreach.
• There is currently a dearth of young forest entomologists and pathologists, owing to a period of attrition during cutbacks. Ironically, this coincides with a sudden increase in need, owing largely to the rapid influx of invasive forest pests.

Milestones

(2004): Initiation of this project is not contingent on any time-sensitive milestones. The requisite methods have been developed during the previous term of this project. Likewise, the collaborative linkages are formed and in full operation.

Projected Participation

View Appendix E: Participation

Outreach Plan

Our ability to incorporate and apply the knowledge gained from the collaborative work that is proposed will necessarily vary with individual management objectives within various agencies. Several outlets for dissemination of information are available to project members. The project members have an established record of publication of results in refereed journals and the proceedings from professional conferences. We will continue publishing in these outlets. In addition, many of the project members are active participants in professional societies such as the Entomological Society of America, the Society of American Foresters, the Western Forest Insect Work Conference and the Western International Forest Disease Work Conference. Members will present results of their research at meetings of these groups. Several members are actively involved with consulting with various land management agencies including university experimental forests (i.e. the University of Idaho Experimental Forest) and tribal lands (i.e. with the Nez Perce). Members are also active participants in various demonstration projects (i.e. the fire, fire surrogate study and SAF field days) that are often attended by practicing land managers. Less formal (but no less important) cooperation with land managers (representing both governmental and private forests) and state forest health extension specialists in implementing the results of the proposed research will also be pursued

Organization/Governance

The governance for the project closely follows the recommended guidelines. Officers will serve a two year term, with a new secretary elected at the annual meeting of the project and the current secretary serving as the chair during the second year. Administrative guidance will be provided by an assigned Administrative Advisor and a CSREES Representative.

Literature Cited

Aukema, B. H., D.L. Dahlsten, & K. F. Raffa. 2000. Exploiting behavioral disparities among predators and prey to selectively target pests: Maximizing bark beetle: predator removal during semiochemically based trap-out. Environmental Entomology 29: 651-660.

Aukema, B.H., D.L. Dahlsten & K.F. Raffa. 2000. Improved population monitoring by incorporating behavioral disparities among predators and prey: relating bark beetle and predator arrival rates to various synthetic lures vs. infested host. Environmental Entomology 29: 618-629.

Aukema, B.H. & K.F. Raffa. 2002. Relative effects of exophytic predation, endophytic predation and intraspecific competition on a subcortical herbivore: Consequences to the reproduction of Ips pini and Thanasimus dubius. Oecologia. 133: 483-491.

Aukema, B.H. & K. F. Raffa. 2004. Gender- and sequence- dependent predation within group colonizers of defended plants: a constraint on cheating among bark beetles? Oecologia. 138: 253-258.

Ayres, M.P., R.T. Wilkens, J.J. Ruel, M.J. Lombardero & E.J. Vallery. 2000. Nitrogen budgets of phloem-feeding bark beetles with and without symbiotic fungi. Ecology 81:2198-2210.

Bentz, B.J. & D.E. Mullins. 1999. Ecology of mountain pine beetle cold hardening in the Intermountain West. Environmental Entomology 28: 577-587.
Biesinger, Z., J. Powell, B.J. Bentz & J.A. Logan. 2000. Direct and indirect parameterization of a localized model for the mountain pine beetle - lodgepole pine system. Ecological Modeling 129:273-296.

Bishop, R. J. & Jacobi, W.R. 2003. Insects associated with black stain root disease centers in pinyon pine stands. Southwestern Entomologist 78: 55-61.
Bonello, P., W.R. McNee, A.J. Storer, D.L. Wood & T.R. Gordon. 2000. Role of olfactory stimuli in host location by twig beetles (Coleoptera: Scolytidae). Ecological Entomology 26:8-15.

Bonello P, A.J. Storer, T.R. Gordon & D.L. Wood. 2003. Systemic effects of Heterobasidion annosum on ferulic acid glucoside and lignin of presymptomatic ponderosa pine phloem, and potential effects on bark-beetle-associated fungi. J. Chem. Ecol. 29:1167-1182

Blodgett, J.T., and P. Bonello. 2003. The aggressiveness of Sphaeropsis sapinea on Austrian pine varies with isolate group and site of infection. Forest Pathology 33:15-19.

Coetzee, M.P.A., B.D. Wingfield, T.C. Harrington, D. Dalevi, T.A. Coutinho & M.J. Wingfield. 2000. Geographic diversity of Armillaria mellea s. s. based on phylogenetic analysis. Mycologia 92:105-113.

