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Wildfire Severity and Postfire Salvage Harvest Effects on Long‐Term Forest Wildfire severity and postfire salvage harvest effects on long-term forest regeneration 1,2, 3 4 1,4 NICHOLAS A. POVAK , DEREK J. CHURCHILL, C. ALINA CANSLER, PAUL F. HESSBURG , 4 4 5 6 VAN R. KANE, JONATHAN T. KANE, JAMES A. LUTZ , AND ANDREW J. LARSON 1USDA-Forest Service, Pacific Northwest Research Station, 1133 N Western Avenue, Wenatchee, Washington 98801-1229 USA 2Oak Ridge Institute for Science and Education (ORISE), Oak Ridge, Tennessee 37830 USA 3Washington State Department of Natural Resources, Forest Health and Resiliency Division, Olympia, Washington 98504 USA 4School of Environmental and Forest Sciences, University of Washington, Box 352100, Seattle, Washington 98195 USA 5Quinney College of Natural Resources & Ecology Center, Utah State University, Logan, Utah 84322 USA 6W.A. Franke College of Forestry & Conservation, University of Montana, Missoula, Montana 59812 USA Citation: Povak, N. A., D. J. Churchill, C. A. Cansler, P. F. Hessburg, V. R. Kane, J. T. Kane, J. A. Lutz, and A. J. Larson. 2020. Wildfire severity and postfire salvage harvest effects on long-term forest regeneration. Ecosphere 11(8):e03199. 10. 1002/ecs2.3199 Abstract. Following a wildfire, regeneration to forest can take decades to centuries and is no longer assured in many western U.S. environments given escalating wildfire severity and warming trends. After large fire years, managers prioritize where to allocate scarce planting resources, often with limited informa- tion on the factors that drive successful forest establishment. Where occurring, long-term effects of postfire salvage operations can increase uncertainty of establishment. Here, we collected field data on postfire regeneration patterns within 13- to 28-yr-old burned patches in eastern Washington State. Across 248 plots, we sampled tree stems <4 m height using a factorial design that considered (1) fire severity, moderate vs. high severity; (2) salvage harvesting, salvaged vs. no management; and (3) potential vegetation type (PVT), sample resides in a dry, moist, or cold mixed-conifer forest environment. We found that regeneration was abundant throughout the study region, with a median of 4414 (IQR 19,618) stems/ha across all plots. Only 15% of plots fell below minimum timber production stocking standards (350 trees/ha), and <2% of plots were unstocked. Densities were generally highest in high-severity patches and following salvage harvest- ing, although high variability among plots and across sites led to variable significance for these factors. Post hoc analyses suggested that mild postfire weather conditions may have reduced water stress on tree establishment and early growth, contributing to overall high stem densities. Douglas fir was the most abundant species, particularly in moderate-severity patches, followed by ponderosa pine, lodgepole pine, western larch, and Engelmann spruce. Generalized additive models (GAMs) revealed species-level climatic tolerances and seed dispersal limits that portend future challenges to regeneration with expected future cli- mate warming and increased fire activity. Postfire regeneration will occur on sites with adequate seed sources within their climatic tolerances. Key words: climatic tolerance; Douglas fir; dry forest; high severity; lodgepole pine; ponderosa pine; regeneration; resilience; salvage harvest; seed dispersal; western larch; wildfire. Received 3 February 2020; revised 17 April 2020; accepted 28 April 2020; final version received 4 June 2020. Corresponding Editor: Carrie R. Levine. Copyright: © 2020 The Authors. This is an open access article under the terms of the Creative Commons Attribution License, which permits use, distribution and reproduction in any medium, provided the original work is properly cited. E-mail: [email protected] v www.esajournals.org 1 August 2020 v Volume 11(8) v Article e03199 POVAK ET AL. INTRODUCTION vegetation in the Klamath Mountains due to their high flammability and the inability of coni- Wildfires are integral in shaping ecosystems fer seedlings to survive recurrent severe fires across the western United States, and large wild- (Odion et al. 2010). While early seral patches are fire years have increased markedly in recent important components of western landscapes years (Stephens 2005, Miller et al. 2009, Wester- (Swanson et al. 2011), large contiguous patches ling 2016), along with the size of stand-replacing can negatively influence plant and animal habitat (syn. high-severity) fire patches (O’Connor et al. and dispersal networks, distribution of future 2014, Stevens et al. 2017). Given the large extent fire severity patch sizes, and forest regeneration of the landscape currently recovering from fire, (Hessburg et al. 2015, 2019). Where these patches much uncertainty