The influence of prescribed fire on wild turkeys in the Southeastern United States: A review and synthesis

Forest Ecology and Management 455 (2020) 117661

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Review and synthesis

The influence of prescribed fire on wild turkeys in the Southeastern United States: A review and synthesis

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Gregory T. Wanna, , James A. Martina, Michael J. Chamberlaina a

Warnell School of Forestry and Natural Resources, University of Georgia, 180 E. Green St., Athens, GA 30602, United States

A R T I C LE I N FO

A B S T R A C T

Keywords: Habitat selection Meleagris gallopavo Movement Pine-grassland Prescribed fire Return interval Survival

The pine-grassland ecosystems once prevalent in the Southeastern United States were dependent on frequent fire events to maintain plant communities and avoid succession to hardwood and shrub-dominant communities. The use of prescribed fire has replaced naturally occurring fires produced from lightning strikes to maintain remaining pine-grassland systems, and to expand and promote restoration into reclaimed areas. Currently, prescribed fire is a widely accepted management tool promoted by both state and federal wildlife and land-management agencies, and is assumed to be beneficial for both game and non-game species. However, a comprehensive set of guidelines related to use of prescribed fire for promotion of wildlife is lacking for most species, including species whose dependence on fire is presumed to be critical. We reviewed available literature on prescribed fire and its influence on wild turkeys (Meleagris gallopavo) in the Southeastern distribution of its range. We reviewed extant literature relative to historical use of prescribed fire for upland gamebird management, and focused on documented effects of fire on life-history characteristics of wild turkeys, including habitat selection, demography, and movement. The literature supported preferential use of areas burned in the previous 3 years, with avoidance of areas lacking a recent fire history. Fire return intervals between 2 and 3 years were generally supported in the literature as optimal to reduce woody shrub encroachment and maintain an herbaceous understory. However, areas infrequently burned, such as mature hardwood forests, provide important habitat during fall and winter, and provide important roosting sites. Contrary to misperceptions among the public, growing-season fire appears to pose little direct risk to wild turkey nests and poults, but research on this topic is limited and only recently initiated. We lack a collective knowledge of the most appropriate spatial scale and extent of prescribed fires for wild turkeys, and a single set of values for these metrics that can be applied throughout the Southeastern U.S. is likely to be unrealistic given variations in local plant communities and landscape composition. Non-target species should be carefully considered before implementing prescribed fire regimes targeted specifically towards wild turkeys, as such fire regimes may not be optimal for other species.

1. Introduction Naturally occurring fire is responsible forshaping vegetative communities in many ecosystems throughout North America, including the Southeastern pine-grassland systems where fire frequency has been altered through fire suppression and landscape changes (Waldrop et al., 1992; Glitzenstein et al., 1995, 2003; Van Lear et al., 2005). Once ecosystems previously maintained by wildfires are lost, rehabilitating them to historic conditions requires implementation of management plans that use prescribed fire and consistent fire events to maintain desired plant communities (Waldrop and Goodrick, 2012). Prescribed fire has been used as a primary management tool to benefit many avian species endemic to ecosystems historically maintained by naturally

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occurring fires (Saab et al., 2005; Russell et al., 2009), including redcockaded woodpecker (Leuconotopicus borealis) and Bachman’s sparrow (Peucaea aestivalis) in pine-grasslands of the Southeast (James et al., 1997; Plentovich et al., 1998; Russell et al., 2009) and prairie chickens (Tympanuchus spp.) in the Great Plains (Kirsch, 1974). Prescribed fires in the Southeastern U.S. have also been used to promote conditions favorable for game species such as northern bobwhite (Colinus virginianus; e.g., McGrath et al., 2017), white-tailed deer (Odocoileus virginianus; Masters et al., 1993), and wild turkey (Meleagris gallopavo; Little et al., 2014). Prescribed fire is now a primary tool used by natural resource managers, and is increasingly used on state and federal lands (Waldrop and Goodrick, 2012). Nonetheless, challenges remain in implementing prescribed fire due to negative perceptions about fire, and

Corresponding author. E-mail address: [email protected] (G.T. Wann).

https://doi.org/10.1016/j.foreco.2019.117661 Received 22 July 2019; Received in revised form 12 September 2019; Accepted 30 September 2019 0378-1127/ © 2019 Elsevier B.V. All rights reserved.