Coetzee, M.P.A., B.D. Wingfield, T.C. Harrington, J. Steimel, T.A. Coutinho, & M.J. Wingfield. 2001. The root rot fungus Armillaria mellea introduced into South Africa by early Dutch settlers. Molecular Ecology 10:387-396.

Cook, S.P. 2002. Beetle (Coleoptera) diversity in mixed pine  hardwood stands in the Ouachita highlands five years following tree harvests. Southwest. Entomol. 27: 269-276.

Cronin, J.T., J.D. Reeve, R.T. Wilkens & P. Turchin. 2000. The pattern and range of movement of a checkered beetle predator relative to its bark beetle prey. Oikos 90:127-138.

Dahlsten, D.L., K.F. Raffa, D.L. Six, B.H. Aukema & D.L. Rowney. 1999. Application of chemical ecology to conservation and augmentation of bark beetle natural enemies, 1997 results for northern California and Wisconsin. In: Hayes, J. & K. Raffa (eds.), Proceedings Bark Beetle Genetics Workshop. USDA FS PNW Gen. Tech. Rept. 466.

Dodds, K.J., and D.W. Ross. 2002. Relative and seasonal abundance of woodborers (Buprestidae, Cerambycidae) and Cucujidae trapped in Douglas-fir beetle pheromone-baited traps in northern Idaho. Pan-Pacific Entomol. 78:120-131.

Dodds, K.J., and D.W. Ross. 2002. Sampling range and range of attraction of Dendroctonus pseudotsugae pheromone-baited traps. Can. Entomol. 134:343-355.
Erbilgin, N. & K.F. Raffa. 2000. Effects of host tree species on attractiveness of tunneling pine engravers, Ips pini (Coleoptera: Scolytidae), to conspecifics and insect predators. Journal of Chemical Ecology 26: 823-840.

Erbilgin N. & Raffa K.F. 2001. Modulation of predator attraction to pheromones of two prey species by stereochemistry of plant volatiles. Oecologia. 127: 444 - 453.

Erbilgin, N. & K.F. Raffa. 2002. Association of declining red pine stands with reduced populations of bark beetle predators, seasonal increases in root colonizing insects, and incidence of root pathogens. For. Ecol. & Manag. 164: 221-236.

Erbilgin, N., E.V. Nordheim, B.H. Aukema, & K.F. Raffa. 2002. Population dynamics of Ips pini and Ips grandicollis in red pine plantations in Wisconsin: Within- and between- year associations with predators, competitors, and habitat quality. Environ. Entomol. 31: 1043-1051.

Erbilgin, N. & K. F. Raffa. 2003. Spatial analysis of forest gaps resulting from bark beetle colonization of red pines experiencing belowground herbivory and infection. For. Ecol. & Manag. 177: 145-153.

Erbilgin, N., J. S. Powell, & K. F. Raffa. 2003. Effect of varying monoterpene concentrations on the response of Ips pini (Coleoptera: Scolytidae) to its aggregation pheromone: Implications to the pest management and ecology of bark beetles. J Ag. & For. Ent. 5: 269-274.

Garriston-Johnston, M.T., J.A. Moore, S.P. Cook, & G.J. Niehoff. 2003. Douglas-fir beetle infestations are associated with certain rock and stand types
in the Inland Northwestern United States. Environ. Entomol. 32: 1354-1363.

Gordon, T.R., A.J. Storer & D.L. Wood. 2001. The Pitch Canker Epidemic in California. Plant Disease 85:1128-1139.

Haberkern., K., Raffa, K.F. & B. Illman. 2000. Ophiostomoid fungi associated with bark beetle species colonizing white spruce in the Great Lakes region. Pgs. 109  110. In: Goheen, E. M. Proc. 1999 Western Internat. Forest Disease Work Conf. And Forest Insect Work Conf. Breckenridge, CO.

Hansen, E.M., B.J. Bentz, and D.L. Turner. 2001. Physiological basis for flexible voltinism in the spruce beetle (Coleoptera: Scolytidae). The Canadian Entomologist 133:805-817.

Harrington, T.C., D.M. McNew, J. Steimel, D. Hofstra, & R. Farrell. 2001. Phylogeny and taxonomy of the Ophiostoma piceae complex and the Dutch elm disease fungi. Mycologia 93:110-135.

Harrington, T. C., N.V. Pashenova, D.L. McNew, J. Steimel, and M.Y. Konstantinov. 2002. Species delimitation and host specialization of Ceratocystis laricicola and C. polonica to larch and spruce. Plant Dis. 86:418-422.

Harrington, T.C. Ecology and evolution of mycophagous bark beetles and their fungal partners. In: Ecological and Evolutionary Advances in Insect-Fungal Associations, F. E. Vega and M. Blackwell, eds. Oxford University Press. In press.

