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View and Print the Publication Ecosystems https://doi.org/10.1007/s10021-020-00548-x Ó 2020 Springer Science+Business Media, LLC, part of Springer Nature Strong Legacy Effects of Prior Burn Severity on Forest Resilience to a High-Severity Fire Lucas B. Harris,1* Stacy A. Drury,2 and Alan H. Taylor3 1Department of Geography, The Pennsylvania State University, 302 Walker Building, University Park, Pennsylvania 16802, USA; 2USDA Forest Service, Pacific Southwest Research Station, Davis, California, USA; 3Department of Geography, Earth and Environ- mental Systems Institute, The Pennsylvania State University, 302 Walker Building, University Park, Pennsylvania 16802, USA ABSTRACT Legacy effects from one disturbance may influence have affected soils, litter cover and shade after the successional pathways by amplifying or buffering 2012 fire and therefore affected seedling establish- forest regeneration after the next disturbance. We ment and survival. A canopy seed bank of unburnt assessed vegetation and tree regeneration in non- cones from trees killed by the 2012 fire potentially serotinous Sierra lodgepole pine (Pinus contorta var. contributed to a strong effect of prior burn severity murrayana) stands after a 1984 wildfire which on regeneration after the 2012 fire despite a lack of burned with variable severity and again after a serotinous or resprouting tree species, although the high-severity subsequent fire in 2012. The legacy influence of this canopy seedbank was likely lim- effects of the 1984 fire were amplified; seedlings ited to the year following the fire. Our results and saplings were abundant in areas initially suggest that a low- to moderate-severity fire in- burned at low severity (1267 stems ha-1) despite creases forest resilience relative to a high-severity high reburn severity, but regeneration was low in fire even when the next fire burns at high severity. areas twice burned at high severity (31 stems ha- 1). Our results suggest that the severity of the 1984 Key words: Repeated fires; Tree regeneration; fire may have influenced post-2012 tree regenera- Legacy effects; Forest resilience; High-severity tion by creating variable fuel loading, which may wildfire; Pinus contorta. HIGHLIGHTS Forests twice burned at high severity had sparse tree regeneration Low-severity fire increased forest resilience to a Severity of a prior fire determined tree regener- subsequent high-severity fire ation after a high-severity reburn INTRODUCTION Received 6 March 2020; accepted 15 August 2020 Successive disturbances, especially when occurring at relatively short intervals, may lead to persistent Electronic supplementary material: The online version of this article (https://doi.org/10.1007/s10021-020-00548-x) contains supplementary and sometimes surprising changes in vegetation material, which is available to authorized users. (Zedler and others 1983; Paine and others 1998; Author Contributions LBH and AHT collected data, LBH performed Buma 2015). Such changes arise in part due to analysis and wrote the first draft of the manuscript, and all authors de- signed the study and contributed substantially to revisions. legacy effects of past disturbances on vegetation *Corresponding author; e-mail: [email protected] structure and composition, which interact with L. B. Harris and others subsequent disturbance to influence vegetation Field-based studies using repeated measurements recovery (Paine and others 1998; Franklin and to quantify vegetation change over two successive others 2000). As climate change leads to increased wildfires (for example, Coop et al., 2016) are scarce fuel aridity and area burned in western US forests but provide key insights on interactions between (Abatzoglou and Williams 2016), a key question is repeated fires, vegetation and potential forest how legacy effects of past disturbances influence recovery. forest recovery after a high-severity fire. Through- In this study, we measured post-fire tree out, we use the term ‘‘burn severity’’ and descrip- regeneration in stands dominated by Sierra tors such as ‘‘high severity’’ to refer to ‘‘vegetation lodgepole pine (Pinus contorta var. murrayana) burn severity’’ meaning the effects of fire on veg- following repeated fires in the southern Cascade etation measured one or more years after the fire Range, California, USA. Sierra lodgepole pine is a (sensu. Morgan and others 2014). widespread subspecies, ranging from Baja Cali- Work in dry pine and mixed-conifer forests of fornia, Mexico, to central Oregon, USA (Critch- the western USA suggest that repeated fires (also field 1980). We conducted this work in low-lying called reburns) reinforce vegetation changes trig- basins representative of those occurring through- gered by past fires, especially when one high- out the southern Cascade Range, in which cold air severity fire leads to vegetation types and structures pooling and nutrient-poor pumice soils favor which are prone to burn at high severity in the lodgepole pine dominance (Youngberg and Dyr- next fire (Odion and others 2010; Coop and