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because of logistical challenges of conducting prescribed fires at large spatial extents necessary to manage pine-grassland ecosystems (Brennan et al., 1998). Furthermore, direct effects (e.g., nest mortality) of prescribed fire on wildlife are still poorly understood, necessitating work to synthesize existing studies on effects of prescribed fire on wildlife. Fortunately, substantive research focusing on responses of wildlife to prescribed fire has accumulated over the past 3 decades (e.g., Fontaine and Kennedy, 2012). Hence, reviewing and synthesizing extant literature is beneficial to inform wildlife management agencies charged with using prescribed fire to manage forest and grassland communities. We focus on the wild turkey (Meleagris gallopavo; hereafter turkey), a species whose range in the Southeastern U.S. is often within fire-managed ecosystems on public lands. We restrict our inference to the fire ecology of turkeys and pine-grassland dominant systems in the Southeastern U.S., because little information exists outside of this region, and because prescribed fire is a common approach used to manage vegetation communities in pine forests of the Southeast (Johnson and Hale, 2002). The wild turkey is the largest galliform in North America and has a distribution ranging from southern Canada to southern Mexico (McRoberts et al., 2014). Turkeys inhabit a wide range of cover types in the Southeastern portion of their range, and habitat use varies seasonally (Miller and Conner, 2007). In general, both forests and treeless cover types are important determinants of turkey space use (Porter, 1992). Turkey populations were threatened with extirpations by the early 20th century resulting from over harvesting and habitat loss. Their historic distribution has since been restored due to restocking and harvest regulations by state and federal agencies (Kennamer et al., 2001). However, since 2000, populations have exhibited declines in both abundance (Eriksen et al., 2015) and productivity (Byrne et al., 2015). State wildlife agencies throughout the U.S. are actively trying to ensure sustainable populations of turkeys, given the economic and social importance of turkeys as a game species (Southwick Associates, Inc. 2003). Therefore, management strategies that can enhance turkey populations by improving habitat conditions are actively pursued by wildlife managers. In the Southeast, pine-grassland ecosystems are critical in supporting turkeys and other wildlife species. Historically, longleaf pines (Pinus palustris P. Miller) were once dominant in savanna and forest communities throughout the Southeastern U.S. (Bridges and Orzell, 1989; Peet and Allard, 1993). Today, < 3% of presettlement old-growth pine habitat remains (Frost, 1993), due primarily to fire suppression (Brockway and Lewis, 1997). The influence of fire on turkeys has been of interest since the 1930s. Herbert Stoddard wrote frequently on using fire to manage northern bobwhite (Way, 2008), and Stoddard’s career spanned a period when the U.S. Forest Service was promoting and practicing complete fire suppression (Way, 2006). Stoddard viewed the use of prescribed fire as essential to maintain productive habitats for game by promoting early successional plant communities conducive for foraging, nesting, and brooding (Stoddard, 1935, 1963). He also recognized that areas lacking frequent fire events quickly developed closed canopies and understories too dense to be used by turkeys (Stoddard, 1963). By the 1970s, use of fire as a management tool for turkeys began being reported by biologists (e.g., Lewis and Harshbarger, 1976). Previously, effects of fire on turkey habitat were only mentioned in passing in the literature and were rarely the primary focus of scientific articles. Herein, we focus on the influence of prescribed fire on turkeys to synthesize the extant literature and identify areas where lack of knowledge is prohibitive to making informed management recommendations. Our objective is to provide state and federal agencies charged with managing turkeys on landscapes with prescribed fire an assessment of how fire influences turkey populations, and identify key uncertainties in best management practices. We have organized the paper into 4 sections. First, we describe fundamental concepts in fire ecology as they relate to land management practices. We discuss dormant and growing season fires, fire return intervals, and spatial extent

of burns. Second, we synthesize extant literature detailing influences of prescribed fire on 3 components of turkey life-history, including habitat use, demographic responses, and movement. Third, we discuss information needs to guide and motivate future research evaluating how prescribed fire influences turkey populations. We offer suggestions relative to reporting results in a consistent manner to ease comparisons among studies. Fourth, we provide practical guidance on managing pine-grassland habitats for turkeys given what is known from extant literature. 2. Fire management Turkeys were historically abundant throughout the Southeastern U.S. (Mosby and Handley, 1943; Kennamer et al., 2001), occupying ecosystems largely composed of fire adapted forests and grasslands (Frost, 1993). Hence, turkeys are presumably well adapted to regularly occurring fires in pine-grassland habitats. A common goal among state agencies is to create and manage historic pine-grassland conditions, which relies on understanding fire ecology. Currently, no region-wide statistics have been published on the extent to which prescribed fire is applied across the landscape. In the 13 states that comprise the Southeastern U.S., the U.S. Forest Service classifies nearly 94 million acres of “fire forest” type habitats, which are composed of pine and mixed-pine stands (data extracted from EVALIDator v1.8.0.00). Of these 94 million acres, roughly 6–9 million acres (6.4–9.6%) were burned in 2017 using prescribed fire. This estimate was based on surveys administered to state forestry agencies (M. Melvin, personal communications). If the average time between fire events (i.e., ‘firereturn interval’, detailed below) is 3 years, then an estimated 19–29 million acres are managed using prescribed fire, accounting for 20–31% of the 94 million acres of fire forest communities in the Southeastern U.S. Therefore, a solid understanding of fire management practices is important given the extent to which they are used. The practical implementation of fire management plans depends on land managers making 3 decisions, including when to burn, the frequency of burns (fire-return intervals), and the spatial extent (scale) of burns. The intensity at which a fire burns is also an important component of fire ecology, but is highly dependent on site-specific conditions and beyond the scope of this review. 2.1. Season of burn Historically, fires across the Southeastern U.S. were ignited by lightning strikes and indigenous humans. There has been considerable debate over the frequency each of these sources contributed to fires in North America prior to European settlement (Robbins and Myers, 1992), and in regards to timing when fires were most prevalent. Fires caused by lightning strikes were undoubtedly most common in the growing season (spring and summer; ∼April–August) when vegetation was driest and lightning strikes frequent (Komarek, 1964). In contrast, fires started by indigenous peoples were likely most frequent in the dormant season (late summer through early spring; ∼September–March), which remains common today as most managers prefer to burn during this time (Stanturf et al., 2002). Dormant season burns are often preferred by managers because they are easier to control, and lowertemperature fires should reduce scorch and mortality of mature trees (Robbins and Myers, 1992, but see Sparks et al., 2002). Likewise, dryer air and lower winds that follow wetting rains during the dormant season produce desirable conditions for fire control and smoke dispersion (Cronan et al., 2015). From a wildlife standpoint, dormant season burns were initially thought to reduce risks to ground- or shrub-nesting gamebirds and songbirds because these fires preceded breeding activities (Robbins and Myers, 1992; Knapp et al., 2009). For example, in the Southeast turkeys begin nesting activities in March with peaks in incubation occurring in April (Williams and Austin, 1988; Little et al., 2014; Yeldell et al., 2

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1–3 year range are needed in a portion of the individual home range.

