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Intensive Selective Deer Browsing Favors Success of Asimina Triloba

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Intensive Selective Deer Browsing Favors Success of Asimina Triloba R E S E A R C H A R T I C L E ABSTRACT: Although white-tailed deer (Odocoileus virginianus Zimmermann) are generalist herbivores, they can have significant effects on species composition and abundance of forest trees, especially when deer densities are high and most plant species are heavily browsed but a few are selectively avoided as browse. We evaluated effects of selective deer browsing on tree species abundance in an old-growth mesic/wet-mesic forest in central Illinois by repeated sampling of permanent study plots in 2003 and 2008 and relating changes in stem density between the two samples to the intensity of deer browsing of • individual species. The study site has high deer density (75 deer km2) during winter months, and initial observations indicated that paw paw (Asimina triloba (L.) Dunal) was strongly avoided as browse. Paw paw density increased in the seedling stratum between the two sample periods. However, nearly all other tree species declined in density; blue ash (Fraxinus quadrangulata Michx.) – a low-use browse species, had a small increase in seedling density. Tree species diversity decreased in the seedling stratum, but Intensive Selective not in the sapling stratum, although sapling density declined for some species. The current trajectory of tree species growth and recruitment suggests that the development of a dense paw paw understory canopy will further decrease species diversity and reduce tree species recruitment. Based on the life Deer Browsing history traits of paw paw, and high deer densities and selective browsing, this issue is of concern for Favors Success of current and future forest communities. Index terms: Acer saccharum, Asimina triloba, deer herbivory, Odocoileus virginianus Asimina triloba (Paw INTRODUCTION large-scale land conversion to agricultural paw) a Native Tree landscapes (Pietsch 1954). Recent high Even though white-tailed deer (Odocoileus deer abundance in Illinois resulted from Species virginianus Zimmermann) are generalist protective legislation that banned deer herbivores, they are a major factor affect- hunting in 1901, and federal and state ing regeneration, species composition, sponsored deer reintroductions during the 1930s and 1940s. As a result, Illinois’ 1,3 and abundance of forest trees (Russell et Mitchell A. Slater al. 2001; Horsley et al. 2003; Côté et al. deer population increased quickly and 1Northern Research Station 2004; Aronson and Handel 2011). Deer nuisance levels were reached in several USDA Forest Service browsing is an issue of particular concern counties by the mid-1950s, resulting in 1831 Highway 169 when it promotes dense understories of the lifting of the state-wide ban on deer Grand Rapids, MN 55744 avoided species that further inhibit other hunting in 1957. Deer populations were plant species (Stromayer and Warren 1997; still low to nonexistent in central Illinois de la Cretaz and Kelty 2002; Horsley et al. at this time (Pietsch 1954); however, deer 2003; Rooney and Waller 2003; Goetsch subsequently re-established themselves Roger C. Anderson2 2011; Kain et al. 2011). We examined in the central Illinois region, and now are the relationship between the intensity of considered to have reached nuisance levels 2School of Biological Sciences white-tailed deer herbivory and tree spe- (Anderson 1997). Illinois State University cies abundance and diversity in a central Normal, IL 61790-4120 Illinois old-growth forest subjected to In central Illinois, vast tracts of agricultural intensive winter deer browsing. Notably, cropland produce nearly limitless quanti- this forest site contains a large abundance ties of food throughout the year; however, of paw paw (Asimina triloba (L.) Dunal), a these croplands supply only seasonal cover, • sub-canopy species, which is known to be which is created in the spring and then well-defended against herbivory by plant destroyed during the fall harvest. This chemical compounds (Zhao et al. 1992; seasonal cover creates the pattern of spring Lewis et al. 1993; Zhao et al. 1994; He dispersal from and winter congregation in et al. 1997) and to experience little or no isolated forest fragments, which is pecu- herbivory (Liang and Seagle 2002; Asnani liar to deer found in this type of habitat 3 et al. 2006). (Sparrow and Springer 1970; Nixon et al. Corresponding author: [email protected]: 1991; Nixon and Hansen 1992). While 218-326-7120 Deer have high densities in Illinois at these deer still obtain a substantial portion the present time and are considered to of their diet in agricultural fields, forag- be a nuisance species in some locations. ing on crop residues and winter wheat, However, deer were virtually nonexistent browsing within forested areas is more in many parts of the state by the 1870s intense during winter and early spring and were likely extirpated in