Duration: 10/01/2016 to 09/30/2021

Administrative Advisor(s):

NIFA Reps:

Non-Technical Summary

Statement of Issues and Justification

Issue(s) and Need:

Eastern white pine (Pinus strobus) is a crucial ecological and economic component of forests in the eastern U.S. and Canada. In the southeastern U.S., white pine is an especially critical associate of forests in the Appalachian Mountains as hemlock trees have been in decline due to the exotic hemlock woolly adelgid (Adelges tsugae). Yet throughout the eastern U.S., from Georgia to Michigan to Maine and adjacent areas in Canada, white pines have experienced unprecedented damage in recent years due to native pests and pathogens that reduce the species’ growth, productivity, and economic value.

For many decades, white pine health has been adversely affected by the non-native white pine blister rust (Cronartium ribicola) and the native white pine weevil (Pisodes strobi).  In this proposal, we describe additional native white pine stressors that have received little or no attention, but are nonetheless linked to declining health of white pine stands throughout North America.

In the northeastern US, Caliciopsis pinea, a stem canker pathogen, poses a significant threat to white pine health. In the past 10-15 years, white pines have succumbed to an increase in the occurrence and severity of C. pinea infestations; these infestations result in excessive resin production, leading to serious growth declines and commercial defects in lumber. External symptoms include crown thinning, cankers and profuse pitching. From recent surveys conducted on 360,000 acres of white pine forest (defined as >75% white pine basal area) in Maine, Massachusetts, and New Hampshire, we know that C. pinea cankers occur on 36% of the surveyed land⁽²⁷⁾.  With white pine being such an important forest product in the northeastern U.S., it is critical to understand the extent to which C. pinea reduces wood quality and lumber yield.

White pine stands in the northeast are also facing an emerging foliar disease complex, “white pine needle damage”, that will likely remain a problem in years with excessive springtime moisture. White pine needle damage results from one or several fungal pathogens native to North America. In addition to brown spot caused by Lecanosticta acicola (=Mycosphaerella dearnessii), several other species of fungi have been isolated from diseased pine needles at a high frequency that may represent novel white pine pathogens. Unfortunately, there is still limited information on the basic etiology, epidemiology, and managment of these pathogens on eastern white pine. While mortality resulting from white pine needle damage has not been reported, continued defoliation will weaken trees, making them more susceptible to other biotic and abiotic stresses.

In the Lake States region, unusual white pine health problems were first detected in 2006 in the Au Sable and Manistee River corridors located in Michigan’s north-central Lower Peninsula. By 2012, the crown symptoms of dying branches also appeared in the Upper Peninsula region, where it was reported in the Munising District of Hiawatha National Forest. Symptomatic white pine has been primarily associated with stem and branch cankers induced by native fungal pathogens (primarily Diplodia spp.) and the native pine spittlebug (Aphrophora parallella).

In the southern Appalachians there has been an increase of dieback and mortality symptoms in white pine since 2006, especially in the mountains of Georgia, Virginia, and West Virginia. These symptoms include branch flagging, canker presence, high resin pitching associated with cankers, and mortality across a range of tree diameters. In Georgia’s Chattahoochee National Forest, one region documented 44% of the white pine trees as dead and another 30% of the trees showed >50% dieback. In both Georgia and Virginia, the majority of mortality was observed on trees with <30 cm diameter. Preliminary examination of cankers yielded primarily C. pinea, which was found on 86% of branches. Embedded within these various cankers were immature pine scale insects, which were later identified via DNA analysis as Matsucoccus macrocicatrices, the first report of the scale in the southeastern U.S.  Surprisingly, M. macrocicatrices is common in the northeastern U.S. and Canada and is known as the “Canadian pine scale,” but the insect has not been investigated as an associate of C. pinea infestations. Because the scale is frequently associated with C. pinea infections on white pine trees in the southeast, this indicates a previously undescribed relationship between insect and fungus affecting white pine health that should be further explored⁽²⁵⁾.

The above reports on declining health of white pine indicate that the stresses may vary from region to region.  However, the reported symptoms are similar, given the frequent reference to canker formation, needle loss, declining growth, dieback, and mortality.  The label “white pine health issues” will be used here to refer to the range-wide phenomenon of declining health in numerous eastern white pine stands. 

White pine has enormous economic value throughout its range.  Over the region, the net volume of white pine saw logs is over 186 billion board feet (USDA Forest Service, Forest Inventory and Analysis).  With a typical market price of $100/1,000 bd ft, the potential value of standing white pine is $18.6 billion.  To help guide the white pine and other forest product markets, market models are being developed for predicting the impacts of changes in forests and the forest products industry, regardless of whether the changes are intentional or not. Models have been used extensively in other parts of the country to help the forest products industry understand the impacts of changes in the resource supply and demand, to better prepare for potential future scenarios, and to help policy makers assess potential impacts of policies. These forward-looking models can be used to assess impacts of any factor affecting wood supply, including diseases and pests that affect tree growth and mortality.  Therefore, the market models should consider the impact of white pine health issues on white pine markets. 

From an ecological perspective, white pine has significant impact on the eastern forest.  White pine attains the largest dimensions of any eastern tree serving as a critical habitat for many species of wildlife that depend on emergent crowns and large snags and downed woody debris.  More information is needed on white pine health issues to better understand how well white pine can continue to provide these ecosystem features and identify effective management strategies to enhance white pine health and resilience in the face of these threats.

