Using Provenance Test Data to Inform Ecological Restoration in the Tahoe Basin. Jessica W. Wright Pacific Southwest Research Station 1731 Research Park Drive Davis, CA, 95618 Phone: 530-759-1742 Fax: 530-747-0241 jessicawwright@fs.fed.us. SNPLMA Theme: Forest Health, Subtheme 1c: Impact of climate change on ecological communities and the evaluation of adaptation strategies Agency personnel who will be directly involved in the project: Thomas Blush, Regional Geneticist, R5, USDA-Forest Service, IFG, Placerville CA. Arnaldo Ferreira, Geneticist, USDA- Forest Service, Placerville Nursery, Camino CA. Phil Cannon, Regional Forest Pathologist, R5, USDA- Forest Service, Vallejo, CA. Richard Sniezko, Geneticist, USDA-Forest Service, Genetic Resource Center, Dorena, OR. Grants Contact Person at PSW: Brian Hanlon, 510-559-6309, bhanlon@fs.fed.us. Total Funding requested- $91,797 Using Provenance Test Data to Inform Ecological Restoration in the Tahoe Basin. J. Wright Proposal Narrative Project abstract Climate change is a major concern for forest managers involved in ecological restoration, in part because they are responsible for planting trees that will be part of a healthy and resilient ecosystem for the next 100-200 years. When beginning an ecological restoration project, managers have to decide what seed stock to plant in a given location. Traditionally, seed zones and elevation bands are used to determine what seed stock to deploy. However, climate change is expected to change the growing conditions in those seed zones and elevation bands. Managers need to know now what seed stock will result in a healthy and resilient forest, long after their ecological restoration project has been completed. Moreover, seed procurement efforts, which are aimed at ensuring that seeds are available for ecological restoration after the next major fire event (wherever or whenever that might be), also need to be informed of where seeds need to be collected from in order to be ready to be deployed. It is the goal of this project to provide information from four decades of provenance tests of Ponderosa, Jeffrey, and Sugar pine that will help guide those decisions. Justification Statement The goal of this proposal is to provide information to land mangers that will inform them in the adaptation strategy of “planting species or genotypes that may be more resilient to climate change” (SNPLMA RFP, subtheme 1c.). Over the past 90 years, the USDA-Forest Service, Pacific Southwest Research Station, Institute of Forest Genetics has been planting, maintaining and measuring provenance tests, the type of study that will provide exactly the kind of data needed to understand species responses to climate change. The goal of this project is to use data already available from these tests to understand how species respond to novel climates. Concise background and problem statement Climate change is expected to impact the forested ecosystems of the Lake Tahoe Basin, and indeed, all of California. Management agencies need to develop adaptation strategies that will ensure healthy and resilient ecosystems into the future (Bolte et al., 2009). Anticipated changes in climate will include changes in temperature, precipitation and length of the growing season. Trees locally adapted to their native climate may not be able to survive or grow in novel climates. It may, in fact, be necessary to look outside the Basin to find trees adapted to the predicted climate in the Basin. What is needed now is research that focuses on tree responses to novel climates. Ecological restoration of forested habitats begins with a simple, but important decision- what seed stock should be deployed in a given location. Managers have to determine what seeds will result in the healthiest and most resilient forests in the decades to come. In order to make informed decisions, they need data on how seed stocks might respond to novel climates- the ones predicted under climate change. 2 Using Provenance Test Data to Inform Ecological Restoration in the Tahoe Basin. J. Wright Provenance tests are studies where genotypes from different sources (or provenances) within a tree species are planted onto a common, relatively homogeneous, site. Often those sources can represent the entire species range- or just a limited subset of that range. The goal of a provenance test is to determine patterns of genetic variation- the “genetic architecture” of a species over its entire range. However, the experimental design of these tests is also ideal for understanding how trees respond to novel climates. Individuals were moved from one climate, planted in a new one, and monitored for survival and growth. Usually these tests comprise multiple test sites, so, we can see how different individuals respond in different climates (a genotype by environment or GxE response). Provenance tests have recently become the focus of increased research attention, as their usefulness for understanding the associations between tree growth and performance and climate is being more fully understood (Mátyás, 1997, 1996, 1994). The work proposed here will focus on three sets of these tests which are highly relevant