Illinois by than during the summer (Nixon et al. the end of the century due to intensive 1991; Kelley 1994). This seasonal pattern Natural Areas Journal 34:178–187 sport and commercial hunting pressure of deer browsing pressure, combined with and destruction of deer habitat from differential preferences for certain plant 178 Natural Areas Journal Volume 34 (2), 2014 species, has likely had substantial effects Study Objectives this time period (1977, 1984, 1994, 2005, on the composition and abundance of tree and 2011) there was a significant positive species in the remaining forest fragments The objective of this study is to examine correlation (R2 = 0.848, P = 0.0264, df = of central Illinois (Strole and Anderson the influence of selective deer browsing 4) between paw paw stem density and deer 1992). High deer densities can prevent on the regeneration success of paw paw killed by fire arms in McLean County in tree species regeneration and, thereby, and other native tree species. We posed the same five years (paw paw stem density alter forest successional patterns (Beals et two questions in this study: (1) Do white- = combined seedlings and saplings ha-1, al. 1960; Ross et al. 1970; Richards and tailed deer have a lower preference for paw natural log transformed, R.C. Anderson, Farnsworth 1971; Anderson and Loucks paw as browse than other tree species? (2) unpubl. data). 1979; Marquis 1981; Horsley and Marquis Did deer herbivory influence changes in 1983; Frelich and Lorimer 1985; Alverson abundance of tree seedlings and saplings Deer exclosures were not available for et al. 1988; Russell et al. 2001; Rooney and of paw paw and other species? this study in Funks Grove, and because Waller 2003). This phenomenon may be of of high deer density, neither were regional particular concern in central Illinois, given METHODS the current high levels of deer abundance, reference forests without deer herbivory. and development of plant communities in Consequently, we examined variation in the absence of deer for nearly a century Study Site intensity of deer browsing on seedlings of (c. 1870 – 1960). all tree species and related this variation to The 27-ha study site is located 35 km changes in seedling and sapling abundances Focal Species southwest of Bloomington, Illinois, within using repeated sampling of permanent the 400- ha Funks Grove (40°21.2’N, study plots. We used this information to Paw paw trees reach a height of approxi- 89°7.7’E). Species composition of the site determine how variation in deer browsing mately 2.1 m after ten years of growth is characteristic of central Illinois mesic/ intensity among species may be shaping the (Larimore et al. 2003), and reach a maxi- wet-mesic forests (Adams and Anderson tree species community at this site and the mum height of approximately 10 m (pers. 1980): sapling and seedling strata at this extent to which paw paw is being avoided observ.); thus, they are understory trees and site are currently dominated by sugar as a browse species. typically do not reach the forest canopy. maple and paw paw, and sugar maple is Although paw paw has not been studied the dominant species in the tree stratum Sampling Design extensively, existing literature suggests that (Smith 2004). it is well-adapted to low-light conditions in Sampling points (n = 105) were located Funks Grove has been affected by high the forest understory. Leaves of paw paw using a stratified random technique (Krebs densities of white-tail deer during the saplings possess photosynthetic properties 1999) within the 27-ha study site (Smith winter months for several years. As an that are similar to those of sugar maple 2004); a 50 × 50 m grid system was estab- example, during the winter of 2007 – 2008, lished and sampling points were emplaced (Acer saccharum Marsh.) (Slater 2011), an aerial deer survey in the Funks Grove at 1-m increments (0 – 9 m, first number) a tree species which is considered to be area estimated a density of > 75 deer to the east or west (even or odd, second highly shade-tolerant (Walters and Reich -2 km (Pers. comm. with E. Smith, Illinois number) using a list of two-digit randomly 2000). Furthermore, the clonal propagation Department of Natural Resources, Spring- generated numbers. At each sampling point, of paw paw decreases the risk of mortality field, 2008), whereas historic deer density measurements of tree diameters at breast by allowing individual ramets to share re- -2 was likely 4 – 8 deer km (McCabe and height [dbh (diameter 1.5 m above ground sources with their parent plants (Hosaka et McCabe 1997). The increased abundance level)], and stem counts of saplings and al. 2005; Hosaka et al. 2008). Given enough of paw paw tracks the local increase in seedlings were taken by species in circular, time, large colonies of paw paw are estab- deer abundance. Using counts of annual nested quadrats (Krebs 1999) with areas lished, which intercept sufficient quantities deer-vehicle collisions (1980 – 2003) on of 0.07, 0.02, and 0.007 ha, respectively.
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