Importance and Consequences if Work is Not Accomplished

Estimating losses due to white pine health issues creates challenges because symptoms can be difficult to recognize and quantify.  Unlike mortality caused by invasive pests, which happens quickly and is easy to recognize, the white pine health issues described in this proposal develop slowly and involve multiple stresses. For example, stresses associated with white pine health issues differ throughout its range.  Losses accumulate over time and are more related to reductions in growth losses and wood quality, losses that are more difficult to quantify than mortality.  As a result, land owners and managers are concerned that loss in tree productivity and allowable cut are occurring without immediate recognition.  Therefore, actions are not being taken to improve productivity by mitigating factors associated with white pine health issues. More targeted efforts are needed to recognize and quantify these losses throughout the range of eastern white pine.  For example, the extent to which C. pinea reduces wood quality and lumber yield remains poorly understood, and potential actions for reducing C. pinea damage have not been tested.  White pine needle damage has recently gained attention, but the long-term effects of repeated defoliation remain undocumented. Further, organisms native to North America, such as needle fungi and M. macrocicatrices, are associated with unprecedented white pine health issues, and it is uncertain if the situation is developing into larger problems that will affect white pine health and productivity on a broader scale. If we fail to quantify white pine losses associated with white pine health issues, the extent and severity of these complexes will continue to be poorly understood, poorly managed, and absent from market models.

What is happening to the white pine?  Native organisms are behaving in unprecedented, unexpected manners that need to be explained.  Explanations will involve a number of factors including distribution and genetic changes in both white pine trees and biotic stress agents, as well as changes in the climate and stand development patterns of white pine forests. Taken together, these factors raise the following questions related to biotic stressors:

1. Why are cankers associated with pinea infections appearing at damaging levels in stands; is this due to increased precipitation in late spring favoring infestations, or is the stand development of white pine in abandoned fields predisposing the tree to infestations?
2. Why are the needle fungi now causing damage, and why are some trees more affected than other trees co-existing in the same stand?
3. To what extent are changes in climate associated with white pine needle damage, and to what extent is infection and subsequent needle loss the result of other stresses predisposing trees to infection?
4. Why is macrocicatrices associated with damage in the south but not the north? Is this a behavior change or has the scale been associated with damage in the north but gone undetected?
5. Is the presence of macrocicatrices and C. pinea in the same stand synergistic to the development of cankers on pine?

Without answers to these and other questions, it will not be possible to assess future losses associated with white pine health issues, and consequently, management recommendations cannot be developed to improve the health and productivity of white pine.

Although the multiple stress agents associated with white pine health issues make it a challenge to study, strategies aimed at improving white pine health are readily available.  White pine has long been successfully regenerated using silvicultural methods, such as shelterwood and seed tree cuts, and responds very well to thinning.  By gaining a better understanding of white pine health issues, we will be able to recommend treatments that reduce the risk of white pine damage due to these stressors, and we can evaluate the economic benefits of silvicultural practices that reduce white pine’s predisposition to health issues.   

Technical Feasibility

Funded projects are currently supporting surveys of white pine health issues in northeastern U.S., southern Appalachia, and Michigan. Market models for forest products are being developed for Maine and will be available to serve as a case study on how to utilize the results of this project in other market models.

Specifically, funded projects are supporting or have supported efforts to better understand (1) factors affecting C. pinea infestations in New England, (2) fungal infections and tree impacts associated with white pine needle damage, (3) interactions of pine spittlebug and stem fungi on white pine in Michigan, and (4) association of M. macrocicatrices with stem fungi and impacts on white pine in southern Appalachia.  Techniques are well established for using dendrochronology to quantify stress impacts on tree growth and onset of infestation, for sampling and identifying fungi and insects, and for measuring the physiological status of pine through stem flow measurements.  Funded projects are also helping to improve quantification of damage on white pine.

The critical role of eastern white pine in the region is well understood, timber values are high and quantifiable, and silvicultural practices are available for regenerating and growing high-value white pine.  As our understanding increases on the development of white pine health issues in the region, the results can be easily incorporated into the best practices for managing the species and reducing damage.  

Critical Role of Multistate Effort

Given the complexity and extent of white pine health issues, it is unlikely that major advances will be accomplished by any single research group working in relative isolation; such advances require collaboration among groups working in parallel. Collectively, potential project members have an enormous amount of expertise in various aspects of white pine health. However, when viewed regionally, the research effort appears fragmented. Bringing members together in a structured and active arrangement would create a synergism in research effort, thereby increasing research efficiency and productivity, with the ultimate goal of improving white pine health throughout the region.

Currently, the potential project members include researchers and outreach professionals from Land Grant Universities (Georgia, Maine, New Hampshire, Vermont), USDA Forest Service (New Hampshire, Georgia, Minnesota), and state agencies (Maine, New Hampshire, West Virginia).  Many of these participants work directly with stakeholders. The combination of researchers and outreach professionals creates an effective means of finding practical solutions for improving the management and health of white pine.  Creating the multistate project will allow project members to:

• Review, synthesize, and share information on white pine health issues.
• Identify critical knowledge gaps that currently hamper efforts to improve white pine health.
• Avoid unnecessary duplication of research effort.
• Establish a collaborative information-exchange network to provide better detection and understanding of factors affecting white pine health.
• Standardize field, laboratory, and analytical protocols across the regions.
• Develop additional multi-institutional research proposals for national-level grants in order to bolster and build upon this initial work.
• Develop silvicultural prescriptions and other management recommendations to help mitigate potential white pine threats.
• Ensure that new findings are communicated with land managers.

