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Simulation Model Suggests That Fire Promotes Lodgepole Pine (Pinus Contorta) Invasion in Patagonia

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Simulation Model Suggests That Fire Promotes Lodgepole Pine (Pinus Contorta) Invasion in Patagonia Simulation model suggests that fire promotes lodgepole pine (Pinus contorta) invasion in Patagonia Authors: Kimberley T. Davis, Bruce D. Maxwell, Paul Caplat, Anibal Pauchard, and Martin A. Nunez The final publication is available at Springer via https://dx.doi.org/10.1007/s10530-019-01975-1. Davis, Kimberley T., Bruce D. Maxwell, Paul Caplat, Anibal Pauchard, Martin A. Nunez. "Simulation model suggests that fire promotes lodgepole pine (Pinus contorta) invasion in Patagonia." Biological Invasions 21, no. 7 (July 2019): 2287-2300. DOI:10.1007/ s10530-019-01975-1. Made available through Montana State University’s ScholarWorks scholarworks.montana.edu Simulation model suggests that fire promotes lodgepole pine (Pinus contorta) invasion in Patagonia Kimberley T. Davis, Bruce D. Maxwell, Paul Caplat, Anibal Pauchard, and Martin A. Nunez Abstract To best understand plant invasions and change. We used a spatially explicit invasion simula- predict unexpected outcomes it is necessary to inte- tion model modified to include fire to explore the grate information on disturbance, the local environ- implications of these complex interactions between ment, and demography. Disturbance by fire has been pine invasions and fire. We asked if fire promoted P. shown to promote invasions worldwide, but precise contorta invasion across a Patagonian steppe site and interactions between fire, native and invading species if this depended on the age of the invasion when it remain unclear. Indeed, trade-offs exist between fire- burned. Our simulations indicated that, although fire induced mortality of seed sources and increased was not necessary to initiate invasion, fire in commu- establishment, driving invasion outcomes. A positive nities where pine invasions were at least 10 years old feedback between lodgepole pine (Pinus contorta) resulted in increased spatial extent and maximum invasions and fire has been identified but only above a invasion density compared to unburned simulations. certain pine density. Above this threshold, fire resulted Fire through younger invasions did not alter the in increased pine dominance at the plot level, however progression of the invasion compared to unburned below this threshold establishment rates did not simulations. Pine invasions should be managed before they reach an advanced stage where positive feed- backs between fire and pine invasion could lead to Electronic supplementary material The online version of dramatic increases in invasion rate. this article (https://doi.org/10.1007/s10530-019-01975-1) con- tains supplementary material, which is available to authorized users. K. T. Davis (&) A. Pauchard Department of Ecosystem and Conservation Sciences, Laboratorio de Invasiones Biolo´gicas (LIB), Facultad de University of Montana, 32 Campus Dr., Missoula, Ciencias Forestales, Universidad de Concepcio´n, Casilla MT 59812, USA 160-C, Concepcio´n, Chile e-mail: [email protected] A. Pauchard B. D. Maxwell Institute of Ecology and Biodiversity (IEB), Las Palmeras Land Resources and Environmental Sciences Department, 3425, Casilla 653, Santiago, Chile Montana State University, Bozeman, MT 59717, USA M. A. Nun˜ez P. Caplat Grupo de Ecologı´a de Invasiones, INIBIOMA, Institute for Global Food Security, School of Biological CONICET, Universidad Nacional del Comahue, Quintral Sciences, Queen’s University Belfast, Belfast, 1250, CP 8400 San Carlos de Bariloche, Argentina Northern Ireland, UK Keywords Biological invasion Á Fire Á Pinus Á 1996). Several invasion simulation models have Positive feedback Á Simulation model Á Tree invasion incorporated disturbance into their simulations and found that the effects of disturbance on invasion depends on the vegetation type and the disturbance regime (e.g. Higgins and Richardson 1998; Pausas Introduction et al. 2006; Stevens and Beckage 2009; Shackelford et al. 2013). Such modeling efforts have also shown Plant invasions that have the capacity to alter fire that including feedbacks between fire and vegetation regimes and create positive feedbacks with fire have in models can lead to abrupt non-linear state changes the potential to cause significant impacts on native (Stevens and Beckage 2009). ecosystems (Mack and D’Antonio 1998; Brooks et al. Invasive pine species introduced to the Southern 2004; Gaertner et al. 2014). Although many studies Hemisphere are thought to create a positive feedback have examined how invasive plants alter fire regimes with fire whereby they alter some aspect of the fire (e.g. van Wilgen and Richardson 1985; Rossiter et al. regime which then promotes their own success over 2003; Pauchard et al. 2008; Balch et al. 2013), few native plants (Veblen et al. 2011). Recent work has have integrated information on invasion-fire feed- shown that one of the commonly invading pine backs to study the effects of these complex interac- species, Pinus contorta, alters fuel loads and structure tions on invasion extent. Integrating feedbacks and compared to uninvaded communities and that this population dynamics is necessary to determine how likely increases fire spread and severity (Co´bar- fire across a landscape will alter invasion spread and Carranza et al. 2014; Taylor et al. 2017; Paritsis consequently invasion impacts on native et al. 2018). The effect of disturbance on invasion communities. success varies when the likelihood of disturbance Disturbance is widely believed to increase invasion differs between uninvaded and invaded areas (Buckley success for many plant species by creating an invasion et al. 2007), as is the case for P. contorta invasions into window with increased resource availability (John- previously fuel-limited steppe systems (Veblen et al. stone 1986; Sher and Hyatt 1999; Davis et al. 2000). 