2017a; Wood et al., 2018a). Although the dormant season is the primary period when prescribed fires are conducted, there has been building momentum over the past few decades to incorporate more growing season burns into management planning (Cox and Widener, 2008). There are several reasons for this scheduling shift. First, objectives in pine-grassland systems are to achieve understories dominated by forb and grass species with little woody vegetation. However, many hardwood saplings, such as oak (Quercus spp.) and sweetgum (Liquidambar styraciflua) are not easily killed by fires during the dormant season (e.g., Boyer, 1993; Glitzenstein et al., 1995). Conversely, hardwood saplings are more susceptible to fire mortality during the growing season, particularly after leaf out (Farrar, 1998). Second, historic conditions of pine-grassland systems may be more achievable when fires occur in the growing season due to effects of fire on vegetation structure. Naturally occurring fires (i.e., non-human produced) were most abundant during the growing season, so it seems reasonable that this is when they should occur if the goal is to recreate historic conditions (Robbins and Myers, 1992). Third, studies examining the influence of growing-season fires on breeding birds indicate their effects on nest loss may be either minor or insignificant (Cox and Widener, 2008), likely due to prescribed fire only affecting a small percentage of the landscape at any given time (for an example with turkeys, see Kilburg et al., 2014; Wood et al., 2018a). Likewise, wild turkeys often renest and frequently initiate clutches in stands not scheduled for prescribed fires when fire return intervals exceed 2 years (Yeldell et al., 2017a; Wood et al., 2018a). However, studies examining effects of growing season fire on non-cavity nesting passerines in the Southeast have been restricted to only a few species, including Henslow’s sparrow (Ammodramus henslowii; Thatcher et al., 2006) and Bachman’s sparrow (Aimophila aestivalis; Tucker et al., 2004; Cox and Jones, 2007). More work is needed before general conclusions can be drawn about influences of growing season fires on a more comprehensive list of ground-nesting or shrubnesting birds.

2.3. Scale of fire Previous studies examining fire effects on turkeys have focused primarily on season of burn and fire-return intervals, but the scale of fire is rarely discussed or sufficiently reported. The scale of fire management can be thought of as forming a hierarchy (Fig. 2). The top-most level is the total area being managed (i.e., total management area), which is comprised of both fire-free and fire-managed habitats. Note that our use of total management area here is synonymous with the more typically used term, study area. Basically, it is the geographic area within which population(s) are studied. The second level of the hierarchy is the area within the total management area that is subjected to fire (i.e., fire managed area) over some defined temporal period (e.g., several years to several decades). Defining the temporal period of fire history is important, otherwise the entire management area would likely be considered a fire management area given the historic prevalence of fires in the Southeast. The third level of the hierarchy is the sum of the area burned each year (i.e., annual burn area). Finally, the fourth level of the hierarchy is the area of individual burns (i.e., burn compartment area). Note that the area of each level of the hierarchy is nested within the area of the next highest level. The term fire rotation is standard and commonly used to describe the amount of time required for an area equal to a defined area of interest to burn. We developed these terms and definitions, with the exception of fire rotation, to explicitly aid with synthesis and development of this paper, but they are not necessarily standard terms used by federal and state land or wildlife management agencies. Very little information exists on the appropriate scale of fire for managing turkey habitat. Stoddard (1963) recommended managing ∼33% of turkey range as fields and pastures based on his experience in Georgia, although it is unclear how he arrived at this number. Managing this percentage of turkey range in early successional plant communities would translate to a fire management area of roughly 33% of the total management area based on our definitions, but does not provide information on the annual burn area. Speake et al. (1975) recommended that spring and summer habitat should include 12–25% of “well dispersed” openings based on turkeys studied in Alabama and Kentucky. This recommendation appears to be based on dispersed openings that resulted in the lowest distances of spring movements into nesting and brood-rearing habitats. If openings are maintained by prescribed fire, this recommendation translates to a fire management area that is 12–25% the size of the total management area. Similar to Stoddard (1963), information was not provided to determine an annual burn area. Hurst (1978) recommended that “one-third of a given forest compartment should be burned annually.” Here, the statistic provided is not particularly helpful, but probably translates to an annual burn area that is 33% of the fire management area. These 3 reported statistics highlight problems caused by a lack of common definitions available for studies evaluating how fire effects turkeys and other wildlife. The scale at which prescribed fire is applied has implications for maintaining the amount of treeless cover types, given that fire is a primary tool used to maintain treeless, early successional plant communities. The average fire management areas have been reported in most studies, although only recent studies reported the average burn compartment area. In 5 studies reporting average burn compartment area, areas ranged from 10 to 485 ha, with an average of 90 ha (area of burned patches presented in Martin et al., 2012; Kilburg et al., 2014; Little et al., 2014; Yeldell et al., 2017c; Wood et al., 2018a). Several studies reported only the fire management area over the period of study (Campo et al., 1989; Sisson et al., 1991; Stys et al., 1992). In these studies, the fire management areas presumably followed management plans, but determining the area optimal for turkey management has not yet been addressed. Furthermore, the size of the average burn

2.2. Fire-return interval Frequency of fire is an important component of habitat management plans in pine grasslands, and the fire-return interval, severity, and timing of fires, determines vegetative response and resulting plant community composition (Thaxton and Platt, 2006). Understory vegetation becomes dominated by hardwoods when fires are excluded from pine-grasslands, and shading eventually limits grasses and herbaceous plants. Frequent fire return intervals (e.g., 1–3 years) are necessary to maintain desirable herbaceous ground conditions associated with pinegrassland communities (Waldrop et al., 1992; Brockway and Lewis, 1997; Glitzenstein et al., 2012). This fire-return interval range is commonly reported for maintaining pine-grassland plant communities. Turkeys use a variety of habitat types throughout their annual cycle, which requires land managers to maintain a range of vegetative conditions. For example, nesting females select dense vegetation with high visual obstruction from predators (Badyaev, 1995; Streich et al., 2015), often near openings in forested and shrub habitats (Byrne et al., 2015). Conversely, brood habitat generally contains relatively less understory vegetation that facilitates foraging by poults and female vigilance (Burk et al., 1990; Jones et al., 2005; Spears et al., 2007). Concealment cover is an important attribute of brood habitat, particularly when poults are < 2 weeks old and roost on the ground (Metzler and Speake, 1985; Speake et al., 1985). Such conditions can generally be found in areas 2 years after burns (Wood et al., 2018a), but once poults begin roosting in trees, areas that have not been burned in 3 or more years may be used (Wood et al., 2018a). During the non-breeding season, turkeys often use hardwood forests in bottomlands, drainages, and swamps (Porter, 1992; Miller and Conner, 2007), which are typically not exposed to regular fire events. Therefore, variable habitat conditions required by turkeys throughout their annual cycle depend on fire history (Fig. 1), and fire-return intervals outside of the commonly cited 3

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Fig. 1. A series of photographs monitoring time-since-fire influences on vegetation structure at a forest stand located at the Jones Ecological Research Center at Ichauway in Southeastern Georgia, USA. A prescribed fire occurred on 21 October 2003, and the first photograph was taken shortly thereafter in November, followed by photos taken at the same time in subsequent years through 2008.