Impacts

Bringing together region-wide expertise on white pine health issues will allow assessment of the species’ responses to a changing environment.  The combination of land use changes and climate change may be initiating transformations in white pine’s responses to its environment that may dramatically alter the appropriate management prescriptions for the future.  The proposed working group will initiate this assessment and develop a range-wide understanding on the future health of white pine.  Finally, our goal of using this initial project to motivate an additional multi-institutional, national-level grant could lead to much broader impact than that outlined above.

Related, Current and Previous Work

Caliciopsis pinea Canker Studies in the Northeast

Caliciopsis pinea was documented on white pine as early as the 1930s ^{(6, 24, 29)}, when initial pathogen epidemiology was conducted. The fungus has been found in West Virginia, New England, and New York, as well as in Europe ^{(2, 7, 24, 29)}. In North America, C. pinea infects Pinus strobus, P. echinata, and P. virginiana ⁽²⁹⁾. In Europe, the pathogen infects P. pinaster and P. radiata ⁽⁷⁾. Other species within the genus C. pinea infect western conifers ^{(15, 16)}. The pathogen attacks thin-barked areas of the bole and branches ⁽²²⁾. Trees of all sizes and age classes are affected. External symptoms include cankers and profuse pitching. Internally, the fungus can cause serious wood defects inside infected trees.

Three funded projects are currently working on C. pinea issues in the northeastern U.S.  A project⁽²⁸⁾ led by I. Munck with the USDA Forest Service in Durham, New Hampshire, has the objectives to 1) develop species-specific primers that will amplify C. pinea DNA amongst plant/microbial DNA in the host, 2) identify areas at greatest risk of C. pinea damage, 3) assess effects of thinning in stands infected with C. pinea, and 4) develop management guidelines for reducing C. pinea canker damage. A second project by the same group ⁽²⁵⁾ has the objectives to 1) quantify the relationship between C. pinea canker severity with lumber yield and quality, and 2) provide forest managers a method for assessing C. pinea infection on standing white pine that can be used to estimate lumber yield and quality.  A third, more detailed study on how C. pinea reduces wood quality and lumber yield is centered at the University of Maine ⁽¹⁰⁾ and has the following objectives to: 1) quantify reduction in lumber yield, grade, and associated economic losses resulting from C. pinea cankers; and 2) develop management recommendations intended to reduce C. pinea damage. By linking work of the three projects, the investigators will provide the justification needed to implement silvicultural practices, such as thinning, to reduce C. pinea cankers in white pine stands.

White Pine Needle Damage

Concern over white pine needle damage has expanded to an international level because important pine growing regions including the U.S., Canada, and Europe have been affected by this problem. In 2010 these fungi were responsible for 60,200 acres of damage to white pine forests in Maine ⁽¹³⁾.  In 2012, needle damage in New Hampshire and Vermont was observed on 12,100 and 3,700 acres of white pine, respectively (Frament per. comm.). Work at the University of New Hampshire ⁽⁶⁾ linked white pine needle damage to the fungal pathogen L. acicola and occasionally to Lophophacidum dooksii (=Canavirgella bandfieldii) and Bifusella linearis.  Three other species of fungi were also recovered from diseased pine needles ⁽⁷⁾.  It is hypothesized that several consecutive years with wetter and warmer than average weather during the spring created an environment conducive to a fungal epidemic. Given many climate-change scenarios ⁽¹⁴⁾, it is likely epidemics of white pine needle damage will become more common, as these fungi are able to survive the less harsh winters and produce more spores during the longer summers. However, not enough information is known about the fungi associated with white pine needle damage, or the climatic factors that favor or inhibit epidemics of these fungi. Having this information will lead to the development of better predictive models and sound management recommendations.

While the extent of defoliation from white pine needle damage has been documented, the impact of the defoliation has not been quantified.  Current studies by members of this group⁽¹⁾ are assessing the timing and magnitude of defoliation events through litter fall analysis across Maine and New Hampshire in order to provide a defoliation metric that may be used to validate annual USFS crown surveys. Additionally, assessments are being made as to what extent defoliations are affecting 1) the hydraulic and microclimate dynamics of infected white pine stands, 2) the measurement of declining growth rates, and 3) the development of a climate based model for predicting disease outbreaks. Warmer winter temperatures and higher than average spring precipitation in the region are thought to be exacerbating white pine needle damage. Severe needle infestation can result in up to a 50% reduction in stem growth and an estimated 18% reduction in water use in white pine⁽¹⁾.  Leaf gas exchange measurements of infected needles have shown evidence of a compensation response through an upregulation of photosynthesis in the current year needles of diseased trees at the expense of lower leaf water use efficiency. These physiological declines associated with white pine needle damage likely compromise tree vigor and secondary defense mechanisms. This pathogen complex should garner greater attention from foresters and researchers alike as it has shown that chronic defoliations lead to decreased stem growth in pine stands in the short term, but perhaps tree mortality in the long term as non-structural carbohydrate reserves are depleted and compounded with other biotic and abiotic stressors.