to the forested ecosystems of the Tahoe Basin. These trees have been growing since the 1970s, 1980s and 1990s and include a great deal of data which is already collected and ready for analysis. The first is a test in Ponderosa (Pinus ponderosa Laws.) and Jeffrey pine (P. jeffreyi Grev. & Balf.). It was planted in 1973. The test is replicated at three sites, including Angora Creek in the Tahoe Basin, Mt Danaher outside the town of Camino and Lincoln Hill on the Eldorado National Forest. It includes four sources of Ponderosa pine (including one from the Basin) as well as three sources of Jeffrey pine- also including one from the Basin. These sources span the elevational range of the species- 500, 4000 and 6000 ft for Ponderosa and 4000, 6000 and 8000 ft for Jeffrey. Each source is represented by progeny from ten randomly selected parent trees of average or better form. Seeds from each of these elevations were collected in very different climates (Table 1). This allows us to examine responses along different axes of climate variation- for example, mean annual temperature and precipitation, and the length of the frost-free period (a measure of the length of the growing season). The Angora Creek test site was burned as a result of the recent fires in that area, however, in the test it was a ground fire, and the canopy was mostly not impacted (see photograph). Only about 10% mortality was observed. This was confirmed by a visit to the site in 2010 (R. Stutts Pers. Com.). This test has not been evaluated since 1990, so, we are requesting funds for a FS-field crew to measure this test. This will also allow us to look at the mortality caused by the recent fire, and see if there was any source-specific effect. 3 Using Provenance Test Data to Inform Ecological Restoration in the Tahoe Basin. J. Wright Table 1. Data for the Planting sites and seed sources in the Ponderosa/ Jeffrey pine provenance tests, planted by IFG Staff in 1973. Planting Sites Angora Creek Lincoln hill Mt Danaher Seed sources Jeffrey pine Lake Valley High Meadows Forest Hill Ponderosa pine Salmon Falls Emerald Bay Silver Fork Forest Hill Elevation (ft) Latitude Longitude MAT (°C) MAP (mm) Frost Free Period (days) Average Height at Angora Creek in 1990 (cm) % alive in 1990 at Angora Creek 6600 4480 3400 38.87 38.66 38.72 120.04 120.50 120.65 6.7 11.4 13.5 938 1363 1271 108 194 210 6300 8080 3840 38.90 38.90 39.10 120.02 119.90 120.77 7.3 4.2 12.3 842 946 1521 110 69 200 199.89 212.34 191.21 91.70% 90.00% 59.17% 477 6300 3950 3840 38.75 38.95 38.77 39.10 121.07 120.10 120.32 120.77 16.2 7.0 12.1 12.3 633 911 1139 1521 291 104 191 200 214.74 210.55 200.41 200.48 47.92% 77.5% 43.80% 75% In Sugar pine (Pinus lambertiana Dougl.) there are two tests available for study- Sugar Pine Heritage (1984) and Harrell (1992). The first includes four test sites, two in California and two in Oregon, the one closest to the Basin is Fitch Rantz, on the Eldorado National Forest. There are 69 sources of Sugar pine included in this test, spanning the entire range of this species, from Mexico to Oregon. Each source is represented by progeny from 10 single-tree collections. There is one source from the Basin. Because of the large number of provenances included in this study, it is possible to look at responses of these single tree collections to climatic conditions that are different from the temperature and precipitation conditions in which they have evolved. Earlier results of this test have been published (Kitzmiller, 2004), however those analyses did not include an in depth analysis of climate, or the latest growth measurements. The second test, Harrell, has only one planting site but 124 sources of Sugar pine are planted on that site. This test was designed in a very practical manner, with seed sources identified by seed zone and elevational band, in the same way that is used operationally by Forest Service Silviculturists. Seeds were bulked from collections made within each seed zone and elevation band. With help from Michael Landram (R5 Regional Silviculturist - now retired) and Matthew Brokash (State and Private Forestry- Forest Health Monitoring), we calculated the averages for a set of climate variables for each of the seed zones and elevation bands in California, so it will be possible to study the impact of the 4 Using Provenance Test Data to Inform Ecological Restoration in the Tahoe Basin. J. Wright changes in climate experienced by moving the provenances to the Harrell trial. This range-wide information, provided by these tests, will be useful to help predict how various seed sources of Ponderosa, Jeffrey and Sugar pine might perform when used for ecological restoration in the Lake Tahoe Basin where a new set of climatic parameters has been predicted. d. Goals, objectives and hypotheses to be tested. Goal: To understand tree species responses to anticipated climate change. This information will inform ecological restoration and seed procurement and deployment efforts on the Basin in light of predicted changes in the local climate. Objective 1: to use data from a provenance test of Ponderosa and Jeffrey pine, including the Angora Creek test