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Forage Availability for White-Tailed Deer Following Silvicultural Treatments in Hardwood Forests
Abstract
Closed-canopy upland hardwood stands often lack diverse understory structure and composition, limiting available nutrition for white-tailed deer (Odocoileus virginianus) as well as nesting and foraging structure for other wildlife. Various regeneration methods can positively influence understory development; however, non-commercial strategies are needed to improve available nutrition in many stands, as some contain timber that is not ready to harvest and others are owned by landowners who are not interested in harvesting timber. Applications of herbicide and prescribed fire have improved availability of food and cover for deer and other wildlife in pine (Pinus spp.) systems. However, this strategy has not been evaluated in hardwood systems. To evaluate the influence of fire and herbicide treatments on available deer forage in upland hardwood systems, we measured forage availability and calculated nutritional carrying capacity (NCC) at 14% crude protein mixed diet, following 7 silvicultural treatments, including controls, in 4 mixed upland hardwood stands July–September 2007 and 2008. We compared NCC among forest treatments and within 4 paired warm-season forage food plots to evaluate the usefulness of food plots in areas where forests are managed. Nutritional carrying capacity estimates (deer days/ha) were greatest following canopy reduction with prescribed fire treatments in both years. Understory herbicide application did not affect species composition or NCC 1 year or 2 years post-treatment. Production of forage plantings exceeded that of forest treatments both years with the exception of early-maturing soybeans and retention cut with fire 2 years post-treatment. We encourage land managers to use canopy reducing treatments and low-intensity prescribed fire to increase available nutrition and improve available cover where needed in upland hardwood systems. In areas where deer density may limit understory development, high-quality forage food plots may be used to buffer browsing while strategies to reduce deer density and stimulate the forest understory are implemented. © 2011 The Wildlife Society.
... Deer and deer hunting, therefore, are an important cultural and economic resource associated with the management of forests. Management of woodlands and forests in the southeastern US for deer often involves increasing the availability of forages by producing and maintaining early successional habitat through timber harvesting, prescribed fire, or a combination of these treatments (Masters et al., 1993b, Lashley et al., 2011, Glow et al., 2019. However, in forests managed for objectives other than deer, such as timber production, or in unmanaged areas, habitat is often marginal because of limited understory biomass production resulting from a dense, often multi-layered tree canopy (Masters et al., 1993b, Sparks et al., 1998. ...
... A basic maintenance diet for an adult deer contains around 6-10% crude protein (CP) (French et al., 1956, Holter et al., 1979, Asleson et al., 1996, National Research Council, 2007. Protein demands are greater at several key life stages for deer including, 14-22% for fawns (Ullrey et al., 1967), 11% for yearling deer (Holter et al., 1979), 11-12% for antler growth (Asleson et al., 1996), and 14% for lactating females (Jones et al., 2009, Lashley et al., 2011, Hewitt 2011. While CP is only one measure of forage quality for deer, it is easily interpreted, well-studied, and correlates well with the key life history processes listed above. ...
... Deer must acquire protein directly from forage as needs cannot be met by body reserves (Sadlier, 1987). Additionally, CP requirements appear more limiting to nutritional carrying capacity than digestible energy requirements in the southeastern US (Jones et al., 2008, Lashley et al., 2011. ...
... Forest management practices that decrease canopy coverage typically increase forage for deer and enhance cover for turkeys. Reductions in canopy coverage are often achieved using various commercial timber harvest methods, and previous work has indicated availability of deer forage improves following implementation of a clearcut, shelterwood harvest, or thinning (Blair and Enghardt 1976, Ford et al. 1994, Peitz et al. 2001, Lashley et al. 2011, Nanney et al. 2018. For landowners without merchantable timber, noncommercial techniques, such as forest stand improvement (FSI), are an option to reduce canopy closure. ...
... For landowners without merchantable timber, noncommercial techniques, such as forest stand improvement (FSI), are an option to reduce canopy closure. Forest stand improvement entails using herbicides to kill undesirable trees within a forest stand and has been shown to increase deer forage biomass and turkey brooding cover in hardwood forests of the Ridge-and-Valley physiographic province (Lashley et al. 2011, McCord et al. 2014. Additionally, FSI treatments that release oak crowns can increase acorn production among remaining trees (Bellocq et al. 2005, Brooke et al. 2019). ...
... However, many upland hardwood forests of the region are composed of tree species that facilitate fire (Kane et al. 2008). Prescribed fire has been applied to hardwood forests managed for deer and turkeys in other regions (Lashley et al. 2011, McCord et al. 2014. Although some managers are hesitant to apply fire that may damage overstory oaks, low-intensity prescribed fire can be applied with little or no damage to residual trees (Brose and Van Lear 1999, Marschall et al. 2014, McCord et al. 2014). ...