White Pine Health Issues in the Great Lakes Region

Drought and intraspecific competition in high density pine stands have been implicated as predisposing factors for increased white pine decline in Michigan ⁽¹⁹⁾. Predisposed trees are more likely to be affected and subsequently form stem and branch cankers caused by fungal pathogens; to date Diplodia and Therrya spp. have been implicated ^{(6, 19, 31, 32)}. Entry and growth of fungal pathogens in trees is promoted by the pine spittlebug which is a sap-feeding pest of white pine in the Great Lakes. Drought likely influences the vulnerability of white pine to the pests ⁽¹¹⁾.

Matsucoccus Scale and Canker Fungi on White Pine in the Southern Appalachians

Studies at the University of Georgia ⁽¹⁷⁾ have identified a number of pathogens from branch and stem cankers of white pine including Fusarium chlamydosporium, F. acuminatum, Diplodia scrobiculata, Pestalotiopsis spp., Phomopsis spp., and C. pinea. Caliciopsis pinea was found on 86% of branches with cankers. Embedded within these various cankers were also immature pine scales (genus Matsucoccus) which were later identified as Matsucoccus macrocicatrices. The discovery of M. macrocicatrices is novel because: 1) this species has never been reported or collected from the southeastern U.S.; it is known only from Massachusetts, New Brunswick, New Hampshire, Nova Scotia, Ontario, and Quebec; 2) it has never been reported with trees that are showing signs of dieback or mortality; and 3) it has never been found associated with pathogens, and only with Septobasidium pinicola, an epiphytic, non-pathogenic and mutualistic fungus ^(4,35). Because the immature stages of M. macrocicatrices are either deeply embedded in the cankers or present on top of the bark with clear necrotic tissue under their feeding area, M. macrocicatrices may be creating wounds for infections by pathogenic fungi such as C. pinea. Relative contribution of the scale insect and fungi to canker formation are currently unknown.

Market Models and White Pine Health⁽¹¹⁾

One application of market models of the forest industry is in assessing the impacts of ecological or biological changes that lead to subsequent changes in management activities. Market models allow for the assessment of the interactions between markets and management activities, positioning the future forest as an outcome of changes in both. In the West, for example, market models have been used extensively to assess the market impacts of removals of large volumes of woody material to reduce wildfire risk. Through the incorporation of current forest supply information and projections of growth and yield in the future, along with endogenously determined prices, market models can be used to assess the market impacts of changes in white pine availability, quality, or prices and the subsequent feedback interactions from market effects to future management decisions.

Key Research Questions

Clearly, we are dealing with previously undocumented insect and fungal complexes on white pine trees in the eastern U.S., risks that cannot yet be assessed for their potential impacts on white pine markets.  At present, there are several critical unanswered questions about these phenomena such as:

1. What is the status of white pine health issues (distribution, impacts on growth and mortality, rates of intensification)?
2. Are these health issues prevalent in certain regions and localities due to changes in land use history and climate conditions?
3. Are there other explanations as to why native insects and pathogens are involved with white pine health issues, especially in natural forest stands?
4. Are 1) fungi found on damaged white pine needles in the Northeast and 2) macrocicatrices in the Southeast recent introductions or just recently discovered?
5. What are the impacts of white pine health issues on lumber quality and forest product markets?

Currently, efforts at answering these important questions remain fragmentary and disconnected. Therefore, we propose to form a USDA Multistate Research Project focusing on white pine health.  The project will involve interdisciplinary and multiagency team members working together to better understand the impacts of the changing environment and of previously undocumented insect and pathogen complexes on this ecologically and economically important tree species.

Objectives

1. Estimate losses associated with affected white pine trees.
   Comments: <br>a. Measure the incidence, range and severity of disease symptoms and fate of symptomatic white pine tree in the eastern US. <br>b. Quantify reduction in lumber yield, grade, and associated economic losses resulting from cankers. <br>c. Provide economic loss information to improve market models.
2. Obtain essential information about pests, pathogens, site conditions, and climate associated with white pine health issues in the eastern US.
   Comments: <br>a. Assess if white pine health varies according to site conditions such as forest structure and composition, various topographic features, land use history, soil conditions, and climate. <br>b. Examine the interactions between insects, fungal pathogens, cankers, and health of eastern white pine. <br>c. Evaluate and characterize the fungi associated with white pine needle damage in the northeastern United States.
3. Develop management recommendations for improving white pine health and reducing economic losses.
   Comments: <br>a. Develop silvicultural prescriptions for increasing resilience/health prior to outbreaks <br>b. Recommend mitigation treatments to infected stands to minimize losses and improve future health <br>c. Propose stand improvement options for diversification of species composition and economic value

Methods

Methods currently being used by project investigators will be summarized below.  Next, an approach will be described that will provide joint planning and coordination among the collaborators to facilitate the better results for all 3 objectives. 

Caliciopsis pinea Canker Studies in the Northeast

PCR assay: A PCR assay for C. pinea will be developed and tested with samples from subset of sites ^{(5, 28)}.

Risk assessment: Detailed forest inventory data were collected for state-owned properties in New Hampshire. Data from these initial surveys will be used to geospatially locate areas at risk of C. pinea canker damage. Variables to be considered will include stand variables such as tree species, density, and basal area as well as site-level variables such as topography and soil type⁽²⁸⁾.

Pathogen incidence and severity on symptomatic white pine will be evaluated in at least 10 sites per State across a range of risk categories. Infection severity will be estimated by counting the sections of the tree (top/mid/bottom/bole on two sides of the tree) with C. pinea canker symptoms.  Data loggers that record climatic variables such as relative humidity and temperature will be installed in sites ranging in pathogen incidence to examine if infections are associated with climate variables ⁽²⁸⁾.