3.1. Habitat use

compartment area is often confounded or not distinguished from the annual burn area, making it difficult to determine which is driving turkey responses.

The hierarchical process of behavioral responses by individuals that leads to disproportionate use of habitats is termed habitat selection (Jones, 2001), whereas habitat use is simply descriptive of where animals are and is the end result of habitat selection. There is an assumption that habitats selected for by individuals provide fitness benefits over those selected against (Martin, 1998). Understanding habitat selection is important because it provides information on areas in need of protection or restoration (for a practical example see Aldridge and Boyce, 2007). Habitat selection may vary by season and demographic class (i.e., age and sex), and often varies by life stage (e.g., prebreeding, nesting, brood rearing). Habitat studies have been extensively reported for turkeys, but many only described habitat use rather than selection, the difference being that habitat selection studies analyzed location data in a used-available context compared to other studies, which simply described habitats where turkeys were found. We summarize findings from habitat selection studies with respect to turkeys in landscapes managed with prescribed fire. In the context of prescribed fire, there are 2 primary questions of interest relating to how it influences habitat selection. First, do turkeys preferentially select or avoid areas with a burn history? In either case, understanding the time scale and at what successional stages these choices occur is important. Second, how do these habitat preferences vary by season and

3. Fire effects Herein, we review direct effects of prescribed fire on turkeys and provide generalized conclusions, while also identifying knowledge gaps. We partition these effects into 2 categories most easily estimated in studies collecting individual-level data (e.g., VHF- and GPS-marked birds): effects influencing how turkeys use the landscape (habitat use and movement), and effects directly influencing population growth (nest and brood mortality). The influence of fire on turkey behavior and demographic rates are of primary interest because they ultimately provide information on what habitat conditions are needed to support turkeys, and if these habitat conditions influence whether individuals survive and reproduce. We reviewed relevant studies using search engines (Google Scholar and Web of Science) and targeted search terms (e.g., “Meleagris gallopavo” and “fire”), reviewed conference proceedings (National Wild Turkey Symposium, Southeastern Association of Fish and Wildlife Agencies) and the Tall Timbers Research Bulletin, and identified and reviewed literature cited in studies located using the aforementioned methods. 4

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Fig. 2. A hypothetical example of the hierarchical scales of a prescribed fire management plan. The full grid in each panel represents the total management area in 3 different years. The fire management area is represented by light yellow (a). Prescribed fire is applied within the fire management area during the first year, and the sum of these burn compartment areas is equal to the annual burn area (b). By the second year 66% of the fire management area has been burned (c), and by the third year all the fire management area has been burned. This example represents a fire management area that is ∼30% of the total management area, with an annual burn area that is ∼33% of the fire management area. A management plan with a 3-year fire return interval and a 3-year fire rotation could produce similar results. (For interpretation of the references to colour in this figure legend, the reader is referred to the web version of this article.)

objectives of each study, they posed comparative challenges for synthesis, so we attempted to coarsely consolidate findings. In some cases, studies could not be directly compared to others because their findings were not reported in a comparable way (e.g., not providing information on habitats selected, but providing information on fire-return-intervals selected, and vice versa). Pre-nesting – The pre-nesting period is often defined as occurring from roughly the beginning of March until onset of laying. Habitat selection may be particularly important during this time period due to effects on fecundity (Badyaev, 1995; Chamberlain and Leopold, 2000). Cover types selected during pre-nesting generally lack understory and mid-story cover. Most recent studies noted that female turkeys selected hardwood forests during pre-nesting (Martin et al., 2012; Little et al., 2016; Yeldell et al., 2017c; Wood et al., 2018b), whereas Kilburg et al. (2015) also found that transitional zones between treeless areas and forests were selected by females. Little et al. (2016) found that shrubscrub communities were avoided in Georgia, whereas Yeldell et al. (2017c) found that mature pine forests were avoided in Louisiana. Only

demographic class? Most habitat selection studies on turkeys have focused on females because of interest in reproduction, although studies have included marked males (Godwin et al., 1992; Martin et al., 2012). We separated studies into pre-nesting (period before egg laying), nesting (laying through incubation), brooding (post-hatching through fledging), and non-breeding (fall and winter) seasons, because this was the most generalizable way it could be reported from the literature. These lifehistory divisions varied across studies. At one extreme, 3 distinct periods were used to distinguish habitat selection throughout the calendar year (Martin et al., 2012), but at the other extreme, 7 distinct categories were used for the breeding season alone (Stys et al., 1992). Moreover, most studies used fixed seasonal windows (e.g., Little et al., 2014) by defining the start date of the egg-laying period as the median of all first reported egg-laying events. Yeldell et al. (2017a, 2017c) used seasonal windows that were unique to each female since high-resolution GPS marks allowed researchers to determine precise dates of each life-history event for each female. While these differences were sensible for 5