Klepzig, K.D., J. Paschke, W. Nettleton & R. Anderson. 2001. Thinning and southern pine beetle. USDA Forest Service, Forest Health Note. 2 p.

Klepzig, K.D., J.C. Moser, F.J. Lombardero, R.W. Hofstetter & M.P. Ayres. 2001. Symbiosis and competition: Complex interactions among beetles, fungi and mites. Symbiosis. 30:83-96.

Klepzig, K.D., J.C. Moser, M.J. Lombardero, M.P. Ayres, R.W. Hofstetter & C.J. Walkinshaw. 2001.

Klepzig, K.D. & C.H. Walkinshaw. 2003. Cellular response of loblolly pine to wound inoculation with bark beetle-associated fungi and chitosan. USDA Forest Service, Southern Research Station, Research Paper SRS-30. 9 p.

Klepzig, K.D., J.Flores-Otero, R.W. Hofstetter & M.P. Ayres. 2004. Effects of available water on growth and competition of southern pine beetle associated fungi. Mycol. Res. 107:1-6.

Logan, J.A. & B.J. Bentz. 1999. Model analysis of mountain pine beetle (Coleoptera: Scolytidae) seasonality. Environmental Entomology 28: 924-934.

Logan, J.A., J.A. Powell & B.J. Bentz. 2000. Evaluating the potential for climate change induced bark beetle invasion of high elevation ecosystems. In A. Menzel (ed) Progress in Phenology, Monitoring, Data Analysis, and Global Change Impacts, Freising, Germany, Oct 4-6, 2000.

Lombardero, M.J., M.P. Ayres, B.D. Ayres & J.D. Reeve. 2000. Cold tolerance of four species of bark beetle (Coleoptera:Scolytidae) in North America. Environmental Entomology 29:421-432.

Lombardero, M.J., M.P. Ayres, P.L. Lorio & J.J. Ruel. 2000. Environmental effects on constitutive and inducible resin defences of Pinus taeda. Ecology Letters 3:329-339.

Lombardero, M.J., M.P. Ayres, R.W. Hofstetter, J.C. Moser & K.D. Klepzig. 2003. Strong indirect interactions of Tarsonemus mites (Acarina, Tarsonemidae) and Dendroctonus frontalis (Coleoptera: Scolytidae). Oikos 102:243-252.

Maijala, P., T.C. Harrington & M. Raudaskoski. 2003. A peroxidase gene family and gene trees in Heterobasidion and related genera. Mycologia 95:209-221.

Malmstrom, C.M. & K.F. Raffa. 2000. Biotic disturbance agents in the boreal forest: considerations for vegetation change models Global Change Biology. 6: 1-14.

McNee, W.R., D.L. Wood, A.J. Storer & T.R. Gordon. 2002. Incidence of the pitch canker pathogen and associated insects intact and chipped Monterey pine branches. Can. Entomol. 134: 47-58

McNee W.R., P. Bonello, A.J. Storer, D.L. Wood & T.R. Gordon. 2003. Feeding response of Ips paraconfusus to phloem and phloem metabolites of Heterobasidion annosum-inoculated ponderosa pine, Pinus ponderosa. J. Chem. Ecol. 29: 1183-1202
McPherson, B.A., D.L. Wood, A.J. Storer, P. Svirha, D.M. Rizzo, N.M. Kelly & R.B. Standiford. 2000. Oak Mortality Syndrome: Sudden Death of Oaks and Tanoaks. Tree Notes: California Department of Forestry and Fire Protection. Number 26, 6 pp.

McPherson, B.A., D.L.Wood, A.J. Storer, N.M. Kelly & R.B. Standiford. 2003. Sudden oak death, a new forest disease in Californai. Integrated Pest Manag. Rew. 6: 243-246.

Otrosina, W.J., C.H. Walkinshaw, S.J. Zarnoch, S. Sung & B.T. Sullivan. 2002. Root disease, longleaf pine mortality, and prescribed burning, pp. 551-557. In K.W. Outcalt (ed.), Proceedings of the Eleventh Biennial Southern Silvicultural Research Conference. USDA Forest Service Gen. Tech. Rep. SRS-48.

Paine, T.D., K.F. Raffa, K.F. and T.C. Harrington. 1997. Interactions among scolytid bark beetles, their associated fungi, and live host conifers. Annual Review of Entomology 42: 179-206.

Pettersson, E.M., B.T. Sullivan, P. Anderson, C.W. Berisford & G. Birgersson. 2000. Odor perception in bark beetle parasitoid Roptrocerus xylophagorum (Ratzeburg) (Hymenoptera: Pteromalidae) exposed to host-associated volatiles. Journal of Chemical Ecology 26: 2507-2525

Powell, J.A., J. L. Jenkins, J.A. Logan, and B.J. Bentz. 2000. Seasonal temperature alone can synchronize life cycles. Bulletin of Mathematical Biology 62:977-998.