Prescribed fire and canopy reduction are accepted forest management practices used to increase forage and cover for white‐tailed deer (Odocoileus virginianus) and wild turkeys (Meleagris gallopavo) in pine systems throughout the southeastern United States. However, use of prescribed fire to improve conditions for deer and turkeys has not been evaluated in upland hardwood forests of the Coastal Plain, and some land managers remain skeptical of the utility of fire in this ecoregion. We designed a manipulative experiment to measure deer and turkey habitat components following canopy reduction and prescribed fire in 4 upland hardwood stands in the Coastal Plain of Alabama, USA, during 2018 and 2019. Specifically, we used herbicide to kill trees with low value to deer and turkeys and retained oaks (Quercus spp.) and other species considered important as part of a forest stand improvement (FSI) operation to reduce canopy coverage. We then applied low‐intensity prescribed fire to half of each treatment unit. One unit in each replicate served as a control. We measured total understory plant coverage, biomass of deer forage, and turkey brooding cover for 2 years following canopy reduction and one year after fire. Coverage of herbaceous plants increased by 134% in FSI/Burn, and coverage of woody and semiwoody plants increased by 33% and 97%, respectively, following FSI only. Deer forage biomass was greater in both FSI and FSI/Burn compared to control, but there was no difference in deer forage biomass between FSI and FSI/Burn. FSI/Burn provided better turkey brooding cover than FSI or control. No overstory trees were killed by fire. We detected minor cambium damage to 13% of water oaks (Quercus nigra) in the FSI/Burn units; other species only showed light bark charring or no sign of burning. We recommend FSI and low‐intensity prescribed fire in Coastal Plain hardwoods to improve brooding cover for turkeys and understory forage for deer while retaining acorn production. © 2020 The Wildlife Society. We applied forest stand improvement and prescribed fire to Coastal Plain hardwoods to evaluate effects on white‐tailed deer forage biomass and wild turkey brooding cover. Our results demonstrate that canopy reduction and fire can be used to increase deer forage biomass and improve structure for turkey broods with minimal damage to retained mast‐producing trees.
... Fortunately, the disturbances responsible for creating and maintaining these communities can also be used for their restoration (McPherson 1997). Prescribed fire stimulates herbaceous plants by making nutrients available for growth (Scharenbroch et al. 2012) and removing layers of leaf litter that inhibit herbaceous germination (Lashley et al. 2011). Fire is essential for oak woodland and savanna restoration (Peterson and Reich 2001), but is slow to alter overstory characteristics (Knapp et al. 2015). ...
... Fire is essential for oak woodland and savanna restoration (Peterson and Reich 2001), but is slow to alter overstory characteristics (Knapp et al. 2015). Mechanical canopy disturbance quickly increases the light available for herbaceous germination and growth (Nielsen et al. 2003;Brewer 2016), but also promotes understory woody vegetation (McCord et al. 2014) that limits herbaceous layer development (Lashley et al. 2011;Barrioz et al. 2013). A long-term regimen of biennial fire can suppress woody undergrowth and maximize herbaceous groundcover and richness (Peterson et al. 2007;Peterson and Reich 2008). ...
... Our results provide insights into the mechanisms behind the positive effects of fire on the herbaceous components of Eastern deciduous forests. Fire promotes herbaceous development by reducing understory woody competition (Knapp et al. 2015), consuming germination-inhibiting layers of leaf litter and debris (Lashley et al. 2011), increasing soil nutrient availability (Scharenbroch et al. 2012), and improving seed germination rates (Emery et al. 2011). In our study, fire reduced litter groundcover by 48% and biennially increased bare ground. ...
Background
In oak-dominated communities throughout eastern North America, fire exclusion and subsequent woody encroachment has replaced the “glitter” of once robust and diverse wildflower and grass layers with leaf-litter dominance. Restoring the important herbaceous components of Eastern oak ecosystems could involve pairing heavy canopy disturbance with growing-season fire, but potential negative effects warrant research. Beginning with 20 ha replicates of closed-canopy forest at three sites across Tennessee and North Carolina, USA, we monitored groundcover response to combinations of thinning (none; light: 14 m ² ha ⁻¹ residual basal area; and heavy: 7 m ² ha ⁻¹ ) and seasonal fire (none; March: pre leaf expansion; and October: pre leaf abscission) from 2008 to 2016.
Results
Before treatments, woody plants and leaf-litter-dominated groundcover and herbaceous plants were rare (<6% groundcover, 118 species). By 2016, herbaceous groundcover averaged 59% after heavy thinning and three biennial burns, and 359 herbaceous species were documented. Only 6% (23) of these species appeared negatively affected by applied disturbances. Across sites, thin-and-burn treatments increased graminoid groundcover 14-fold, forb groundcover 50-fold, herbaceous richness 9-fold, and herbaceous diversity 10-fold, relative to unmanaged stands. These increases were often greater where fire was repeatedly applied, and only after repeated fire was herbaceous response greater in heavily thinned stands relative to lightly thinned stands. Burn-only treatments rarely affected herbaceous metrics, and thin-and-burn treatments more than doubled woody groundcover. This suggests that canopy reduction, leaf-litter consumption, and pulses of bare ground were more related to positive herbaceous responses than to the control of woody competition in the understory. Fire season effects were not observed, but herbaceous response after less intense October fires was comparable to that following more intense March fires.
