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Ips Typographus and Dendroctonus Ponderosae Models Project Thermal Suitability for Intra- and Inter-Continental Establishment in a Changing Climate

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Ips Typographus and Dendroctonus Ponderosae Models Project Thermal Suitability for Intra- and Inter-Continental Establishment in a Changing Climate ORIGINAL RESEARCH published: 15 March 2019 doi: 10.3389/ffgc.2019.00001 Ips typographus and Dendroctonus ponderosae Models Project Thermal Suitability for Intra- and Inter-Continental Establishment in a Changing Climate Barbara J. Bentz 1*, Anna Maria Jönsson 2, Martin Schroeder 3, Aaron Weed 4, Renate Anna Irma Wilcke 5 and Karin Larsson 2 1 USDA Forest Service, Rocky Mountain Research Station, Logan, UT, United States, 2 Department of Physical Geography 3 Edited by: and Ecosystem Science, Lund University, Lund, Sweden, Department of Ecology, Swedish University of Agricultural 4 Sigrid Netherer, Sciences, Uppsala, Sweden, USDI National Park Service, Northeast Temperate Inventory and Monitoring Network, 5 University of Natural Resources and Woodstock, VT, United States, Rossby Centre, Swedish Meteorological and Hydrological Institute, Norrköping, Sweden Life Sciences Vienna, Austria Reviewed by: Climate change is altering legacies of native insect-caused disturbances and contributing Petr Doležal, to non-native invasions globally. Many insect fitness traits are temperature dependent Biology Centre (ASCR), Czechia Matthias Schmidt, and projected climatic changes are expected to cause continued alterations in Northwest German Forest Research insect-caused tree mortality, with uncertain consequences for forest ecosystems and Institute, Germany their management. Dendroctonus ponderosae in Pinus habitats of western North *Correspondence: Barbara J. Bentz America and Ips typographus in European Picea are among the most significant tree [email protected] mortality agents on each continent. Changing climate is influencing both species in their native habitats, although thermal suitability if they should invade new continents Specialty section: This article was submitted to and novel forest habitats has not been investigated. We assessed thermal suitability Forest Disturbance, for intra- and inter-continental establishment using physiological models that describe a section of the journal evolved, temperature-dependent traits of each species. Models were driven by Frontiers in Forests and Global Change projections from two Global Climate Models representing RCP 8.5. Simulations suggest Received: 07 November 2018 that for both species the common phenological strategy of one generation annually Accepted: 13 February 2019 (univoltine) will shift northward with warming throughout this century. As optimum habitat Published: 15 March 2019 for I. typographus univoltinism shifts northward, habitat supporting a 2nd generation, Citation: Bentz BJ, Jönsson AM, Schroeder M, a historically common strategy in warm European Picea forests, expands on both Weed A, Wilcke RAI and Larsson K continents. In contrast, a 2nd D. ponderosae generation has been historically rare due (2019) Ips typographus and to traits that evolved for phenological synchrony in its cool native habitats. As thermal Dendroctonus ponderosae Models Project Thermal Suitability for Intra- habitat for D. ponderosae univoltinism shifts northward, suitability for a 2nd generation is and Inter-Continental Establishment in limited to the warmest Pinus forests on both continents. In the near future (2011–2040), a Changing Climate. Front. For. Glob. Change 2:1. models project extensive thermal suitability for inter-continental establishment of both doi: 10.3389/ffgc.2019.00001 species, highlighting the need for effective mitigation policies and continued monitoring Frontiers in Forests and Global Change | www.frontiersin.org 1 March 2019 | Volume 2 | Article 1 Bentz et al. Bark Beetle Inter-Continental Establishment at ports in an era of climate change and increasing global trade. Throughout the century, thermal suitability remains high for I. typographus population success on both continents, for D. ponderosae in warm areas of Europe, and for D. ponderosae expansion into novel North American Pinus habitats. Portions of the historical D. ponderosae range, however, are projected to become thermally unsuitable. Keywords: climate change, voltinism, phenology, bivoltine, invasive, range expansion, mountain pine beetle, European spruce bark beetle INTRODUCTION during the period 1958–2001 and has increased in recent decades (Schelhaas et al., 2003; Seidl et al., 2011). Living forests are critical components of the earth’s climate Both species feed and reproduce in tree phloem, typically system. Growing trees absorb carbon dioxide (CO2) and can store killing the host when at epidemic levels, and they select mature it for hundreds to thousands of years. Increasing atmospheric trees within the appropriate genera that are