Raffa, K.F., B. H. Aukema & D. L. Dahlsten. 2000. Using semiochemical disparities among bark beetles and predators to improve the monitoring and conservation of natural enemies. pp. 37  39. In: Goheen, E. M. Proc. 1999 Western Internat. Forest Disease Work Conf. And Forest Insect Work Conf. Breckenridge, CO.

Raffa, K.F. 2001. Mixed messages across multiple trophic levels: The ecology of bark beetle chemical communication systems. Chemoecology 11: 49-65.
Seybold, S.J., Bohlmann, J. & K.F. Raffa. 2000. Biosynthesis of coniferophagous bark beetle pheromones and conifer isoprenoids: an evolutionary perspective and synthesis. Can. Entomol. 132: 1-57.

Six, D.L. and B.J. Bentz. 2003. The fungi associated with the North American Spruce Beetle, Dendroctonus rufipennis. Canadian Journal of Forest Research. 33: 1815-1820.

Six, D.L., T.C. Harrington, J. Steimel, D. McNew, and T.D. Paine. 2003. Genetic relationships among Leptographium terebrantis and the mycangial fungi of three western Dendroctonus bark beetles. Mycologia 95: 781-792.

Sullivan B.T., E.M. Pettersson, K.C. Seltmann & C.W. Berisford. 2000. Attraction of the bark beetle parasitoid Roptrocerus xylophagorum (Hymenoptera: Pteromalidae) to host-associated olfactory cues. Environmental Entomology 29: 1138-1151.

Sullivan, B.T., M.J. Dalusky & C.W. Berisford. 2003. Interspecific Variation in Host-Finding Cues of Parasitoids of the Southern Pine Beetle (Coleoptera: Scolytidae). J. Entomol. Sci. 38: 631-643

Sullivan, B.T., C.J. Fettig, W.J. Otrosina, M.J. Dalusky & C.W. Berisford. 2003. Association between severity of prescribed burns and subsequent activity of conifer-infesting beetles in stands of longleaf pine. For. Ecol. & Manag. 185: 327-340

Schowalter, T.D. and G.M. Filip 1993. Beetle-Pathogen Interactions in conifer Forests. Academic Press, 252 pp.

Storer, A.J., P. Bonello, T.R. Gordon & D.L. Wood. 1999.Evidence of resistance to the pitch canker pathogen (Fusarium circinatum) in native stands of Monterey pine (Pinus radiata). Forest Scienci 45: 500-505.

Storer, A.J., D.L . Wood, T.R. Gordon & W.J. Libby. 2001. Restoring native Monterey pine forests in the presence of an exotic pathogen. Journal of Forestry 99(5):14-18.

Storer, A.J., D.L. Wood & T.R. Gordon. 2002. Epidemiology of pitch canker disease of Monterey pine in California. Forest Sci. 48: 694-700

Veysey, J.S., M.P. Ayres, M.J. Lombardero, R.W. Hofstetter & K.D. Klepzig. 2003. The effect of alternative host species on reproductive success of Dendroctonus frontalis (Coleoptera: Scolytidae). Environ Entomol. 32:668-679.

Wikler, K., A.J. Storer, W. Newman, T.R. Gordon & D.L. Wood. 2003. The dynamics of an introduced pathogen in a native Monterey pine (Pinus radiata) forest. For. Ecol. & Manag.179: 209-221

Wood, D.L. 2000. Insect/Pathogen/Tree Interactions - Where Do We Go From Here? "An Overview of Western Regional Research Project W-110 and W-187. Proc. 51st Ann. Mtg., Western Forest Insect Work Conference, pp. 33-36.

Wallin, K.F. & K.F. Raffa. 2001. Host - mediated interactions among feeding guilds: Incorporation of temporal patterns can integrate plant defense theories to predict community level processes. Ecology 82: 1387-1400.

Wallin, K.F. & K.F. Raffa. 2002. Density-mediated responses of bark beetles to host allelochemicals: A link between individual behavior and population dynamics. Ecol. Entomol. 27:484-492.

Zhou, J., D.W. Ross, and C.G. Niwa. 2001. Kairomonal response of Thanasimus undatulus (Say), Enoclerus sphegeus F. (Coleoptera: Cleridae), and Temnochila chlorodia (Mannerheim) (Coleoptera: Trogositidae) to bark beetle semiochemicals in eastern Oregon. Environ. Entomol. 30:993-998

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