Conclusions
Our results conflict with warnings concerning the potential negative effects of disturbance on herbaceous diversity east of the prairie–woodland transition zone. Canopy disturbance and repeated fire, regardless of season, widely restored herbaceous groundcover and diversity in Eastern oak ecosystems. Herbaceous components were resilient to extended periods of fire exclusion, but current conservation programs often prioritize existing, high-quality sites. Our results suggest that such policies may overlook the tremendous restoration potential present in otherwise inconspicuous understories of closed-canopy oak forests throughout eastern North America.
... Food and cover resources can be manipulated through forest management activities. Implementing canopy reductions allows light to reach the forest floor thereby improving understory food abundance and structure (Hurst, Campo, & Brooks 1980;Greenberg, Perry, Harper, Levey, & McCord 2011;Lashley, Harper, Bates, & Keyser 2011). However, without periodic disturbances, woody encroachment leads to degraded structure and forage availability for many wildlife species (Crawford 1971;Harper 2007;Jackson et al. 2007;Harper, Ford, Lashley, Moorman, & Stambaugh 2016). ...
... However, without periodic disturbances, woody encroachment leads to degraded structure and forage availability for many wildlife species (Crawford 1971;Harper 2007;Jackson et al. 2007;Harper, Ford, Lashley, Moorman, & Stambaugh 2016). Coupling canopy reductions with periodic prescribed fire controls woody encroachment, creating a two-tiered forest structure with an open overstory and an understory consisting of grasses, forbs, and woody plants (Pack, Igo, & Taylor 1988;Masters, Lochmiller, Engle, Bulletin, & Winter 1993;Lashley et al. 2011;McCord, Harper, & Greenberg 2014). Another common management technique is combining canopy reductions and thinning with selective herbicide application, which removes the hardwood midstory and increases herbaceous vegetation dominance in the understory (Godfrey & Norman 1999;Edwards, Demarais, Watkins, & Strickland 2004). ...
... Given that fine-scale spatial habitat heterogeneity can influence realized patterns of animal space use (Boyce et al. 2003), and that forest management can evoke changes in vegetative structure (Harper 2007;Lashley et al. 2011), the natural conclusion is that common forest management strategies could induce changes in the distribution of associated wildlife communities. To explore this possibility, we implemented multiple forest management practices in a southern pine (Pinus spp.) ecosystem to evoke increased spatial heterogeneity in vegetative conditions within and across forest management units. ...
Habitat quality is often evaluated based on food availability. However, ecological theory suggests cover should be a more important decision rule when food is not a proximate threat to fitness, as cover mediates predation risk as well as other important factors of fitness. In reality, vegetation characteristics related to food availability and cover are rarely coupled with animal use in the same space and time to determine their relative influences on habitat use. Using an array of 81 camera traps in a matrix of forest management strategies used to deliberately cause a wide disparity in vegetation characteristics, we monitored intensity of use by white-tailed deer (Odocoileus virginianus) and wild turkey (Meleagris gallopavo). We measured vegetation characteristics related to food and cover at each camera trap location then used a generalized additive model to determine how vegetation characteristics specific to the location affected intensity of habitat use by animals at the location. Consistent among both species, cover best explained intensity of habitat use. Contrastingly, food did not explain intensity of habitat use well for either species. Some vegetation simultaneously provides cover and food, and our data indicate that areas with vegetation characteristics providing both resources had the greatest influence on intensity of habitat use by both species. Our results suggest deer and turkey may perceive cover as a more important habitat component when food is not a proximate fitness threat.
... Disturbance techniques, such as canopy reduction, prescribed fire, and herbicide applications, may increase forage availability and improve forage quality for elk and whitetailed deer. Canopy reduction methods, such as clearcutting, shelterwood harvest, improvement cuts, and thinning operations, allow increased sunlight to the forest floor, which stimulates additional browse, and herbaceous forage (Collins and Urness 1983, Ford et al. 1993, Strong and Gates 2006, Lashley et al. 2011, Cook et al. 2016). Characteristics of closed-canopy forests in the eastern United States often make it necessary to couple canopy disturbance with prescribed fire to achieve increased forage for cervids (Masters et al. 1993, Sachro et al. 2005, Van Dyke and Darragh 2007, Shaw et al. 2010, Lashley et al. 2011. ...