site to understand how trees respond to novel climates on a broad scale. Objective 2: to use data from two provenance tests in Sugar Pine to understand tree responses to novel climates on a narrow scale. Hypothesis: While trees are expected to perform best (grow, survive and reproduce well) in their native climates, there may well be a range of climates where populations will have nearly-optimal performance. There will also be climates (or specific climate variables) that tree species respond negatively to. These tests will allow us to more accurately define seed transfer guidelines in the Basin, informing both seed procurement operations, as well as ecological restoration. Approach, methodology and location of research To achieve our Objectives we will use a combination of growth and survival data along with climate data for the test sites as well as the source populations- both for the current climate, as well as projected future climate. We will use this data to examine how tree species might respond to climate change. We will relate these data to the Basin directly by examining test sites planted on the Basin, along with provenances collected on the Basin as well as using current and projected climate models for areas of the Basin currently occupied by pine forests. Survival and growth data, including insect and disease responses, has been collected at our field sites since their original planting, and at regular intervals ever since. There are ten growth and survival measurements taken at all four sites in the Sugar Pine Heritage experiment, the most recent being 2009. The Ponderosa/Jeffrey tests were measured four times, with the most recent being 1990. Here we request funding to complete a 2012 measurement of 39-year growth data. Maps and data sheets of all three test sites are on file at the Institute of Forest Genetics, PSW- Placerville. In addition, many of the trees in this trial were tagged (which are reported to have survived the Angora fire). Since much of the data is ready to analyze now, we anticipate a very quick start to this project once funding commences. Fortunately there is a large literature available to guide our analyses (eg. Rahi et al. 2010; Leibing et al., 2009; Thomson et al., 2009; Thomson and Parker, 2008; Chuine et al, 2006; Sáenz-Romero et al 2006; 5 Using Provenance Test Data to Inform Ecological Restoration in the Tahoe Basin. J. Wright Rehfeldt 2004; Rehfeldt et al, 2002; Rehfledt et al 1999). However, each Provenance test is unique, and these methods will have to be adapted for our particular study. We will use survival analyses (including Cox regression models which can incorporate co-variates such as climate variables into the models (Allison, 1995)) to look at the association between survival and provenance. We expect survival to be strongly influenced by climate, as well as other indirect responsesincluding White Pine Blister Rust (WPBR). In 2009 and 2010, all of the Sugar Pine test sites were surveyed for WPBR (R. Sniezko, unpublished data). As described below (Relationship to Previous Research), the two groups will work together to put both pieces of the puzzle together- response to introduced pathogens as well as climate. As an example of a climate-focused analysis, we compared survival curves to the change in mean annual precipitation for individuals moved from their source provenance to Fitch Rantz. The results were very clear that trees for which Fitch Rantz was a drier habitat, survival was higher. Trees that came from a habitat drier than Fitch Rantz (Fitch Rantz was a wetter habitat) survival was lower (Figure 1). However, these analyses were based on broad climate variables (mean annual precipitation), and a more in depth analysis using more sophisticated climate variables- for example, ones that describe when precipitation fell over the course of the year- winter snows vs summer rains- will shed more light on tree survival. 1 Figure 1. Survival of Sugar pine trees at the Fitch Rantz (FR) test site (6500 ft). Black lines indicate provenances for which FR is a drier habitat. Blue lines indicate provenances for which FR is a wetter habitat. Overall, trees from wetter habitats (for which FR was a drier habitat) performed better at Fitch Rantz. 0.9 Proportion surviving 0.8 0.7 0.6 0.5 0.4 600-800 mm less 200-400 mm less 0-200 mm more 400-600 more > 800 mm more 0.3 0.2 0.1 400-600 mm less 0-200 mm less 200-400 more 600-800 mm more 0 0 5 10 15 Years from planting 20 25 Growth data will be analyzed first using traditional Generalized Mixed Models (GMMs). Preliminary analyses of the Fitch Rantz data showed that trees from the different regions had different heights after 22 years of growth. Using a multivariate regression model (using a stepwise selection process) comparing growth and climate, location and elevation, we showed that longitude is the best predictor of plant growth. These models can be improved with better climate data. Location: We request funds to support a full-time Postdoctoral Researcher who will focus on completing these analyses and publishing the results. The Postdoctoral Researcher will be based in Davis, California, and will complete their analyses and writing there. They will make at least monthly (and probably more frequent) trips to Placerville to interact with Thomas Blush and Arnaldo Ferreira as their analyses are progressing. These interactions will ensure that management stays up to date with the latest findings, and that the questions that managers ask are addressed. 