Thinning effects and loss of economic product ^{(10, 27)}: Three locations, two in New Hampshire and one in southern Maine, with C. pinea infections were thinned 5-15 years ago. At each location, symptomatic stands will be located in thinned and adjacent unthinned sites.  Six trees will be harvested in each stand, three with low infection rates and three with high (canker severity > median value for region).  Stems are then cut at 1 foot intervals and examined for surface and internal cankers.  A representative sample of up to 33% of cankered discs (radial stem cross-sections) per tree will be collected from the sections and returned to the lab for dendrochronological analysis, including yearly canker incidence and changes in growth responses.  In addition, wood chips from symptomatic tissues will be collected from each stand for later DNA analysis to detect the presence of C. pinea.  Another ten trees in each stand will be harvested, cut into commercial lengths, and scaled for useable volume.  Trees will then be cut into dimensional lumber, dried, planed, and graded to calculate quantitative relationships between wood volume and quality, C. pinea infection rates, and thinning. 

Results from the C. pinea work will be combined to produce management guidelines.

White Pine Needle Damage

Diagnostic assay development⁽⁶⁾:  Sequence data from the ITS1-5.8S-ITS2 region of the rDNA will be used to develop primers specific to each species of fungi associated with white pine needle damage. A multiplex quantitative polymerase chain reaction (Q-PCR) will be employed for the identification and quantification of each of the species associated with white pine needle damage simultaneously. This diagnostic assay will also allow for the detection and monitoring of latent infections, which is the state when a host is infected with a pathogen but does not show any symptoms that have been hypothesized to be the inoculum source for the subsequent years’ epidemic

White pine needle loss, transpiration, and radial growth⁽¹⁾.  Four locations in New Hampshire and Maine are being used for litterfall measurements. Litterfall traps have been collected monthly throughout the 2014 and 2015 growing seasons (May-October).  Transpiration was monitored during the 2014 growing season on a subset of trees to continuously measure the rate of sapflow in order to quantify changes in tree water use associated with white pine needle damage defoliations.  Additionally, leaf gas exchange was measured on current year and 1 year old needles of both diseased and health trees during the summer of 2014 in order to detect changes in photosynthesis, stomatal conductance, and water use efficiency resulting from white pine needle damage.  Increment cores from eight locations throughout Maine, New Hampshire, Vermont, and Massachusetts have been analyzed for wood growth declines. Tree ring analysis will be used to quantify growth reductions from the current outbreak, attempt to detect the initiation of decline, and relate growth trends with climate and potentially past outbreaks.

Silvicultural management experiment ⁽²⁾:  During the summer of 2015 two long-term study plots were established in pine stands in Hillsborough and West Ossippee, New Hampshire in order to test the effects of silvicultural intervention on white pine needle damage incidence and severity. These plots each consist of 2 blocks, with each block consisting of three 1 acre treatment plots: a high density thinning (100 ft²/ac), a low density thinning (60 ft²/ac), and a control plot (no thinning). Pending the winter of 2015-16 these plots will be thinned to the desired levels in an effort to increase growing space and light exposure between infected crowns. During the 2015 pre-treatment year a forest inventory was conducted and all tallied white pine were evaluated for dbh, height, live crown ratios, crown diameter, foliar transparency, crown die-back, crown position, light exposure, and severity of white pine needle damage within the crowns. These inventory metrics will be repeated each subsequent year of the silvicultural thinning in order to monitor crown response in addition to high resolution dendrometer measurements of stem growth. The primary goal of this study is to determine a management strategy for mitigating white pine needle damage in New England.

Matsucoccus Scale and Canker Fungi on White Pine in the Southern Appalachians

Field studies. Thirty-three permanent plots in Georgia, Virginia, and West Virginia are currently being monitored every six months. Using guidance from FIA data, field scouting, and incoming reports of white pine health issues, 40 additional sites and 120 plots were established on federal lands in Georgia, Virginia, West Virginia, North Carolina, and Tennessee. Within each site, five white pine seedlings were collected to quantify canker and scale densities. Data collected included the number scales at each life-stage, the number and sizes of cankers, and the presence/absence of C. pinea fruiting bodies. Results of the recent studies and continued observations on permanent plots also allow us to examine potential relationships between the cankers, scale, and leaf area.

Laboratory studies: A subset of cankers without C. pinea fruiting bodies were collected from all diameter classes of white pine and plated on three different media in an attempt to isolate and identify any pathogens. Subsequent pathogenicity tests will be conducted with isolates of unknown pathogenicity on white pine saplings. Pathogenicity tests will typically involve two isolates per fungal species and five inoculated white pine saplings in four plots.

Market Models⁽¹¹⁾

The development of a detailed market model requires extensive input data preparation on: current and future supply inventory under common management regimes, current and future demand at mills and concentration facilities, cost and timing of management activities, the transportation costs that link supply and demand, and estimation of elasticities of demand for forest products at processing centers. Forest Inventory and Analysis data (USDA-USFS) of current standing forest structures along with predictions of future stand trajectories developed using FVS or other growth and yield modeling program will comprise the bulk of input data. Once input data are compiled, a dynamic, spatial-equilibrium price endogenous linear programming market model will be programmed using GAMS (General Algebraic Modeling System). Once constructed, the market model can be maintained through minor periodic data updates and used well into the future to enable understanding of potential responses within the forest products market to changes in resource, demand, or policy environment. Changes in white pine abundance and lumber quality can be evaluated using this model.  The model will be the first of its kind in the Northeast but has been used elsewhere effectively. Model effectiveness will be evaluated by scholars and stakeholders familiar with the forest products industry in Maine to ensure that outputs follow reality.

