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Risk of Genetic Maladaptation Due to Climate Change in Three Major European Tree Species

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Risk of Genetic Maladaptation Due to Climate Change in Three Major European Tree Species Received: 27 February 2017 | Accepted: 13 June 2017 DOI: 10.1111/gcb.13802 PRIMARY RESEARCH ARTICLE Risk of genetic maladaptation due to climate change in three major European tree species Aline Frank1 | Glenn T. Howe2 | Christoph Sperisen1 | Peter Brang1 | J. Bradley St. Clair3 | Dirk R. Schmatz1 | Caroline Heiri1 1Swiss Federal Institute for Forest, Snow and Landscape Research WSL, Birmensdorf, Abstract Switzerland Tree populations usually show adaptations to their local environments as a result of 2Department of Forest Ecosystems and natural selection. As climates change, populations can become locally maladapted Society, Oregon State University, Corvallis, OR, USA and decline in fitness. Evaluating the expected degree of genetic maladaptation due 3Pacific Northwest Research Station, USDA to climate change will allow forest managers to assess forest vulnerability, and Forest Service, Corvallis, OR, USA develop strategies to preserve forest health and productivity. We studied potential Correspondence genetic maladaptation to future climates in three major European tree species, Nor- Aline Frank, Swiss Federal Institute for Forest, Snow and Landscape Research WSL, way spruce (Picea abies), silver fir (Abies alba), and European beech (Fagus sylvatica). Birmensdorf, Switzerland. A common garden experiment was conducted to evaluate the quantitative genetic Email: [email protected] variation in growth and phenology of seedlings from 77 to 92 native populations of each species from across Switzerland. We used multivariate genecological models to associate population variation with past seed source climates, and to estimate rela- tive risk of maladaptation to current and future climates based on key phenotypic traits and three regional climate projections within the A1B scenario. Current risks from climate change were similar to average risks from current seed transfer prac- tices. For all three climate models, future risks increased in spruce and beech until the end of the century, but remained low in fir. Largest average risks associated with climate projections for the period 2061–2090 were found for spruce seedling height (0.64), and for beech bud break and leaf senescence (0.52 and 0.46). Future risks for spruce were high across Switzerland. However, areas of high risk were also found in drought-prone regions for beech and in the southern Alps for fir. Genetic maladaptation to future climates is likely to become a problem for spruce and beech by the end of this century, but probably not for fir. Consequently, forest manage- ment strategies should be adjusted in the study area for spruce and beech to main- tain productive and healthy forests in the future. KEYWORDS Abies alba, climate change, Fagus sylvatica, genecology, local adaptation, Picea abies, quantitative traits, relative risk of maladaptation, seedling common garden experiment, water availability 5358 | © 2017 John Wiley & Sons Ltd wileyonlinelibrary.com/journal/gcb Glob Change Biol. 2017;23:5358–5371. FRANK ET AL. | 5359 1 | INTRODUCTION of local maladaptation to future climates for populations and species. Relative risk quantifies the difference between populations adapted Tree species of temperate and boreal regions often exhibit multiple to two different climates, e.g., past and future climates, taking into genetic adaptations to their local climates (Alberto et al., 2013; Bus- account the amount of within-population genetic variation (Camp- sotti, Pollastrini, Holland, & Bruggemann,€ 2015). For example, the bell, 1986; St. Clair & Howe, 2007). The quantitative genetic statis- timing of bud break and bud set typically varies along latitudinal and tics and climate associations needed to calculate relative risk of elevational gradients, and drought tolerance appears to be higher in climate change can be obtained from common garden experiments populations from dry environments. Such climatic adaptations are (e.g., St. Clair, Mandel, & Vance-Borland, 2005). In addition, high- considered to result from diversifying natural selection (Savolainen, resolution climate projections are needed, particularly changes in Pyh€ajarvi,€ & Knurr,€ 2007). As local climates change, however, tree temperature and precipitation. species may become maladapted if evolutionary adaptation does not In this study, we focused on Norway spruce (referred to as keep pace with ongoing environmental changes (e.g., St. Clair & “spruce”), silver fir (“fir”), and European beech (“beech”, Fagus sylvat- Howe, 2007). The resulting genetic maladaptation could lead to ica L.), three major European tree species whose ranges partly over- reduced fitness or even local extinction