exists regarding the develop- occur following uncharacteristically large and ment of future forest canopies, particularly after severe wildfires, accelerating tree establishment moderate- and high-severity fires. Of special con- and regrowth of conifers may be desirable for cern are forests that were historically dominated habitat restoration, erosion control, water quality, by low- and mixed-severity fires; ecosystems that carbon sequestration, or other ecosystem ser- are less well-adapted to the effects of large-scale vices. and severe wildfires (Perry et al. 2011, Hessburg Postfire recovery is influenced by biotic and et al. 1999, 2016). abiotic factors operating across multiple spatial Seedling establishment delays result from and temporal scales. Recovery is contingent lengthy dispersal gradients within large high- upon local viable seed source, favorable seedbed severity patches and the effects of these long dis- conditions, limited interspecific competition, tances on conifer species that rely on wind or ani- favorable long-term climate, and near-term post- mal seed dissemination. Replacing conifers in fire weather conditions (Larson and Franklin, these areas are species well-adapted to fire, with 2005, Harvey et al. 2016, Kemp et al. 2016). Broad the ability to resprout (Lawes and Clarke 2011, environmental gradients of western U.S. moun- Pausas et al. 2016), or that depend on fire to ger- tain ranges can significantly influence forest minate seeds stored in soils (Schwilk and Ackerly regeneration, with moist and cool environments 2001, Keeley et al. 2011). In the western United favoring abundant regeneration, while arid con- States, few coniferous species regenerate through ditions present challenges for seedling establish- adventitious sprouting or by serotinous cones, ment and growth (Dodson and Root 2013, suggesting that hardwood trees, shrubs, and Savage et al. 2013, Tepley et al. 2017). For exam- other nonforest species may be favored in large ple, Dodson and Root (2013) showed significant stand-replacement patches (Lydersen and North increases in postfire regeneration density along a 2012, Tepley et al. 2017). This, in turn, may lead broad elevational gradient in eastern Oregon; to shifts in dominant species or physiognomic tree establishment at low elevations was unlikely conditions (Romme et al. 1998, Odion et al. 2010) due to elevated evaporative demands. However, that can last for decades to centuries. within environments, local variations in topogra- Regeneration failures are documented where phy and edaphic properties can create favorable postfire conditions were unfavorable to tree microsites for regeneration (Donato et al. 2009). establishment and survival (Coop et al. 2016, Ste- Similarly, periods of benign weather following vens-Rumann et al. 2018), and studies suggest fire can provide opportunities for trees to regen- that systems can persist in alternative nonforest erate (Littlefield 2019). However, long-term states (Odion et al. 2010) or landscape traps (Lin- regeneration success is not guaranteed, particu- denmayer et al. 2011). Where nonforest commu- larly where trees develop at the edge of their nities (e.g., grasslands, shrublands, and physiological tolerance (Parks et al. 2019). Under- savannahs) establish after disturbance, their standing the conditions conducive to regenera- dominance can self-regulate through feedbacks tion and factors that limit long-term success is with future reburns to the detriment of conifer key to postfire management, especially as the cli- establishment (Odion et al. 2010, Coop et al. mate changes (Tepley et al. 2017). 2016, Coppoletta et al. 2016). For example, short Management can foster the development of fire-return intervals favored shrubland forest vegetation through planting, mechanical v www.esajournals.org 2 August 2020 v Volume 11(8) v Article e03199 POVAK ET AL. removals, prescribed burning, and soil scarifica- Forest regeneration is a long-term process that tion to expose mineral soil (Shive et al. 2013, Ste- depends on post-disturbance climate and vens et al. 2014). However, current trends in weather conditions, the variability in physiologi- burned area are outpacing the ability of some cal traits and life history strategies of regenerat- managers to mitigate the worst impacts of severe ing species, and on the environmental context for wildfires (North et al. 2019). After large fire regeneration (Coop and Schoettle 2009, Coop years, managers prioritize the allocation of scarce et al. 2010, Donato et al. 2016, Malone et al. 2018, resources, often with limited information on fac- Stevens-Rumann et al. 2018). Inter- and tors that drive forest establishment (Calkin et al. intraspecific competition and facilitation can 2005). There is therefore a critical need to docu- affect establishment and growth
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