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Southeast, habitats burned within the previous 2 years appear to be important for nesting females. Brood-rearing – The brood-rearing period is a dynamic time when females must make habitat selection decisions based on changing resource requirements of poults as they age and become able to roost in trees (Williams and Austin, 1988; Healy, 1992). During the day, understories that allow poults to move freely and secure arthropods are important (Hurst, 1978; Metzler and Speake, 1985; Wood et al., 2018b). Little et al. (2016) reported that females preferred hardwood forests, pine forests, and shrub-scrub cover during brood-rearing, while avoiding treeless cover types. Martin et al. (2012) found that females selected hardwood forests and avoided pine forests during broodrearing. Sisson et al. (1991) also found that females in Georgia avoided pine forests during brood-rearing, but in contrast to Little et al. (2016), noted that females selected for treeless cover types. Wood et al. (2018b), in agreement with Martin et al. (2012) and Little et al. (2016), found that brooding females selected hardwood forests. They also found that treeless cover types, young pine forests, and mature pine forests were selected for during brood-rearing. Broods also roosted at sites not recently burned (3–6 years post-fire), but selected diurnal use areas burned 2 years prior. Yeldell et al. (2017c) reported that females selected against hardwood forests and mature pine forests burned > 3 years prior during brood-rearing. Campo et al. (1989) reported brood-habitat use in managed pine forests in east Texas and noted greater selection by females with broods in burned areas, but the burn history was not provided. Regardless, a certain percentage of the landscape with a 2-year fire-return interval in forested cover types appears to produce vegetative conditions selected for by brooding females. However, forests lacking such a fire-return interval are also likely important to broods once roosting in trees occurs at 2 weeks of age (Barwick et al., 1970; Phalen et al., 1986). Collectively, these results indicate general selection for forests (hardwood and pine) and treeless cover types with relatively recent burn histories (generally ≤ 2 years). Non-breeding – During the non-breeding season, insects, soft mast, and leafy vegetation become less prevalent in the diet of turkeys, while hard mast begins to dramatically increase (McRoberts et al., 2014). Not surprisingly, hardwood forests are recognized as being critical nonbreeding habitat to wild turkeys, because of foraging resources and availability of roosts (Chamberlain and Leopold, 2000; Miller et al., 2000). Unfortunately, most contemporary studies did not examine habitat use in areas with frequent burns during the non-breeding season, although Little et al. (2016) reported that hardwood forests and pine forests were selected for during this time, whereas shrub-scrub cover types were selected against. Yeldell et al. (2017c) reported that hardwood forests, mixed forests, and treeless cover types were selected, but these cover types had no recent fire history. Other periods – One study of GPS-marked female turkeys in Louisiana examined the relationship between use of recently burned areas and presence of escape cover (Yeldell et al., 2017b). The authors estimated the probability that turkeys would use burned areas within 250 days post-fire. Turkeys readily used recently burned stands and were sometimes found in burned stands the same day a burn occurred. Use of burned areas increased up to 141 days after fires occurred, before declining, and there was a relationship between the size of a burn and turkey use. Turkey space use within burned areas declined as distance to unburned areas increased, suggesting that turkeys favored edges of burned and unburned areas that could serve as escape cover. The strength of this relationship declined as time-since-fire increased. The authors concluded that smaller burn units were likely to be beneficial for turkeys because they provide more edge cover in the form of shrubs and trees for concealment.

one study reported on habitat preferences of males during pre-nesting; Martin et al. (2012) found that males in Georgia did not preferentially select any habitat types during this period. However, habitats were sometimes selected based on their fire histories during pre-nesting. Martin et al. (2012) reported that females selected hardwood drain cover types burned in the previous 2 years. Kilburg et al. (2015) reported that females selected edges between forests and treeless areas burned during the same growing season in North Carolina, and Yeldell et al. (2017c) reported that females avoided mature pine forests not burned (during either dormant or growing season) in the previous 2 years. Other studies examined habitat selection of turkeys in areas managed with prescribed fire, but the successional stage of selected cover types was not reported. Godwin et al. (1992) found that both juvenile and adult males in Mississippi preferentially selected forested cover types in spring (dates of which corresponded roughly to pre-nesting), but fire history of forested cover types selected was not reported. In Mississippi, Palmer and Hurst (1998) found that females used areas with lower ground cover height, more herbaceous vegetation, and less woody vegetation, conditions found in areas burned 0–1 year prior. Females used areas burned > 2 years prior less than their availability. Across studies, other cover types with varying fire-return intervals were not reported as being selected for or against. Therefore, it appears that forested edges along treeless cover types with relatively recent fire histories (≤2 years) were selected by females during pre-nesting. Nesting – Habitat selection during nesting is perhaps the most important factor influencing fecundity of female turkeys as reproductive success is a primary driver of population sustainability (Vangilder, 1992; Palmer et al., 1993; Roberts and Porter, 1996; Thogmartin and Johnson, 1999; Park et al., 2001). In contrast to pre-nesting, females typically select habitats with more shrub cover during nesting (Streich et al., 2015). However, Davis et al. (1995) found that females in South Carolina did not preferentially select any cover type for nesting, but did find that nearly 90% of nests were located in stands subjected to periodic prescribed burning (33% in stands burned < 1 year before nesting, 28% in stands burned 1–2 years before nesting, 5% in stands burned 2–3 years before nesting, and 23% in stands burned > 3 years before nesting). Kilburg et al. (2014) reported that transitional zones between pine and hardwood forests were selected by nesting females, whereas Little et al. (2016) and Martin et al. (2012) both reported that hardwood forests were selected during nesting. Little et al. (2016) also reported that nesting females selected shrub-scrub cover types and pine forests, whereas Martin et al. (2012) found that females avoided pine forests. Little et al. (2014) reported that nesting females selected areas burned the same season (within the previous 2 months). Martin et al. (2012) reported that nesting females selected areas burned in the previous 2 years, and Wood et al. (2018b) reported nesting females in Georgia preferred treeless cover types and mature pine forests, the latter of which was preferentially selected for when fire occurred within the previous year. Yeldell et al. (2017a, 2017c) also reported that mature pine forests were preferred, and stands burned 2 years prior were selected (Yeldell et al., 2017a). Mixed oak-pine forests burned the previous year were also selected during nesting (Yeldell et al., 2017c), in addition to mature pine forests burned 2 years prior (Yeldell et al., 2017a). Across studies, females exhibited greater plasticity in habitat selection during nesting than pre-nesting. Notably, habitats burned ≤2 years prior were often selected by nesting females. Stys et al. (1992), in a study of radio-marked females in Mississippi pine plantations, reported that older forests were preferred for nesting (stands burned < 1 year at time of nesting were avoided, whereas those burned the prior 1–6 years were preferred). These results were similar to Burk et al. (1990), who found that nesting females in Mississippi pine plantations generally selected habitats burned an average of 3 years prior. Conversely, previous contemporary studies all occurred in areas where pine plantations were absent or only a minor percentage of available habitat. Therefore, in non-plantation pine forests of the