of current tree populations. lap in Central Europe (EUFORGEN, 2009). The climatic conditions in This has the potential to affect forest composition, structure, and Central Europe are expected to change markedly by the end of the stability, with potential negative consequences for the provision of century (2051–2080) compared to the second half of the 20th cen- forest goods and services (Lindner et al., 2010). tury (1951–2000), with mean summer temperatures increasing Different levels of climate-induced maladaptation are expected between 1.3 and 2.7°C, and summer precipitation decreasing by up to occur in different tree species. Adaptive specialists, such as lodge- to 25% (Lindner et al., 2014). The impact of climate change will likely pole pine (Pinus contorta), Douglas-fir (Pseudotsuga menziesii), and vary among regions and locations, and is considered to be especially Norway spruce (Picea abies [L.] Karst.), show high levels of climate- pronounced in mountainous areas such as Switzerland (Pepin et al., related population differentiation. They are likely at higher risk of 2015). Here, under the A1B scenario, mean summer temperatures maladaptation than are adaptive generalists such as white pine (Pinus are projected to increase by more than 4°C by the end of the 21st monticola), western redcedar (Thuja plicata), and silver fir (Abies century compared to the period 1980–2009 (CH2011, 2011). The alba Mill.) (Frank, Sperisen, et al., 2017; Rehfeldt, 1994; St. Clair & expected changes in temperature and precipitation may affect Howe, 2007). At the population level, the degree of maladaptation growth, fitness, and the distribution of all three tree species (Gessler might vary across the landscape, depending on the amount of et al., 2007; Hanewinkel, Cullmann, Schelhaas, Nabuurs, & Zimmer- within-population genetic variation, environmental heterogeneity, mann, 2013; Lebourgeois, Rathgeber, & Ulrich, 2010; Meier, adaptational lag, and the local extent of climate change (St. Clair & Edwards, Kienast, Dobbertin, & Zimmermann, 2011; Nothdurft, Howe, 2007). However, estimates for climate-induced maladaptation 2013; Nothdurft, Wolf, Ringeler, Bohner,€ & Saborowski, 2012). between and within species are rare. Recent results from a seedling common garden study have shown Knowledge of trees’ maladaptation to future climates is valuable that spruce, fir, and beech in Switzerland are characterized by dis- for developing and refining forest management strategies and tools, tinct genecological patterns (Frank, Pluess, Howe, Sperisen, & Heiri, such as seed transfer guidelines, that could help to mitigate negative 2017; Frank, Sperisen, et al., 2017). Genetic clines for spruce are climate change impacts on forest ecosystems (Park et al., 2014). Tra- pronounced along temperature gradients, whereas for fir and beech, ditionally, seed transfer guidelines and seed zones have been used genetic clines are weaker, and are mostly found along gradients of to conserve or enhance forest productivity and timber quality (Lan- temperature and water availability. These genecological patterns sug- glet, 1971). Such guidelines should now be reconsidered to preserve gest vulnerability to climate change is larger for spruce than for fir forest health and productivity in potentially warmer and drier cli- and beech. However, quantitative estimates of maladaptation to cli- mates. Forest managers may, for example, select seed sources that mate change are lacking for these and most other tree species (but match the future local climate of a particular forest site, and use see St. Clair & Howe, 2007). In particular, we have no information such “preadapted” plant material for reforestation or admixture about the differences in potential maladaptation between spruce, fir, within existing stands (a.k.a., assisted migration or assisted gene and beech, or the variation in risk across the landscape. Such infor- flow; Aitken & Whitlock, 2013; Williams & Dumroese, 2013). For mation, however, could form the basis for science-based recommen- that purpose, forest managers need to know which species and dations for forest management, particularly for establishing regions are most vulnerable to climate change, and which stands guidelines of climate change adjusted seed transfer. Therefore, we could serve as sources of reproductive material preadapted to future addressed the following main questions using a genecological climates. approach: (1) Are current populations of spruce, fir, and beech in We used “relative risk of genetic maladaptation” to assess the Switzerland genetically maladapted to climate change? (2) Which vulnerability of trees to climate change. This index, which was origi- species and regions are most vulnerable to future maladaptation, and nally developed by Campbell (1986) to evaluate the genetic risk of why? We then discuss potential forest management practices
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