... Canopy reduction methods, such as clearcutting, shelterwood harvest, improvement cuts, and thinning operations, allow increased sunlight to the forest floor, which stimulates additional browse, and herbaceous forage (Collins and Urness 1983, Ford et al. 1993, Strong and Gates 2006, Lashley et al. 2011, Cook et al. 2016). Characteristics of closed-canopy forests in the eastern United States often make it necessary to couple canopy disturbance with prescribed fire to achieve increased forage for cervids (Masters et al. 1993, Sachro et al. 2005, Van Dyke and Darragh 2007, Shaw et al. 2010, Lashley et al. 2011. Varying seasonality (dormant, early growing season, and late growing season) and frequency of prescribed fire changes vegetation composition, which can affect forage quantity and quality for cervids (Gruchy et al. 2009, VanderYacht et al. 2017. ...
... Forage availability in timber harvest treatments increased up to tenfold in comparison to mature forest stands. Studies in similar regions of the southern Appalachians also reported increases in forage availability and NCC for white-tailed deer following canopy disturbance (Beck and Harlow 1981, Ford et al. 1993, Lashley et al. 2011. Researchers in western forest systems have reported similar increases in summer forage availability and NCC for elk following timber harvest (Hett et al. 1978, Collins and Urness 1983, Strong and Gates 2006. ...
Closed‐canopy forests dominate the landscape across much of the eastern United States and often lack a well‐developed understory, which limits nutrition available for cervids. We evaluated the influence of timber harvest combined with prescribed fire, herbicide treatment, or fire and herbicide treatment in young mixed‐hardwood forests on forage availability and nutritional carrying capacity (NCC) for elk (Cervus canadensis) and white‐tailed deer (Odocoileus virginianus) in the Cumberland Mountains, Tennessee, USA, July–August 2013–2015. We compared forage availability, NCC using a 12% and 14% crude protein nutritional constraint, and vegetation composition in untreated mature forest stands, reclaimed surface mines, and 6 timber harvest treatments (timber harvest only, with early growing‐season fire, with late growing‐season fire, with herbicide only, with herbicide and early growing‐season fire, and with herbicide and late growing‐season fire). Forage availability in treatments involving timber harvest was greater than in untreated mature forest stands and reclaimed surface mines. Forage availability estimates in treatments involving herbicide and prescribed fire were less than all other timber harvest treatments. Nutritional carrying capacity estimates at the 12% and 14% crude protein constraints were greater in timber harvest treatments and on reclaimed surface mines than in untreated mature forest stands. Herbaceous species coverage was greater and woody species coverage was less on reclaimed surface mines and in timber harvest treatments involving herbicide and prescribed fire than in all other timber harvest treatments and untreated mature forest stands. Greater coverage of herbaceous forage species in treatments involving herbicide and prescribed fire and on reclaimed surface mines compensated for reduced forage availability and resulted in NCC estimates similar to all other timber harvest treatments. Our data indicate using periodic prescribed fire and following an herbicide application with prescribed fire are effective techniques to transition young mixed‐hardwood forest communities to early successional communities and maintain increased forage availability and NCC for elk and deer. © 2018 The Wildlife Society. Timber harvest followed with periodic prescribed fire and strategic herbicide applications can greatly increase nutrition available for cervids in mixed hardwood forest systems.
... Biennial fire maximizes community heterogeneity and herbaceous species richness in the understory by suppressing woody vegetation (Peterson et al., 2007;Peterson and Reich, 2008). Increases in understory light can promote a dense layer of woody saplings and shrubs (McCord et al., 2014) which can limit herbaceous layer development (Lashley et al., 2011;Barrioz et al., 2013). Although expensive, the mechanical removal of this vegetation can restore gradients in light, moisture, and nutrients that encourage increases in herbaceous cover and diversity (Brudvig and Asbjornsen, 2009). ...
... However, regionally-specific evaluations of herbicides have narrowly focused on wildlife response to woodland restoration (e.g. Lashley et al., 2011;McCord et al., 2014;Greenberg et al., 2016), regeneration of woody species (e.g. Schweitzer and Dey, 2011), or utilized soil-active herbicides in the absence of a residual overstory (Nanney, 2016). ...
... Controlling these stems with fire is difficult (Arthur et al., 2015), so relaxing the lower size restrictions held during herbicide application could improve results. The understory can be dominated by seedlings even after effective broadcast application of herbicide (Lashley et al., 2011;McCord et al., 2014). In contrast, Nanney (2016) successfully transitioned a young forest dominated by woody regeneration into an early successional community dominated by herbaceous plants after spot-spraying all woody vegetation three years following a clearcut harvest. ...