Opportunities for Collaborative Work

Studies that have been limited to regional areas could be expanded to the range of white pine by the multistate project members.  Work that could be planned for additional funding include:

1. The white pine Matsucoccus scale is now confirmed as affecting white pine throughout its range, but current studies on the insect are limited to the south. The white pine multistate project could utilize the experience gained from the southern studies and begin similar scale studies in the Northeast and Lake States regions. 
2. More intensive studies on pinea cankers are occurring in the Northeast, but the fungus is distributed throughout the range of white pine. Characteristics of cankers associated with C. pinea infections (high hazard sites, canker severity as related to tree size and age, climatic conditions associated with higher canker incidence) can be expanded to the other regions to look for consistencies and differences.
3. Studies in the south indicate that Matsucoccus scale and pinea are both found in the same damaged tissue suggesting that Matsucoccus scale feeding favors C. pinea infection.  This relationship needs a high priority for study throughout the range of white pine.
4. A primary fungus associated with white pine needle blight, acicula, is a common needle pathogen associated with brown spot needle disease on southern pines. There are abundant studies that can be proposed for L. acicula such as genetic and behavioral similarities and differences between the regions and between species of pine. 
5. Range-wide studies on health issues affecting white pine will follow gradients in temperature and precipitation. Examining how health issues vary between cooler temperatures of northern regions to the warmer temperatures of southern regions will help clarify how climate change will affect the health issues, especially if warmer temperatures will affect the northern regions of white pine.
6. Develop white pine management recommendations across its distribution.

Joint Planning and Coordination

Annual meetings will be used as the primary method to facilitate cooperation, foster teamwork, and create synergism among project collaborators.  In addition to the initial collaborators, additional scientists will be encouraged to participate in the project to expand the scope and regional extent of the multistate project.  A web site, shared “cloud –drive”, and virtual meetings will be used to facilitate communications and data sharing during the year. 

The Multi-state Project meetings will have two functions:  1) Provide an opportunity to improve the effectiveness of current projects, and 2) develop a multi-institutional research proposal to be submitted to a national-level competitive research grant, such as the AFRI program.  

The joining of resources and expertise will expand the potential for accomplishments among the group.  For example, project members can expand sampling potential by sharing sampling expertise, by sharing scale and fungal identification expertise, and by making available dendrochronological processing and analysis available to more scientists.   

Measurement of Progress and Results

Outputs

• Standardization of methods (Objectives 1 & 2): Comments: <ul><li>Site selection and plot measurements <li>Canker quantification <li>Sampling for Matsucoccus <li>Sampling for canker fungi <li>Foliar measurements</ul>
• Sharing of expertise, equipment, and facilities (Objectives 1-3) Comments: <ul><li>Scale identification <li>Fungal identification <li>Dendrochronological techniques <li>Stem flow <li>Lumber processing equipment (mill, kiln, planer, etc) <li>Silvicultural recommendations</ul>
• Pooling of data (Objectives 1 &2) Comments: <ul><li>Data will be shared across the white pine region so that regional and range-wide trends can be recognized. This sharing will be especially important for assessing if climatic trends are driving any of the white pine health issues.</ul>
• Development of management recommendations (Objective 3) Comments: <ul><li>Members will present proposed management recommendations for addressing white pine health issues in each region. <li>Common strategies as well as regional differences will be clarified <li>A white pine health management publication will be developed</ul>

Outcomes or Projected Impacts

• Forest health specialists and the state and federal level are familiar with factors involved in white pine health issues. Damage and causes can be recognized and properly identified. <ul><li>Members will present proposed management recommendations for addressing white pine health issues in each region. <li>Common strategies as well as regional differences will be clarified <li>A white pine health management publication will be developed</ul>
• Forest managers and lumber mill operators can make realistic estimates of losses due to white pine health issues and use the information for making decisions. <ul><li>Members will present proposed management recommendations for addressing white pine health issues in each region. <li>Common strategies as well as regional differences will be clarified <li>A white pine health management publication will be developed</ul>
• Forest managers can take preemptive and reactive actions to minimize damage due to white pine health issues. <ul><li>Members will present proposed management recommendations for addressing white pine health issues in each region. <li>Common strategies as well as regional differences will be clarified <li>A white pine health management publication will be developed</ul>
• Market models include data concerning white pine health issues to provide better output for decision-making <ul><li>Members will present proposed management recommendations for addressing white pine health issues in each region. <li>Common strategies as well as regional differences will be clarified <li>A white pine health management publication will be developed

Milestones

(2017):<br>Development of a research proposal by project members will occur over a multi-year time frame (Objectives 1 & 2): <ul><li>Have presentation by an invited speaker who will provide key factors needed for successful AFRI submissions. <li>Project members will identify current strengths and weakness in knowledge and understanding. <li>The discussions will identify priorities for developing a research proposal that could potentially involve a project across the range of white pine <li>Responsibilities wil be assigned for developing specific parts of the proposal </ul> <p>A white pine health management publication will be developed (Objective 3): <ul><li>Development of recommendations</ul>

(2018):<br>Development of a research proposal by project members will occur over a multi-year time frame (Objectives 1 & 2): <br>• Draft of proposal is ready and reviewed at the meeting. Submission planned for 2018. <p>A white pine health management publication will be developed (Objective 3): <br>• Development of recommendations

(2019):<br>Development of a research proposal by project members will occur over a multi-year time frame (Objectives 1 & 2): <br>• If needed, proposal revisions are discussed. Resubmit in 2019 <p>A white pine health management publication will be developed (Objective 3): <br>• Agreement on publication content, assignment of sections to write.

