Kristin Zouhar Jane Kapler Smith Steve Sutherland Chapter 2: Effects of Fire on Nonnative Invasive Plants and Invasibility of Wildland Ecosystems Considerable experimental and theoretical work has resource availability. In the context of invasion, this been done on general concepts regarding nonnative is the availability of resources needed by a nonnative species and disturbance, but experimental research species to establish, persist, and spread. Morphological on the effects of fire on nonnative invasive species is properties, phenological properties, and competitive sparse. We begin this chapter by connecting fundamen- ability of native species influence resistance to invasion, tal concepts from the literature of invasion ecology to while the same properties of nonnative species influ- fire. Then we examine fire behavior characteristics, ence potential to invade. Native and nonnative plant immediate fire effects, and fire regime attributes in responses to fire, such as damage or stimulation from relation to invasion potential. These concepts form heat and increases or decreases in postfire years, are the basis for examining the literature that supports particularly important for our discussion. Propagule or refutes several common generalizations regarding pressure is the availability, abundance, and mobility fire effects on nonnative invasives. We conclude with of propagules in and around a plant community. a summary of management implications regarding In this chapter, we examine several generalizations fire effects on nonnative invasive plants. that have been suggested about wildland invasion by nonnative species after fire. We treat these general- izations as questions that can be examined in light of Invasion Ecology ________________ current research. Invasion ecology is influenced by interactions of • Question 1. Does fire generally favor nonnative ecosystem properties, properties of native and non- over native species? native plant species, and nonnative propagule pres- • Question 2. Do invasions increase with increas- sure (Lonsdale 1999) (fig. 2-1). Ecosystem properties ing fire severity? include disturbance regimes and fluctuations in USDA Forest Service Gen. Tech. Rep. RMRS-GTR-42-vol. 6. 2008 7 Figure 2-1—Susceptibility of a plant community to invasion by nonnative species after fire depends on properties of the ecosystem itself, properties of plant populations (both native and nonnative) and availability of nonnative plant propagules (following Lonsdale 1999). • Question 3. Does additional disturbance ­(before, their historic fire regime (Keeley 2001; Keeley and during, or after fire) favor invasions? others 2003). Furthermore, divergence from the • Question 4. Do invasions become less severe historic fire regime in the “opposite” direction—with with increasing time after fire? reduced fire frequency or severity—may also increase • Question 5. Do invasions increase with disrup- invasibility. For invasion to actually occur, community tion of the presettlement fire regime? susceptibility and resources adequate for the spread • Question 6. Are postfire invasions less common of nonnative species must coincide with availability of in high elevation ecosystems? propagules of nonnative species that can successfully We will return to these questions after reviewing the compete with native vegetation for those resources connections between invasion concepts and fire. (Davis and others 2000). Ecosystem Properties and Resource Properties of Native and Nonnative Plants Availability Several plant characteristics influence their suscep- tibility to fire injury, ability to recover and compete Invading species must have access to resources, for resources following fire, and changes in cover and including light, nutrients, and water, so community dominance over time after fire. Fire has less potential susceptibility to invasion can be explained to some to kill individuals and impact populations if the species’ extent by changes in resource availability. A species meristem tissues, buds, and seeds are protected from will “enjoy greater success in invading a community heat-caused damage. Avoidance of heat damage can be if it does not encounter intense competition for these based on structural features, location of meristematic resources from resident species” (Davis and others tissues, or phenology (see “Influence of Fire Season 2000). Therefore, a plant community becomes more and Plant Phenology on Postfire Invasions” page 18). susceptible to invasion when the amount of unused Elevated buds, thick bark, and underground vegeta- resources increases. Fire can increase resource avail- tive structures can provide protection. ability by reducing resource use by resident vegetation Fire survivors and species that form persistent (through mortality or injury) or by altering the form and seed banks, native or nonnative, have early access to availability of nutrients. Reports of postfire increases resources on a burned site (see “Influence of Fire on in nonnative species, often attributed to increased light Resource Availability and Interactions Between Plant or other resources, are available in the literature (for Species” page 10). They can spread by regenerating example, D’Antonio 2000, review; Hunter and others from surviving structures or establishing from the seed 2006; Keeley and others 2003). bank, then producing abundant seeds that establish Disturbed areas are often considered vulnerable to on the exposed mineral soil seedbed (see “Downward invasion (Sakai and others 2001), and burned areas are heat pulse effects on plant survival” page 14). Many no exception. However, some plant communities that nonnative invasives are annuals or biennials with have evolved with recurring fire, such as California short generations, ability to self pollinate, and low chaparral, are not considered highly invasible under shade tolerance (Sutherland 2004). A review by 8 USDA Forest Service Gen. Tech. Rep. RMRS-GTR-42-vol. 6. 2008 Barrett (2000) highlights the relationship between and abundance. Where burns are associated with an- disturbance and opportunistic or invasive species with thropogenic disturbance, they are likely to be subject short life cycles, well-developed dispersal powers, and to greater propagule pressure and may therefore be high reproductive output. These typically ephemeral more susceptible to postfire invasion than burns in less species can establish on burned sites only if abundant disturbed areas (see “Question 3” page 22). The gener- propagules are available from the soil bank or nearby ally positive relationship between nonnative invasive unburned areas. Unless they alter ecosystem processes species and anthropogenic disturbance (for example, to perpetuate early-seral conditions, ephemerals are Dark 2004; Johnson and others 2006; McKinney 2002; often replaced by perennials within a few years after Moffat and others 2004) may have implications for fire (see “Question 4” page 25). plant communities and bioregions that currently show relatively little effect of fire on nonnative plant inva- Nonnative Propagule Pressure sions. Fire appears only weakly related to spread of nonnative species in the Northeastern bioregion (chap- The spatial distribution of nonnative source popula- ter 5), but the plethora of nonnative invasive species tions and their mode of propagule dispersal influence present in this region (Mehrhoff and others 2003) and their establishment and spread in new areas (Amor the prevalence of anthropogenic disturbance suggest and Stevens 1975; Giessow and Zedler 1996; Wiser that, if burning increases, impacts from nonnative and others 1998), including burns (Keeley and others species may increase as well. Similarly, while fire- 2003). Some seeds are heat tolerant and therefore may caused increases in nonnative invaders are currently survive a fire onsite (Volland and Dell 1981). Postfire uncommon in Alaska (chapter 10), expanding human establishment and spread of nonnative species depends, influences on wildlands coupled with climate change in part, on propagule pressure (sensu Colautti and may increase problems with nonnative plants in that others 2006; Drake and Lodge 2006; Lockwood and state. Similar concerns have been voiced regarding in- others 2005)— the abundance of nonnative propagules vasive species in Colorado shortgrass steppe (Kotanen occurring onsite and within dispersal distance of the and others 1998) and may apply in many areas of the burned area. D’Antonio and others (2001b) contend United States. that variation in propagule supply interacts with the “ecological resistance” of an ecosystem (sensu Elton 1958) such that when resistance is low, few propagules Influence of Fire on Invasions _____ are needed for successful invasion, and as resistance The responses of plants to fire depend on both fire increases it takes proportionately more propagules for attributes and plant attributes relating to survival invaders to establish. In a meta-analysis designed to ex- and establishment (Pyke and others, in review). Non- amine characteristics of invasiveness and invasibility, native plants that survive on site, establish from the Colautti and others (2006) found that while propagule seed bank, or disperse seed into burns soon after fire pressure was rarely considered in studies of biological have early access to resources that are more plenti- invasions, it was a significant predictor of invasibility. ful or more available after fire. Fire behavior char- More disturbance and higher resource availability