6 Using Provenance Test Data to Inform Ecological Restoration in the Tahoe Basin. J. Wright The field sites are located at the Angora Creek planting site in the Basin, as well as the Mt Danaher and Lincoln Hill test sites. To my knowledge, there are no provenance tests in Sugar pine currently growing on the Basin. Given the importance of Sugar pine in the forested ecosystems of the Basin, I feel it is critical to include this species in our analyses, even if that means using tests sites not found on the Basin. Relationship of the research to previous and current relevant research, monitoring, and/or environmental improvement efforts. Results for the Sugar pines concerning White Pine Blister Rust (WPBR) will be tied in with all other measurements and assessments that have been made in studies on this disease across the entire range of trials in the Western United States where five-needle-pines have been studied for their vulnerability to this disease. For example, Richard Sniezko, Geneticist at the Dorena Genetic Resource Center (Dorena, OR), completed a detailed WPBR assessment of the trees at all five of the Sugar Pine test sites in 2009 and 2010. We will work with Richard’s group to bring together analyses focusing on Climate with those focusing on WPBR. Given the importance of Sugar Pine, and WPBR in the Lake Tahoe Basin, such a collaborative effort is critical to ensure a complete understanding of the environmental stresses experienced by Sugar pine trees on the Basin. SNPLMA and the Nevada Division of State Lands (NVDSL) Lake Tahoe License Plate Program, has funded previous work examining the Conservation Genetics of Sugar pine on the Basin (Evaluation of montane forest genetic resources in the Lake Tahoe Basin: Implications for conservation, management and adaptive responses of Pinus lambertiana to environmental change- Vogler, Maloney and Neale). That work has produced detailed data on variation at molecular genetic markers across the valley- giving insight to fine scale genetic variation. The work proposed here will also focus on the conservation genetics of Sugar pine, but using a more trait-focused approach. However, with additional funding (budget available upon request- approximately $50 per sample, with a minimum of 700 samples), it would be very simple to sequence the same molecular genetic data at the Sugar Pine Heritage tests as used by Vogler et al., allowing a comparison of genetic variation in trees sampled across the Basin to the entire species range. If this side-project is funded, Detlev Vogler, Patricia Maloney and David Neale have agreed to share their molecular genetic data with me to allow for an integration of the two projects. Strategy for engaging with mangers and obtaining permits Mangers have been involved in various aspects of this project for the last several years. For example, last year, Phil Cannon (R5 Regional Pathologist) organized a meeting at the Regional Office to discuss the provenance test resources in the Region, and how we could get answers from those tests. Thomas Blush (R5 Regional Geneticist), and Arnaldo Ferreira’s predecessor (Geneticist in charge of seed procurement) were also at the meeting, along with Michael Landram (Regional Silviculturist, now retired). Therefore much of the motivation for preparing this proposal is to address management needs that have already been strongly expressed. This study has the potential to change land management strategies in the 7 Using Provenance Test Data to Inform Ecological Restoration in the Tahoe Basin. J. Wright Basin. For example, the Forest Service might use the results to change their recommendations for the best source of planting stock for ecological restoration activities in the Basin. Having a Postdoc funded through SNPLMA will speed the process of data analysis and manuscript publication, and bring additional expertise to the project. The Postdoc will meet regularly with Forest Service Geneticists in what should be an open exchange of results and ideas. These meetings will be expanded as needed and desired to include managers from the Basin who are responsible for ecological restoration. There is special interest in moving these studies along so that results can be presented at the Western International Forest Disease Work Conference (WIFDWC) which will hold its next week-long conference at Lake Tahoe during the second week of October, 2012. About 100 forest managers and forest pathologists are expected to attend. Forest Service (Forest Health Protection) pathologists from R5 have also agreed to help measure these studies and in particular those parameters that relate to forest health, survival and growth in response to White Pine Blister Rust, following fire (at Angora Creek) and in response to changing climates. The research proposed here