2011; Paritsis et al. 2013). Pinus contorta experiences For pine species (genus Pinus) introduced to the high levels of fire-induced mortality (Baker 2009), and Southern Hemisphere, invasion is often correlated it recovers rapidly following fire in its native range in with human-caused or natural disturbances (Richard- western North America (e.g. Turner et al. 1997; Pierce son and Bond 1991; Richardson et al. 1994). In and Taylor 2011; Kemp et al. 2016). However, we particular, fire has been found to promote the invasion have found that in the introduced range, fire only of Pinus radiata, P. pinaster, and P. halepensis into promotes P. contorta establishment when the invasion South African fynbos (Richardson and Cowling 1992) density prior to the fire was high (Taylor et al. 2017). and P. halepensis into the Argentine Pampas grass- In other words, a positive feedback between pine lands (Zalba et al. 2008). Additionally, areas domi- invasions and fire is likely to form above an invasion nated by tall shrubs that are generally resistant to density threshold (Taylor et al. 2017). invasion by Pinus contorta (Taylor et al. 2016a) were Given the potential for P. contorta to alter fire significantly more vulnerable after fire (Taylor et al. behavior in invaded systems and to respond differently 2017). Other studies suggest that when disturbance is to fire depending on pre-fire invasion density, it is more likely in invaded than uninvaded areas it may unknown how fire will affect invasion success across a actually decrease the likelihood of further invasion, site. For example, although fire through a dense due to tradeoffs between increased invader habitat invasion will increase the quality of the seedbed and quality in disturbed areas and disturbance-induced the rate of pine establishment, it may also destroy a invader mortality (Buckley et al. 2007). Therefore, it is large part of the invasion seed source. Given the necessary to consider both the negative and positive threshold effect we found, we may expect that fires effects of disturbance on invasion when predicting through a young, low density invasion would reduce landscape-level changes in invasion as a result of post-fire invasion rates, whereas fire through an older, disturbance. Examining these interactions may be best high density invasion would increase invasion rates. achieved through simulation modeling (Higgins et al. Additionally, the invasion threshold at which a positive feedback forms with fire likely depends on the because P. contorta germinates best on bare mineral native vegetation. To explore the interactions between soil, is a prolific seed producer, exhibits high germi- P. contorta invasions and fire we created a spatially nation and establishment rates, and benefits from explicit invasion-fire simulation model. This study reduced competition with herbaceous species and aims to answer two questions through simulations with lower canopy cover following fire (Lotan and Critch- the invasion-fire model. First, will fire through an field 1990; Despain 2001; Ledgard 2001). Most P. invaded system promote further invasion and does this contorta seed germinates in the first year and seeds are depend on the age of the invasion when it burns? likely not viable in the seedbank beyond three to four Second, will changes in the invasion density threshold years (Ledgard 2001). that results in increased post-fire P. contorta estab- lishment alter invasion density and extent? Study site We hypothesized that fire early in an invasion would reduce P. contorta occupied cells (invasion Our model simulates a well-studied site in Coyhaique extent) and mean densities by killing invading trees Alto, Chile (CA) (Langdon et al. 2010; Taylor et al. but not promoting high post-fire establishment rates 2016a). CA is a Patagonian steppe site with a cold dry (due to low pre-fire invasion densities and reproduc- climate dominated by Festuca sp. and cushion plants tive output). We expected that fire through older such as Baccharis sp., Mullinum sp., and Acaena sp. invasions ([ 10 years, mainly reproductive trees) Pine plantations (P. contorta and P. ponderosa) were would result in higher invasion densities, but reduced planted in this area in the late 1980s and P. contorta spatial extent in the short term. but not P. ponderosa has invaded the surrounding native vegetation (Langdon et al. 2010; Taylor et al. 2016a). It is unclear which variety of P. contorta was Methods planted at this site, but we did not observe serotinous cones on any P.
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