(2020):<br>Development of a research proposal by project members will occur over a multi-year time frame (Objectives 1 & 2): <br>• Work on funded project <p>A white pine health management publication will be developed (Objective 3): <br>• Final draft approved and submitted for publication

(2021):<br>Development of a research proposal by project members will occur over a multi-year time frame (Objectives 1 & 2): <br>• Work on funded project <p>A white pine health management publication will be developed (Objective 3): Year 5 <br>• Revise and submit manuscript if needed.

Projected Participation

View Appendix E: Participation

Outreach Plan

The findings of the multistate project on white pine health issues will be communicated to two audiences, stakeholders who value white pine and researchers. 

To reach the stakeholders, we can use the network of U.S. Forest Service and state agency specialists who want to be involved in the multi-state project.  These people work on forest health in Georgia, Maine, Minnesota, New Hampshire, Virginia, and West Virginia and have direct connection to stakeholders. The team members also have annual meetings with colleagues from other states during which information can be shared.  As reports and publications are generated by the project, the information can be immediately distributed through the existing network of forest health professionals at the state and federal level.

To reach researchers, team members on the multi-state project will attend conferences related to forest health (Entomological Society of America, American Phytopathological Society, Ecological Society of America, Society of American Foresters) to report on new findings as they develop.  Developing research proposals and subsequent publications in peer-reviewed journals will communicate the project’s findings to our research peers. 

Organization/Governance

The project will use the standard form of governance.  Participating institution will send at least one representative to an annual meeting, and meeting attendees will be considered the Technical Committee.  The meeting will be held in conjunction with another professional meeting or workshop, such as the Northeast Forest Pest Council or the Southern Forest Insect Work Conference.  The Technical Committee will elect officers of chair, chair-elect, and secretary at the annual meeting. The chair-elect normally succeeds to chair the following year. Each officer is eligible for re-election.  Sub-committees focused on particular objectives will confer as needed by teleconference to plan and coordinate details of the multi-state project.