others ness 1965; Cochran and Berntsen 1973;Volland 2016; Lauvaux and others 2016). Repeated fires 1985). Vegetation and post-fire tree regeneration have the potential to drive persistent vegetation were assessed in 2008 following a 1984 wildfire change even in more mesic, higher-elevation that burned at a range of fire severities, and then mixed-conifer and subalpine forests by dealing a again in 2019 following a 2012 wildfire that double blow: a further reduction in live trees to act burned mainly at high severity. Building on an as seed sources and mortality of much of the tree earlier analysis of tree regeneration at this site by regeneration (that is, seedlings and saplings) that Pierce and Taylor (2011), we assessed the influ- established after the previous fire (Gray and ences of burn severity, stand structure and mi- Franklin 1997; Busby 2019; Turner and others crosite characteristics (for example, cover of 2019). shrubs, logs and rocks) on tree seedling and sap- The characteristics of repeated fires may affect ling density following the 2012 fire. Through these tree regeneration and therefore future forest cover. analyses, we evaluated the legacy effects of the Prior research suggests that repeated high-severity 1984 fire on subsequent forest change and whe- leads to lower tree seedling density compared with ther the second, higher-severity 2012 fire dimin- a low-severity fire followed by a high-severity fire, ished, reinforced or amplified these effects. We because a past high-severity fire may more com- hypothesized that tree seedling and sapling den- pletely eliminate the seedbank and seed sources sity would be strongly correlated with density of (Stevens-Rumann and Morgan 2016). Such results and distance to cone-bearing trees following the suggest that legacy effects from a past fire can 2012 fire, because Sierra lodgepole pine is not influence forest resilience to the next fire (John- serotinous and therefore has to seed in from stone and others 2016). Here we define forest re- nearby surviving trees following a fire (Dahms silience as the degree of perturbation a forest can 1963; Pierce and Taylor 2011). Consequently, we withstand before transitioning to a new state expected the second, predominantly high-severity (Holling 1973; Gunderson 2000), and legacies as 2012 fire to diminish any legacy effect of the 1984 the organisms and biologically derived patterns left fire because of the large reduction in density of after a disturbance which influence future ecosys- surviving seed trees. tem dynamics (Franklin and others 2000). A growing body of research has examined suc- METHODS cessional pathways over successive fires (Donato and others 2009; Coop and others 2016; Stevens- Study Area Rumann and Morgan 2016; Turner and others This study was conducted within Lassen Volcanic 2019). However, most studies on repeated fire ef- National Park (LVNP), California. As reviewed by fects on forest change rely on space-for-time sub- Parker (1993), Sierra lodgepole pine is common at stitution (Pickett 1989) where nearby sites with 1800–2200 m elevation in LVNP and occurs in different fire histories are surveyed and results are three primary settings: recently disturbed areas, assumed to represent a time series at one location. Strong Legacy Effects of Prior Burn Severity on Forest Resilience to a High-Severity Fire mixed with other tree species in uplands and as the vals than surrounding mixed-conifer forest because dominant tree in lowlands. Sierra lodgepole pine is they are less productive and slower to recover from highly tolerant of wet and cold conditions and disturbance and therefore tend to be more fuel- therefore often dominates low-lying areas experi- limited (Merschel and others 2018). These fire encing cold air drainage and wet soils within LVNP history data indicate the 28-year interval between (Taylor 1990; Parker 1993). fires in the present study is relatively short but In 1984 the Badger Fire burned 563 ha of LVNP within the historical range of variability for this and the adjacent Lassen National Forest. This area forest type. burned again in the 2012 Reading Fire (10,876 ha). In 2008 Pierce and Taylor (2011) assessed vegeta- Field Data tion change caused by the Badger Fire within three Patches that burned at low, moderate and high lodgepole pine-dominated study sites (40.552° N, severity in the 1984 Badger Fire had been selected 121.404° W) selected to be homogenous with re- for study using 2005 aerial photographs as part of spect to terrain and soils so that the influence of an earlier analysis by Pierce and Taylor (2011), and burn severity on tree regeneration could be iso- burn severity in these burn patches was verified in lated. These sites were in flat, low-lying basins in the field. Low-, moderate- and high-severity fire which cold air pooling and potentially coarse or was defined as areas with less than 25% tree ca- low-nutrient substrate often exclude trees species nopy cover loss, 25–75% loss and 75% or more other than lodgepole pine (Cochran and others loss, respectively.
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