will not require permits, as the focus of the work will be on already collected data, and non-invasive tree measuring (heights and diameters). Appropriate Forest Service staff will be informed whenever researchers will be on site, and we will work with them if there are ever any issues regarding the tests. Description of deliverables The analyses completed as part of this research will give us new insight to the response of different provenances of several of California’s most significant tree species to new climates to which they may soon be exposed. This insight will be disseminated in a number of ways. Peer-reviewed journal articles will be one approach, of course. However, land managers will learn about those results long before they are seen in published journals- we will be interacting with the R5 Regional Geneticist and his staff throughout the process of analyzing data and manuscript preparation. Together we will determine the most effective way to disseminate results to resource Managers on the Basin. This will include workshops and presentations- focusing on VTC and Webinars to avoid Travel Restrictions faced by many Forest Service employees. Also, all data from this project will be made publically available on the Center for Forest Provenance Data website (http://cenforgen.forestry.oregonstate.edu/). This will make the data available to other researchers for future analyses. 8 Using Provenance Test Data to Inform Ecological Restoration in the Tahoe Basin. J. Wright Timeline Start date: July 2012- 3rd Quarter of 2012- milestones reported by Quarter below. July 2012- Funding commences- Postdoctoral researcher is hired, brought up-to-date on data analysis. Meetings held with Blush and Ferriera. October 2012- Data analysis ongoing. Meetings with managers ongoing. 1st Quarterly report completed. January 2013- Data Analyses in progress, manuscript preparation beginning. Meetings with managers ongoing. Quarterly report completed. April 2012- Manuscripts submitted and in review. Meetings with managers ongoing. Quarterly report completed. July 2012- Manuscripts accepted for publication. Meetings with managers ongoing. Quarterly/final report completed. 9 Using Provenance Test Data to Inform Ecological Restoration in the Tahoe Basin. J. Wright IV. Literature Cited Allison, P.D. 1995. Survival Analysis Using SAS: A Practical Guide. Cary, NC: Sas Institute Inc. Bolte, A., Ammer, C., Löf, M., Madsen,P., Nabuurs, G., Schall, P., Spathelf, P., and Rock, J. 2009. Adaptive forest management in central Europe: Climate change impacts, strategies and integrative concept. Scandinavian Journal of Forest Research. 24:473-482. Chuine, I., Rehfeldt, G.E. and Aitken, S.N. 2006. Height growth determinants and adaptation to temperature in pines: a case study of Pinus contorta and Pinus monticola. Canadian Journal of Forest Research. 36:1059-1066. Kitzmiller, J.H. 2004. Adaptive genetic variation in Sugar pine. USDA Forest Service Proceedings, RMRS-P32. Leibing, C., Zonneveld, M.V., Jarvis, A., and Dvorak, W. 2009. Adaptation of tropical and subtropical pine plantation forestry to climate change: Realignment of Pinus patula and Pinus tecunumanii genotypes to 2020 planting site climates. Scandinavian Journal of Forest Research. 24:483-493. Mátyás, C. 1994. Modeling climate change effects with provenance test data. Tree Physiology. 14:797804. Mátyás, C. 1996. Climate adaptation of trees: rediscovering provenance tests. Euphytica. 92:45-54. Mátyás, C. 1997. Assessment of effects of environmental change on tree populations. In: Encycopedia of environmental control technology. P. Cheremishinoff, Ed. New Jersey, Gulf Publishing. Thomson, A.M., and Parker, W.H. 2006. Boreal forest provenance tests used to predict optimal growth and response to climate change: 1. Jack pine. Canadian Journal of Forest Research. 38:157-170. Thomson, A.M., Riddell, C.L., and Parker, W.H. 2009. Boreal forest provenance tests used to predict optimal growth and response to climate change: 2.Black spruce. Canadian Journal of Forest Research. 39:143-153. Rahi, A.A., Bowling, C. and Simpson, D. 2010. A red pine provenance test in northwestern Ontario: 48year results. The Forestry Chronicle. 86:348-353. Rehfeldt, G.E. 2004. Interspecific and intraspecific variation in Picea engelmannii and its congeneric cohorts: Biosystematics, genecology and climate change. USDA-Forest Service. RMRS-GTR-134. Rehfeldt, G.E., Tchebakova, N.M., Parfenova, Y.I, Wykoff, W.R., Kuzmina, N.A and Milyutin, L.I. 2002. Intraspecific responses to climate in Pinus sylvestris. Global Change Biology. 8:912-929. 10 Using Provenance Test Data to Inform Ecological Restoration in the Tahoe Basin. J. Wright Rehfeldt, G.E., Ying, C.C., Spittlehouse, D.L., and Hamilton, Jr., D.A. 1999. Genetic responses to climate in Pinus contora: Niche breadth, climate change and reforestation. Ecological Monographs. 69:375-407. Sáenz-Romero, C., Guzmán-Reyna, R.R., and Rehfeldt, G.E. 2006. Altitudinal genetic variation among Pinus oocarpa populations in Michoacán, Mexico. Implications for seed zoning, conservation, tree breeding and global warming. Forest Ecology and Management. 229:340-350. 11