of sufficient size and concentrations of CO2, largely caused by industrial activities, bark/phloem thickness for offspring production. Both species are contributing to global climate change (IPCC, 2013) and have relatively large geographic distributions that follow that forest carbon sequestration and storage have become important of their primary host trees, Picea abies for I. typographus mitigation strategies (Dugan et al., 2018; Xu et al., 2018). Multiple and Pinus species for D. ponderosae. Across their vast ranges, factors affect in situ forest carbon storage capacity including phenological plasticity in thermal- and photoperiod-regulated tree growth and decay rates, management, and tree mortality traits allow for adjustments in lifecycle, and hence generation due to deforestation and natural disturbances (Breshears and timing, depending on year-round available heat, day length, Allen, 2002 Li et al., 2003; Canadell and Raupach, 2008). and timing of cold temperatures that can induce offspring Insects are significant tree mortality agents globally, and bark death (Bentz et al., 2014; Schebeck et al., 2017; Schroeder and beetles (Coleoptera: Curculionidae, Scolytinae) are among the Dalin, 2017). Many of these physiological traits are directly most important in forest ecosystems (Hicke et al., 2013; Mezei influenced by climatic changes and can differ between the species et al., 2017). Bark beetles are integral ecosystem components (Bentz and Jönsson, 2015). Moreover, although both species that promote heterogeneity and resilience across multiple scales have adapted to feed and breed in living tree tissues, their (Kulakowski et al., 2017), yet can also impact economic, societal tolerance to active plant defenses and density dependent traits (Grégoire et al., 2015) and short-term carbon sequestration also differ (Kärvemo and Schroeder, 2010). (Pfeifer et al., 2011; Hansen et al., 2014) goals. The bark beetle The distribution of I. typographus follows the entirety of and forest carbon feedback to climate change highlights the its host tree Picea abies which has a continuous range in importance of understanding future climate influences on global Scandinavia, north-eastern Europe and western Russia, and at distributions of bark beetle populations (Kurz et al., 2008; higher latitudes in central Europe (Farjon and Filer, 2013). Raffa et al., 2008; Adams et al., 2010). Climatic changes have The current distribution of D. ponderosae, in contrast, is not already influenced patterns and timing of bark beetle population as extensive as species within its host genus Pinus, which are outbreaks in their native ecosystems (Buotte et al., 2016; Senf contiguous further north and east in Canada, further east in the and Seidl, 2018), in addition to intra- and inter-continental range US, and further south in Mexico than the current distribution of expansion and invasion of novel habitats by some species (Liu D. ponderosae. Canada was covered in ice 21,000 years BP, and as et al., 2014; Cooke and Carroll, 2017). Predicting tree-killing glaciers retreated several Pinus species colonized the new habitat bark beetle population responses, including range expansion (Godbout et al., 2008; Roberts and Hamann, 2015). Despite the and potential contraction, to future climate projections is availability of Pinus host trees, the northernmost areas remained an important component of forest management and carbon thermally unsuitable for D. ponderosae (Safranyik, 1978). Recent sequestration strategies globally. warming in western Canada resulted in a rapid northward The European Ips typographus (L.) and North American (NA) expansion of D. ponderosae into Pinus habitat of northern Dendroctonus ponderosae (Hopkins) are considered among the British Columbia and western Alberta (de le Giroday et al., most important disturbances affecting tree mortality on each 2012). Climate change-induced range expansion of D. ponderosae continent. Due to thermally-dependent fitness traits and positive southward has not occurred, despite the availability of known responses to drought-induced tree stress, population activity has Pinus hosts. Multiple Pinus species are found at low elevations increased recently with a changing climate (Weed et al., 2015; in Arizona, Mexico, and Central America, yet the most Kolb et al., 2016; Marini et al., 2017; Senf and Seidl, 2018). In southern locations where D. ponderosae has been observed western Canada and the United States (US) recent tree mortality is high elevation forests of southern Arizona, US, northern due to D. ponderosae exceeded 28 Mha (Hicke et al., 2015; Cooke Chihuahua, Mexico, and northern Baja California Norte, Mexico and Carroll, 2017). In Europe tree mortality caused by bark (Armendáriz-Toledano et al., 2017; Bentz and Hansen, 2017; beetles, mainly I. typographus, exceeded 2.88 million m3 per year Dowle et al., 2017). Frontiers in Forests and Global Change | www.frontiersin.org 2 March 2019
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