3.2. Nest and brood survival Direct effects of prescribed fire on turkeys is of great interest 6

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to fire annually if they were randomly located within the study area. However, they noted this was an inflated exposure rate because not all nests in burned areas are lost to fire (e.g., nests located in depressions or fire shadows). Overall, literature suggests that risk of nest exposure to prescribed fire is low and few nests are lost to fire. Nonetheless, valid concerns exist about influences of growing season burns, especially those during peak nesting, given variation in fire frequency and scale of fires. Earlier studies suggested that because females preferentially nest in areas with greater concealment, and because these conditions typically occur in patches not recently burned, these patches may be more likely to be on schedules for burning during nesting season (Burk et al., 1990; Sisson et al., 1990). However, as noted above, contemporary studies suggest the opposite; females preferentially nest in stands burned within the previous 3 years as these areas contain sufficient concealment cover. Sisson and Speake (1994) concluded that spring burning posed a threat to wild turkey nests, but in light of recently published works reviewed, the risk was probably overstated and must be considered in the context of the percentage of nests likely to be exposed to fire. We note that nest exposure to fire, as reported in contemporary literature, was minimal, but prescribed fires occurred at greater frequencies during the dormant season when nests were not active – an important point to consider. Broods – Wild turkey poults are precocial and capable of moving considerable distances from the nest within 24 h of hatching (Healy, 1992). This mobility inherently makes broods less susceptible to fire, although fire does pose risks to younger poults incapable of flight. Less is known about effects of fire on broods compared to nests, and as with nests, dormant season fires pose no direct risks to broods. Little et al. (2014) studied brood survival (n = 34) in southern Georgia over a 3-year period. Prescribed fires were primarily conducted during January–July and were distributed throughout the study areas in a mosaic pattern. One of 34 (∼3%) broods was lost to prescribed fire. The authors concluded that prescribed fire had little impact on brood survival, and that population-level impacts resulting from nest and brood loss were likely to be dampened as a result of high renesting rates of females. Wood et al. (2018a) estimated brood (n = 29) survival in southern Georgia over a 2-year period, on a site managed primarily with dormant season fire in one year (63%) and growing season fire the next (92%). No broods or poults were exposed to prescribed fire. Likewise, Yeldell et al. (2017a) noted that none of the monitored broods (n = 12) were exposed to fire on a site managed primarily (71%) with dormant season (December–March) fires. Collectively, existing literature suggests that broods on fire-managed landscapes rarely suffer direct mortality from fire, and broods regularly select recently burned stands (Wood et al., 2018a), often immediately after fires occur (Yeldell et al., 2017b).

because of potential links to fitness. This interest has prompted recent studies, although some authors essentially dismissed the potential for direct effects of prescribed fire on turkeys (Brennan et al., 1998). Direct effects of fire on turkeys can be separated into potential effects on nests and poults, as direct effects on adults is highly unlikely given their mobility. Nests – Moore et al. (2005, 2010) summarized previous studies conducted in the upper coastal plain of South Carolina using radiomarked females to monitor nests in the presence of prescribed fire (studies conducted by Moore et al., 2002; Carlisle, 2003). Prescribed fire was applied annually to 1.3% of the total management area during the growing season, and 7.5–10% during the dormant season (i.e., annual burn area was 8.8–11.3% of total management area). The authors stated that fires were planned on a 3–5 year rotation (not fire-return interval) to enhance conditions for red-cockaded woodpeckers. Of 22 females monitored, 2 (9%) had nests destroyed by prescribed fire, and one female renested after her first nest was destroyed. The authors concluded that the population was likely to be minimally affected by growing season prescribed fires given the small areas burned during this period. Some of the most comprehensive studies on effects of fire on turkey nests have been conducted in southern Georgia. Little et al. (2014) detailed influences of fire on nests on a site with fire return intervals ranging from 1 to 3 years. The authors monitored 78 nests and found that only 5 were exposed to prescribed fire, and 2 of these 5 nests were successful. Hence, ∼3% of nests located in cover types actively managed with prescribed fire were directly lost to fire, but the frequency of fire during this period was not reported. The authors concluded that the risk of fire to turkey nests was minimal due to low exposure rates, and most likely mitigated by the propensity of females to renest. A second study conducted in an area which overlapped with the study areas of Little et al. (2014) was conducted by Wood et al. (2018a). The authors monitored 76 nests in areas treated with prescribed fire primarily during the dormant and early growing season and found that no nests were exposed to fire throughout the incubation period, although 4 (5.2%) would have been, had they not been lost to predation. Likewise, Wood et al. (2018b) found that female turkeys nested in areas with successional stages < 3 years post fire, suggesting that females typically do not choose to nest in areas scheduled for burns. Yeldell et al. (2017a) monitored 69 nests of radio-marked female turkeys in west-central Louisiana to evaluate nest site selection on a site with fire return intervals of 3–5 years. The annual burn area was ∼ 20% of the total management area. Prescribed fire was applied during the dormant (December–March) and growing seasons (April–July), although most (> 70%) occurred during the dormant season. One nest was exposed to fire during egg-laying, but the nest was not destroyed or abandoned. No nests were exposed to prescribed fire during incubation. Because most prescribed fire occurred during the dormant season prior to nesting, exposure of nests to fire was limited. However, most females selected nest sites in forest stands that were at successional stages of 2 years after fire events. The authors concluded that even if a greater proportion of the landscape had been burned during the growing season, exposure of nests to fire was unlikely given that fire would occur in stands where females were not nesting. Exposure rates of turkey nests to fire was formally estimated by Kilburg et al. (2014). Female turkeys were radio-marked in North Carolina on a forested site managed with both dormant and growing season fires. Unlike other studies, growing season fires were the most frequent and occurred from March–June. The annual burn area was approximately 25% of the total management area, and roughly 20% of the total management area was burned when nests were active. The authors calculated the percentage of nests exposed to fire each week of the nesting season by multiplying the percentage of the land area burned each week by the percentage of females actively nesting. They used 30 nests in the analysis, 1 (∼3%) of which was lost to prescribed fire. The authors estimated that 6% of nests would have been exposed