Establishing herbaceous groundcover is essential for oak woodland and savanna restoration. In the Appalachian region, woody vegetation in the understory can persist through many fires and interfere with achieving this goal. Herbicide applications could reduce such vegetation and interact with canopy-disturbance and fire to accelerate restoration. In stands thinned to woodland (16 m 2 ha −1 , 75% canopy closure) or savanna (5 m 2 ha −1 , 24% canopy closure) conditions and burned biennially in the fall (October) or spring (March), we economically applied triclopyr (Garlon® 3A) to understory woody plants using foliar and cut-surface techniques in the fall between fires. From 2011 to 2013, only minor differences in vegetation were observed between areas managed with canopy-disturbance and fire (CF) and areas where herbicides were also used (CFH). Small-sapling (≥1.4 m tall, < 7.6 cm DBH) density in CF was 2,566 stems ha −1 greater than CFH in 2012. This difference was (1) the only woody control CFH attained beyond CF, (2) only lasted a single growing-season because it was mostly fire-sensitive species top-killed by subsequent fire, and (3) only led to increased herbaceous groundcover in savannas burned in the fall. This included the greatest reported increase in herbaceous groundcover (graminoid +18.2%, forb +8.0%) to be associated with herbicide applications under partial oak canopies in the Appalachian region. Expanding herbicide target constraints, completely removing undesirable seed-sources, increasing triclopyr concentration, exploring tank-mixes, and alternative application timing (e.g., prior to canopy disturbance) could improve effectiveness; however, fire suppression throughout the Appalachian region has increased the dominance of fire-sensitive woody species. Our results demonstrate how such composition can reduce the utility of herbicides relative to fire during oak woodland and savanna restoration.
... To measure diet selection in herbivores, availability and use of foods must be quantified (Norbury and Sanson 1992). Quantifying food availability is commonly accomplished by measuring available biomass of plant species in a given area (Edwards et al. 2004;Wam and Hjeljord 2010;Lashley et al. 2011) or by counting the relative number of stems available of each species (Lashley et al. 2011;Lashley and Harper 2012). Use of foods is measured using three predominant methods: ...
... To measure diet selection in herbivores, availability and use of foods must be quantified (Norbury and Sanson 1992). Quantifying food availability is commonly accomplished by measuring available biomass of plant species in a given area (Edwards et al. 2004;Wam and Hjeljord 2010;Lashley et al. 2011) or by counting the relative number of stems available of each species (Lashley et al. 2011;Lashley and Harper 2012). Use of foods is measured using three predominant methods: ...
... The ability to assess diet selection by vegetation type was important in comparing the relative importance of plants within treatments (e.g. Lashley et al. 2011) and within a single vegetation type (e.g. Lashley and Harper 2012). ...
Context Diet selection is studied in herbivores using three predominant methods: (1) microhistological surveys (identification of plants cell walls remaining in gut contents or faecal excretions); (2) direct bite counts (of tame animals); and (3) indirect bite counts (identifying herbivory on damaged plant tissues). Microhistological surveys and direct bite counts are accurate and provide the potential advantage of linking diet selection to particular individuals. Also, they allow diet selection to be measured in systems with sympatric herbivores more easily than indirect bite counts. However, they require expertise in cell wall structure identification or access to tame animals, and generally require greater expense than indirect bite counts. Conversely, indirect bite counts have the advantages of relatively low cost and time commitment for gathering data and do not require animal observation, but may not be accurate. Aims We tested for similarity between diet-selection estimates calculated by indirect bite counts and microhistological surveys. Methods We performed concurrent indirect bite count and faecal microhistological surveys on white-tailed deer (Odocoileus virginianus) at Fort Bragg Military Installation, NC. Key results The indirect bite count survey assignment of selection was 48% similar to assignments derived from the microhistological survey, based on Jaccard's similarity index. Out of 23 plant species determined to be selected by indirect bite counts, 15 of those species were selected according to microhistological surveys. According to the microhistological survey, eight of the selected plants made up 51% of the overall diet, and seven of those eight were selected according to the indirect bite counts. Conclusions Our data indicate that indirect bite counts may provide a relatively accurate index of the deer-selected plants most important in the white-tailed deer diet, but may be less appropriate to determine selection of plants that infrequently occur in their diet, plants that are typically consumed in entirety, or plants where herbivory damage is poorly identified. Implications Indirect bite counts are a relatively inexpensive and time-efficient tool that may be useful to determine plant species most important to white-tailed deer within a forested landscape, particularly if additional research can improve on associated inaccuracies.
... However, "wildlife" is an ambiguous term (Hunter and Schmiegelow 2011), and burning is not necessarily "good." Indeed, prescribed fire at some application and frequency may promote or enhance habitat for diverse taxa, such as reptiles , Greenberg 2000, Keyser et al. 2004, wild turkey (Meleagris gallopavo L.; McCord et al. 2014), and white-tailed deer (Odocoileus virginianus Zimmerman; Lashley et al. 2011), but its occurrence can be negative for these and other species, such as ovenbird (Seiurus aurocapilla L.; Rush et al. 2012) or eastern box turtle (Terrapene carolina L.; Howey and Roosenburg 2013), depending on when and how the burn is conducted. We contend that a major limitation in accomplishing land management goals related to fire use is failure to articulate precise management objectives or outcome assessment to determine their success. ...