Literature Cited

1. Asbjornsen, H.   Assessing tree species' ecohydrological functions and climate change impacts on New Hampshire's forest ecosystems.  McIntire-Stennis Project, New Hampshire Experiment Station, Durham.  PROJ NO: NH00071-M
2. Asbjornsen, H., I. Munck, and C. McIntire. 2015. Impact of White Pine Needle Damage and management interventions on forest health across the Northeastern United States. Project funded by a USDA Forest Health Monitoring grant.
3. Asaro, C. 2011. What is killing white pine in the highlands of Western Virginia? in: Forest Health Review May 2011 Virginia Department of Forestry, Charlottesville, VA.
4. Booth, J.M., and P.J. Gullan. 2006. Proceedings of the Entomological Society of Washington 108: 749-760.
5. Broders, K.D., G.J. Boland. 2010. Molecular diagnostic assay for detection of the butternut canker pathogen Sirococcus clavigignenti-juglandacearum. Plant Disease 94:952-958.
6. Broders, K.D., I. Munck.   White Pine Needle Damage (WPND) in the northeast: Characterization and distribution of fungi associated with needle damage and effect of climate on disease development and spread.  Evaluation Monitoring Program, Forest Health Management, USDA Forest Service, 2012-2015.
7. Broders, K., I. Munck, S. Wyka, and B.E. Iriarte. 2015. Characterization of fungal needle pathogens associated with white pine needle damage (WPND) in the Northeastern North America. Forests. 6: 4088 -4104
8. Capretti, 1978. Caliciopsis pinea Peck on Pinus pinaster Ait and Pinus insignis Dougl in Italy. Phytopathologia Mediterranea 17:101-104.
9. Chhin, S., J. O’Brien.   White pine decline in the Great Lakes region.  Evaluation Monitoring Program, Forest Health Management, USDA Forest Service, 2013-2014.
10. Costanza, K.L., W.H. Livingston, S. Fraver.   Impact of Caliciopsis canker on white pine wood quality and lumber yield.  USDA-ARS Forest Products R&U, University of Maine Forest and Agricultural Experiment Station. Funded 2015-2017.
11. Crandall, M.   Forest management for forest and human community health in a time of transition.  Maine Agricultural and Forest Experiment Station, University of Maine, Orono, Project No. ME041521.
12. Desprez-Loustau, M.-L., B. Marcais, L.-M. Nageleisen, D. Piou, and A. Vannini. 2006. Interactive effects of drought and pathogens in forest trees. Annals of Forest Science, 63: 597-612.
13. Frament B., R. Lilja, and M. Weeks. 2011. 2010 Aerial Detection Survey Results: Durham Field Office, Forest Health Management, USDA Forest Service.
    http://www.na.fs.fed.us/fhp/pubs/aerial/aerial_survey_results_dfo10.pdf
14. Frumhoff PC, McCarthy JJ, Melillo JM, Moser SC, and J. WD (2007) Confronting Climate Change in the U.S. Northeast. Synthesis report of the Northeast Climate Impact Assessment (NECIA). Cambridge, MA: Union of Concerned Scientist (UCS).
15. Funk, A. 1963. Studies in the genus Caliciopsis. Canadian Journal of Botany 41:530-543.
16. Funk, 1964. Extensions of the host ranges and distribution of Caliciopsis species on western conifers. Plant Disease Reporter 48.
17. Gandhi, K.J.K., M. Cram, F. Koch, C. Asaro, D. Coyle, A. Mech. Monitoring the health and effects of scale/pathogen complex on eastern white pines in the southern Appalachians.  Evaluation Monitoring Program, Forest Health Management, USDA Forest Service, 2013-2016.
18. Gandhi, K.J.K., D.R. Coyle, K. Klepzig, F. Koch, L. Morris, J.T. Nowak, B. Otrosina, B. Smith. 2014. Evaluating the role of rhizophagous insects and associated fungi in pine health issues in southeastern U.S. Entomological Society of America Annual Conference, 15-18 Nov 2014, Portland, OR.
19. Griesmer, R., and Adams, G.C. 2012. Investigation of the roles of spittlebugs, cankers, stem density and ground water in branch dieback and mortality in regenerating white pine. Michigan Academician, 40 (3).
20. Lancaster, K.F., W. Leak. 1978. A silvicultural guide for white pine in the Northeast. Gen. Tech. Rep. NE-41. Broomall, PA: U. S. Department of Agriculture, Forest Service, Northeastern Forest Experiment Station. 13 p.
21. Leak, W.B., M. Yamasaki. 2013. Effects of low-density thinning in a declining white pine stand in Maine. Res. Note NRS-170. Newton Square, PA: U.S. Department of Agriculture, Forest Service, Northern Research Station. 6p.
22. Lombard, K. 2003. Caliciopsis canker (pine canker) Caliciopsis pinea. in: UNH Cooperative Extension Publication UNH, Durham, NH.
23. Lowell, E.C., S.A. Willits. 1998. Lumber recovery from beetle-killed spruce trees, Kenai Peninsula, Alaska. Western Journal of Applied Forestry 13:54-59.
24. McCormack, H.W. 1936. The morphology and development of Caliciopsis pinea. Mycologia 28(2): 188-196.
25. McIntire, C., H. Asbjornsen, I. Munck, K. Broders, B. Livingston. 2014. Interactive effects of White Pine Needle Damage and climate change on forest health across the Northeastern United States. XXIV IUFRO World Congress, Salt Lake City, UT. Oct. 5-11.
26. Mech, A.M., C. Asaro, M.M. Cram, D.R. Coyle, P.J. Gullan, L.G. Cook, and K.J.K. Gandhi. Matsucoccus macrocicatrices (Hemiptera: Matsucoccidae): First report, distribution, and association with symptomatic eastern white pine in the southeastern United States.  Journal of Economic Entomology 106: 2391-2398. 
27. Munck, I., K. Lombard, W.D. Ostrofsky, K. Broders, W. Livingston.   Lumber yield based on Caliciiopsis symptom severity ratings.  STDP Program, Forest Health Management, USDA Forest Service, 2015-2017. 
28. Munck, I., K. Lombard, W.D. Ostrofsky, K. Broders, W. Livingston, T. Luther. 2014. Extent and severity of Caliciopsis canker of white pine: Risk assessment, evaluation of silvicultural management tools, and diagnostic assay Evaluation Monitoring Program, Forest Health Management, USDA Forest Service, 2014-2017.
29. Ray, W.W. 1936. Pathogenicity and cultural experiments with Caliciopsis pinea. Mycologia 28(3):201-208.
30. Schulz, A., C. Asaro, D.R. Coyle, M. Cram, R. Lucardi, A.M. Nech, K.J.K. Gandhi. 2014. Mapping the distribution of a potentially new tiny terror in southern white pine forests. Entomological Society of America Annual Conference, 15-18 Nov 2014, Portland, OR.
31. Smith, D.R., and G.R Stanosz. 1995. Confirmation of two distinct populations of Sphaeropsis sapinea in the north central United States using RAPDs. Phytopathology, 85:699-704.
32. Solheim, H., T.B. Torp, and A.M Hietala. 2013. Characterization of the ascomycetes Therrya fuckelii and pinii fruiting on Scots pine branches in Nordic countries. Mycological Progress, 12: 37-44.
33. Speers, J.H. 2010. Fundamentals of Tree Ring Research. Univ. Arizona Press. 352 p.
34. 2011. A/R Methodological Tool: Demonstrating appropriateness of allometric equations for estimation of aboveground tree biomass in A/R CDM project activities. Version 01.0.0.
35. Watson, W.Y., G.R. Underwood, and J. Reid. 1960. The Canadian Entomologist XCII: 662-667.
36. Wendel, G.W., and H.C. Smith. 1990. Eastern White Pine inS. Burns and B.H. Honkala (eds).  Silvics of North America, Vol. 1, Conifers.  USDA Forest Service, Agricultural Handbook 654.

Attachments

Land Grant Participating States/Institutions

CT, GA, MA, ME, MI, NH, NV, VT, WI

Non Land Grant Participating States/Institutions

Forest Service, USDA Forest Service, USDA Forest Service - Minnesota