3.3. Movement The most immediate and necessary behavioral response of turkeys to prescribed fire is movement out of stands where burns occur. Work on turkey movements in response to fire is recent and has been aided by use of high-resolution spatial data collected with GPS transmitters. Cohen et al. (In Press) used spatial location data from GPS-marked female turkeys captured in southern Georgia and west-central Louisiana to investigate how individuals responded to fire in areas recently burned (< 250 days). Both study areas were subjected to fire within the dormant (December-March) and growing seasons (April-July). The authors found that females often immediately moved back into stands after fires occurred. Individuals tended to loaf more (i.e., remain stationary) in burned areas as time-since-fire increased; this positive relationship suggested that conditions immediately after fire were probably not conducive to loafing given a lack of concealment cover, but conditions progressively improved post-fire. Turkeys were also more likely to walk through recently burned areas rather than loaf as distance from unburned stands increased, indicating the denser peripheral 7

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vegetation was important for escape cover. One important insight from this study was that interior areas > 250 m from unburned areas were unlikely to be extensively used by turkeys given their distance from escape cover was too great; hence, the authors suggested burning smaller patches and working to maximize perimeter to area ratios when implementing burn regimes, the same conclusion reached by Yeldell et al. (2017b).

exist. In fact, such situations are likely the norm and not the exception, even if the management plan does not call for such variability. Nonetheless, we hope readers find this simplified example useful as a heuristic for understanding and describing prescribed fires in their study systems.

4. Information needs

The appropriate scale at which to apply fire to create optimal habitat conditions for turkeys is unknown. Determining the most appropriate scale is perhaps the most difficult question to answer, because it is unlikely that a single metric can be developed and used throughout the highly variable plant communities of the Southeast. Future studies could clarify the appropriate scale of habitats managed with prescribed fire by addressing the following questions. What is the optimal fire management area? Answers to this question will vary depending on the landscape composition of the total management area. If the total management area consists only of closed-canopy forests, then the fire management area is likely small to nonexistent and will need to be expanded. Reductions likely need to occur if the fire management area has eliminated trees producing cover and mast utilized in winter. What is the optimal burn compartment area? The work by Cohen et al. (In Review) and Yeldell et al. (2017b) indicates that large burns produce interior areas mostly unused due to a lack of adjacent cover. However, use of the entire burn area will continuously increase as time since fire increases. Therefore, in the case of larger burns, the temporary loss of the core burn area must be weighed against the habitat it later provides. Conducting a series of smaller burn areas may maximize the immediateuse to future-use tradeoff, but this relationship and recommendations for the appropriate shape and sizes do not exist in published studies. What is the optimal annual burn area? Answering this question will depend on the habitat conditions managers want to create. If the goal is to create more open habitats, the annual burn area will necessarily be high, which will generally correspond to a higher average fire-return interval.

4.2. Scale of fire

Studies examining the response of turkeys to prescribed fire events have produced valuable information for habitat management, but we identified information needs necessary for informed management. We categorize the greatest information needs and review them sequentially, although order does not reflect importance. 4.1. Reporting results and terminology We found general inconsistencies reporting results among studies, and a consistent terminology used to describe different components of fire management was largely lacking. Across studies, fire-return intervals were often reported as ranging between 1 and 7 years. To our knowledge, burning the same forest stands annually is uncommon relative to longer fire-return intervals. Furthermore, we noted that some studies reporting fire-return intervals of 1 year likely were referring to fire-return intervals in certain patches occurring between 1 and 2 years, perhaps due to changes from dormant-season to growing-season fire schedules, or logistical constraints with conducting fires on scheduled burn dates. Reporting a median or mean fire-return interval in months along with the range and standard deviation (e.g., x¯ = 14 months, σ = 7 months, range = 7–20 months) would provide more precise and useful information. We also found that growing season and dormant season prescribed fires were not always explicitly defined in terms of when they occurred, which caused the delineations to have no meaning. Furthermore, statistics noting the frequency of each type of fire were often lacking (i.e., percentage of fire management area and annual burn area attributed to dormant and growing seasons during the study period). Finally, for habitat selection results, time-since-fire successional stages were sometimes reported as being selected for, but the associated cover type they occurred in was not reported. The opposite was also true, as some habitats were reported as being selected for, but the time-since-fire stages in these cover types selected for was not stated. In most studies we reviewed, areas over which fires occurred were reported in ways that lacked context to understand relative scales of fire management. For example, it was common for authors to report fire management areas, but not report the total management area where the study was conducted, although presumably the studied populations may have also occurred in areas lacking a burn history. Similarly, annual burn areas and burn compartment areas were often excluded from site descriptions. Reporting fire-return intervals or burn rotations alone is not particularly useful if the area over which they occur is omitted. Therefore, statistics should be reported for each level of the hierarchy, including a measure of area (e.g., in units of hectares), as well as a percentage or proportion of the burned area relative to the total managed area and fire managed area. For example, if the total management area is 10,000 ha, and 40% of the total management area is fire management area, then the fire management area is 4,000 ha. A possible point of confusion regarding terminology may be the hypothetical example we present in Fig. 1. For simplicity and ease of understanding, this is clearly an idealized management scenario where each year the annual burn area comprised ∼ 33% of the fire management area, which corresponds to a scenario where all burn compartment areas have a 3-year fire-return interval. In reality, logistical constraints will generally preclude adhering to strict fire schedules for every burn compartment area, and a far more variable fire history will