... However, without manipulation of the light environment, understory response following low-intensity fire can be negligible (Shaw et al. 2010, but see Schuler et al. 2013). Lashley et al. (2011) reported that forage availability for whitetailed deer was doubled by implementing low-intensity, early growing season fire under closed-canopy hardwoods, but coupling fire with canopy disturbance resulted in eight times as much deer forage as compared to unthinned and unburned areas. Through time, continued low-intensity fire will remove the midstory, but some level of canopy removal is necessary to allow sufficient sunlight (at least 20 % full sunlight) to the forest floor to stimulate extensive groundcover (Royo et al. 2010, McCord et al. 2014, Knapp et al. 2015. ...
... Many silvicultural practices can be used with prescribed fire in hardwood systems to create, maintain, or enhance food and cover resources for wildlife. Regeneration harvests (e.g., shelterwood and group selection), various types of thinnings, and midstory removal via herbicide applications may be used in combination with fire to manage habitat for wildlife (Brose and Van Lear 1998, Johnson et al. 2009, Lashley et al. 2011, Bakermans et al. 2012, McCord et al. 2014, Kendrick et al. 2015. The amount of canopy removal and resultant increased sunlight directly influences plant composition and structure, as well as habitat for different wildlife species. ...
Fire is being prescribed and used increasingly to promote ecosystem restoration (e.g., oak woodlands and savannas) and to manage wildlife habitat in the Central Hardwoods and Appalachian regions, USA. However, questions persist as to how fire affects hardwood forest communities and associated wildlife, and how fire should be used to achieve management goals. We provide an up-to-date review of fire effects on various wildlife species and their habitat in the Central Hardwoods and Appalachians. Documented direct effects (i.e., mortality) on wildlife are rare. Indirect effects (i.e., changes in habitat quality) are influenced greatly by light availability, fire frequency, and fire intensity. Unless fire intensity is great enough to kill a portion of the overstory,
burning in closed-canopy forests has provided little benefit for most wildlife species in the region because it
doesn’t result in enough sunlight penetration to elicit understory response. Canopy reduction through silvicultural treatment has enabled managers to use fire more effectively. Fire intensity must be kept low
in hardwoods to limit damage to many species of overstory trees. However, wounding or killing trees
with fire benefits many wildlife species by allowing increased sunlight to stimulate understory response, snag and subsequent cavity creation, and additions of large coarse woody debris. In general, a fire-return interval of 2 yr to 7 yr benefits a wide variety of wildlife species by providing a diverse structure in the understory, increasing browse, forage, and soft mast, and creating snags and cavities. Historically, dormant-season fire was most prevalent in these regions, and it still is when most prescribed fire is implemented in hardwood systems as burn-days are relatively few in the growing season of May through August because of shading from leaf cover and high fuel moisture. Late growing-season burning increases the window for burning, and better control on woody composition is possible. Early growing-season fire may pose increased risk for some species, especially herpetofauna recently emerged from winter hibernacula (April) or forest songbirds that nest in the understory (May to June). However, negative population-level effects are unlikely unless theburned area is relatively large and early growing-season fire is used continually. We did not find evidence that fire is leading to population declines for any species, including Endangered Species Act (ESA)-listed species (e.g., Indiana bat [Myotis sodalis Mill. Allen] or northern long-eared bat [M. septentrionalis Trouess.]). Instead, data indicate that fire can enhance habitat for bats by increasing suitability of foraging and day-roost sites. Similarly, concern over burning and displacement of woodland salamanders (Plethodontidae), another taxa of heightened conservation concern, is alleviated when fire is prescribed along ecologically appropriate aspect and slope gradients and not forced into mesic, high site index environments where salamanders are most common. Because topography across the Central Hardwoods and Appalachians is diverse, we contend that applying fire on positions best suited for burning is an effective approach to increase regional landscape heterogeneity and biological diversity. Herein, we offer prescriptive concepts for burning for various wildlife species and guilds in the Central
Hardwoods and Appalachians.
... Forbs were generally more available in MDP and YP at the low deer density site, but forb coverage was relatively limited at all sites. Lashley et al. (2011) and Shaw et al. (2010) noted limited forb availability in forested habitat because of limited sunlight and competition among woody species. Ritchie et al. (1998) and Ritchie and Tilman (1995) found legumes and woody plants more abundant in enclosures where deer browsing had been eliminated, but grasses and forbs were more abundant in the presence of deer. ...
... This is surprising considering the amount of important deer forage plants available in this vegetation type, and especially considering forbs often represent more than half of a deer's diet during spring and summer (Whittington 1984. Forbs are important contributors to deer carrying capacity estimates (Iglay et al. 2010, Shaw et al. 2010, Lashley et al. 2011). ...
... Both forb and trees-shrubs showed a vegetation × season interaction effect, which was a result of canopy closure (Table 6). Lashley et al. (2011) showed less forage available in closedcanopy forest as compared to stands that allowed more sunlight through the canopy following retention cutting and prescribed fire. There was no forb interaction effect in MH because forbs were essentially absent during both winter and summer (Table 7). ...
Given the white-tailed deer’s (Odocoileus virginianus; deer) popularity and potentially negative impact on forested systems; Arnold Air Force Base (AAFB) in Tullahoma, Tennessee, USA has made minimizing negative deer impacts on biodiversity a priority. To address these management issues, I initiated a study on AAFB to investigate deer survey techniques and the effects of deer density on forage availability across vegetative communities.
Current use of infrared-triggered cameras (camera) for estimating deer populations does not provide a measure of precision critical for density estimation. I conducted a camera survey for deer in Wildlife Management Area (WMA) Units 1 and 2 at AAFB, August 2010 and used Program DENSITY to fit a spatial detection function of capture-recapture (spatial modeling) data from the camera surveys of bucks. Spatial modeling can provide reliable estimates of buck density and facilitate our understanding of biases associated with camera surveys for deer.
I compared population and precision estimates from spotlight, ground thermal infrared imaging (ground imaging), and aerial vertical-looking infrared (aerial imaging) surveys in the Security Area (SA) of AAFB, January–February 2010. All 3 techniques provided a precise estimate of deer density. However, the high cost of ground imaging does not justify its use. I also found the potential of road bias in distance sampling to invalidate the technique, unless random transects representative of the study area can be applied. Aerial imaging is less susceptible to road bias, but use should be restricted to large areas where high cost can be justified.
I evaluated the effects of 2 deer densities on forage availability and quality within 4 vegetative communities on WMA Units 1 and 2, and the SA of AAFB 2010. Forage availability was consistently greater during summer verses winter and within middle-aged and young pine stands at the low deer density site versus the high deer density site. Both crude protein and total digestible nutrient values were similar regardless of deer density. I recommend managers consider implementing management practices that would reduce deer density and increase forage availability when forage availability beings to decline and deer density estimates approach levels seen detrimental in literature.
... We confi rmed that Fort Bragg scored an 8 out of 26 on the soil quality index, which is lower in productivity than Ͼ 80% of the soil classes ranked on this scale in the US. Additionally, high-quality forage quantity at Fort Bragg was relatively low in comparison to reports in higher productivity soils (Edwards et al. 2004, Jones et al. 2009, Shaw et al. 2010, Lashley et al. 2011, Lashley 2014. ...
... availability) and number of stems browsed (i.e. use) in a 1.5 ϫ 1.2 ϫ 1.2 m plot (Lashley et al. 2011). We used the structure of damage in remaining forage tissues and the foraging ecology of deer and other wildlife to distinguish herbivory between wildlife species (Lashley et al. 2014b). ...
... We used the structure of damage in remaining forage tissues and the foraging ecology of deer and other wildlife to distinguish herbivory between wildlife species (Lashley et al. 2014b). We calculated a selection index (Chesson index;Chesson 1978, 1983, Edwards et al. 2004, Lashley et al. 2011, Lashley and Harper 2012 by dividing the ratio of use and availability for a given species by the sum of ratios for all species (index cutoff value ϭ 0.007). Th is allowed us to group plants based on the selectivity of white-tailed deer on the site. ...
Cervid studies have demonstrated body weight and lactation may be limited in areas with poor-quality soils, with the underlying mechanism often attributed to poor forage quality resulting from poor soil quality. However, if highly nutritious foods are produced but in limited quantities, selective foraging may alleviate nutritional stressors associated with poor soil productivity when adequate quantities of high-quality forage are obtained. We tested whether poor soil productivity limits forage from being high quality or conversely limits the abundance of high-quality forages. To do so, we determined whether nutrient concentrations in selected and non-selected forages on our poor soil study site met the nutritional requirements of lactating white-tailed deer Odocoileus virginianus assuming adequate amounts of forage are obtained. Also, we compared body weight of yearling males at a high density (13 – 17 deer km Ϫ 2) and low density (3 – 5 deer km – 2), because previous studies concluded soils on the study site were too poor for morphometrics (e.g. body weight) to be density-mediated. Deer selected plant species from each of fi ve forage classes that would meet their nutritional requirements (i.e. assuming adequate forage intake) but also selected for diff erent nutrients across forage classes. Phosphorus was limited in most forages, but deer selected forages that met P-requirements 10 times more than expected. We demonstrated body weight was 7.3% greater when deer density was low than high. Contrary to previous reports from poor productivity soil regions, deer on our study site should be able to meet lactation requirements when the quantity of high-quality forage is not limiting, and similarly body weight should be density-mediated. Hence, management strategies that alter the amount of forage per animal (i.e. decreasing animal density and/or increasing forage abundance) are viable options to promote desirable ungulate morphometrics in poor soil regions.