4.3. Growing season fire Managers now have research to help inform decisions about appropriateness of growing season burns in their habitat management programs. However, most studies presenting results on fire exposure events occurred in study areas where dormant season fires were more common than growing season fires. Thus, risks of nests and broods to fire are expected to be low in these studies. Works by Kilburg et al. (2014) and Wood et al. (2018b) were the exceptions because growing season fires were more frequent at these sites (during one year of Wood et al., 2018b). Results from both works indicated exposure of nests and broods to prescribed fire were low, although there was temporal overlap between fire, nesting, and brood rearing. More studies assessing survival of nests and broods under management regimes where growing season fires are used at higher frequencies will be beneficial. To date, the current information available indicates low occurrence of nest and brood mortalities in areas where prescribed fire and breeding events overlap. 4.4. Demographics and growth Most studies reporting responses of turkeys to prescribed fire have focused on females, but age classes of females have not been distinguished and estimated responses of yearlings (< 1 year) and adults (≥1 year) were pooled. However, habitat use and home ranges of turkeys may vary by age and sex class (Badyaev et al., 1996). Further research detailing age and sex-specific responses of turkeys to prescribed fire is necessary, because potential differences could be predictive of a population-level response to prescribed fire. Producing predictions of such population-level responses requires population 8

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models and understanding relationships between fire effects and vital rates, which are responsible for variation in abundance and growth. Ultimately, managers are interested in how turkey populations respond to prescribed fire via changes in turkey abundance. Herein, we summarized reports of direct effects of fire on nests and broods, but estimates of individual survival, fecundity, and population change in response to prescribed fire are lacking. An ideal study design would be comparative in nature and estimate demographic rates independently for populations occurring in areas managed with and without prescribed fire. Comparisons among sites with varying levels of prescribed fire would be highly informative, but may be challenging to implement in practice. Nonetheless, information obtained from experimental studies is valuable and provides improved inferences (Morrison et al., 2008).

29% of the landscape within their annual range, and Farrell et al. (2019) found that the probability of occurrence of turkeys stabilized when the proportion of hardwood forests exceeded 20%. Accordingly, a coarse rule-of-thumb might be to manage forests such that the fire management area does not reduce hardwoods below 30% of the landscape. Therefore, any forest cover types above this percentage can be managed with prescribed fire to create turkey habitat. 6. Conclusions The use of prescribed fire as a management tool for managing, maintaining, and reclaiming pine-grassland ecosystems throughout the Southeastern U.S. is likely to continue, and perhaps increase. This offers continuing opportunities to improve and expand pine-grassland habitats, which is beneficial for wildlife. However, it is important that wildlife managers understand how prescribed fire influences the populations they manage. Doing so will require carefully thought out study designs and measuring population responses to habitats manipulated with prescribed fire. We encourage researchers and wildlife managers to work together to devise studies aimed at addressing the information needs outlined. Presenting results from these studies in a consistent way using common terminology will greatly benefit our collective understanding of prescribed fire effects on turkeys and other wildlife.

5. Management recommendations Season of burn – Most studies reviewed noted that dormant season fires were prevalent in habitats used by turkeys (Yeldell et al., 2017a, 2017b, 2017c; Wood et al., 2018a, 2018b; Cohen et al., 2019). We offer that use of growing season fires for turkey management hinges on the ability to apply these fires to patches of habitat where turkeys are unlikely to be nesting. We recommend that growing-season fires be considered as part of management plans where turkeys are a focal species, particularly where managers are unable to meet management objectives with dormant-season prescribed fire alone. However, we urge caution given that literature detailing influences of growing season fires on turkey nests and poults are relatively recent and more replicated studies are needed. We also urge managers to use data generated from studies monitoring turkeys with VHF or GPS units in areas where growing-season prescribed fires are implemented. Reporting rates of nest and brood loss from fire events is important to understand fire risk. Managers should obviously consider other species co-occurring with turkeys and consider potential effects of growing-season burns on these species. Growing-season fires applied on a fire-return interval ≤ 2 years may pose risks to turkey nests and broods, as these stands are preferentially selected by females during these reproductive stages. Fire-return interval – Several studies have provided recommendations on fire-return intervals needed to maintain habitat conditions selected by turkeys in the Southeast. In 8 studies providing recommendations, 6 suggested fires should occur ≤ 3 years (Little et al., 2014, 2016; Streich et al., 2015; Yeldell et al., 2017a, 2017b; Wood et al., 2018a), 1 suggested 3–4 years (Miller et al., 2000), 1 suggested 3–6 years (Stys et al., 1992), and 1 suggested 4–5 years in upland stands (but every 2–3 years in lowland thickets; Kilburg et al., 2014). It should be noted that these recommendations were often specific to create habitat for certain life-stage conditions during reproductive periods (e.g., nesting or brood rearing), and were not necessarily comprehensive recommendations to maintain conditions across all habitats used throughout the annual cycle. We agree with the conclusions of others (e.g., Little et al., 2016; Wood et al., 2018a,b) that fire-return intervals of ≤ 3 years distributed in a mosaic pattern throughout fire-managed areas are appropriate for turkeys. Proximity of recently burned patches to unburned areas is important, as noted in several studies detailed herein. Scale of fire – No studies produced recommendations for optimal size of burn compartment areas, but presumably there is an optimal range of sizes that balances trade-offs between available cover and burn area. For example, Yeldell et al. (2017b) found that areas in the interior of a burn > 250 m from the edge were avoided. Therefore, it may be beneficial to create burn compartment areas with geometries that leave most of the burn area below this distance to edge (e.g., rectangular shapes with high width to length ratios). Likewise, managers should consider avoiding fire management areas large enough to exclude hardwood forests. Davis et al. (2017) found that male turkeys in Mississippi reached highest abundance when hardwood forests composed

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