... Previous studies have reported increases in biomass and quality of Japanese honeysuckle (Lonicera japonica) following fertilization to improve forage for deer (Segelquist et al. 1975, Dyess et al. 1994, as well as an increase in woody browse production (George and Powell 1977). However, forbs generally provide greater nutritional content for deer than the woody species examined in the previous studies (Lashley et al. 2011, Nanney et al. 2018. Shaw et al. (2010) applied soil amendments to a variety of native plants within closed-canopy forests and observed limited changes in forage quality or quantity, but low sunlight availability may have buffered vegetation response. ...
... Past studies on wild plant fertilization for deer focused on vines, shrubs, and trees (Segelquist et al. 1975, George and Powell 1977, Wood and Tanner 1985, Dyess et al. 1994), whereas we sampled four forbs and one bramble species. Deer select forbs when they are available, and they tend to be higher in crude protein than browse (Lashley et al. 2011, Nanney et al. 2018. We were interested in how fertilization might influence quality of forbs and brambles, as they are important diet components of deer during spring and summer when nutritional demands are greatest. ...
Old-field plant communities provide habitat components for several game species, including white-tailed deer (Odocoileus virginianus) and wild turkey (Meleagris gallopavo). Prescribed fire, herbicide application, and disking are commonly applied to improve forage and cover within old-fields, but plant response on sites with nutrient-poor soils is not always favorable. Although it is reasonable to expect vegetation to respond to liming and fertilization, little information exists on how forage nutrient content and vegetation structure of old-field plants are influenced by soil amendment. We designed an experiment to test the effects of three amendment treatments (lime, fertilizer, lime+fertilizer) on four fields across Tennessee. We tested soils during spring 2017 and 2018 and applied treatment amendments based on soil test recommendations. During summer 2018, we measured vegetation structure and collected young and old forage for nutritional analysis from four commonly-occurring early successional plants: (common ragweed (Ambrosia artemisiifolia), horseweed (Conyza canadensis), Canada goldenrod (Solidago canadensis), pokeweed (Phytolacca americana), and blackberry (Rubus canadensis). The effect of amendment treatment varied based on species and nutrient, but crude protein in old goldenrod leaves was the only forage/nutrient combination that soil amendment raised to meet minimum nutrition requirements of a lactating doe with twin fawns that was not already in excess of the minimum requirement for a lactating doe. Although soil amendment failed to consistently raise most nutrient values in old-fields, it did increase average vegetation height by 71% following fertilization and 65% following fertilizer+lime application. Additionally, visual obstruction from 50-100 cm, 100-150 cm, and 150-200 cm was greater following fertilizer and fertilizer+lime applications. In fields where cover is limited because of low soil productivity, amendments can be applied to increase vegetation structure for various wildlife species.
... Even with readily digestible, high-energy forage or mast available, deer may select other food items depending on nutritional needs (Dykes et al., 2020;Provenza et al., 2003). As such, deer likely have a strong effect on oak fitness and the broader plant community by consuming acorns and seedlings of many species, especially because deer remain abundant during poor mast years (Harlow et al., 1975;Lashley et al., 2011;Steiner 1996). ...
... Third, we transplanted oak and blackgum (Nyssa sylvatica) seedlings near parent trees with and without acorn additions to test the hypothesis that masting (seed density) would generate either negative growth in oak seedlings or strengthen deer-mediated apparent competition with blackgum (i.e., oak mast seeding attracts deer, a shared predator, that suppresses relative growth of competing blackgum seedlings). We selected blackgum because this species is a preferred deer browse and a common competitor of oak seedlings in eastern forests (Iverson et al. 2008, Lashley et al. 2011, Nowacki and Abrams 2008. Finally, we characterized the local plant community near parent trees with and without acorn additions to test the hypothesis that mast seeding would elevate local herbivory pressure, thus providing a spatiotemporal niche opportunity for herbivory tolerant plants and causing understory plant communities to diverge in composition. ...
Mast seeding is a resource pulse that is thought to be a strategy in plants to satiate obligate seed predators. Behavioral responses of facultative consumers receive less attention in mast seeding studies despite evidence that they may be more important to community-level indirect effects. We designed an acorn addition experiment to test the hypothesis that mast seeding in oaks (Quercus spp.) generates indirect effects on the plant community by generating a behavioral response in white-tailed deer (Odocoileus virginianus)—the largest herbivore in the ecosystem and a facultative seed predator. Deer activity at masting trees increased substantially for months following acorn addition. Deer may require multiple dietary sources to satisfy energetic and nutritional demands and are consumers of oaks at multiple life stages. As such, acorn addition generated opposite density-dependent effects on seeds and seedlings. In acorn addition plots, we documented increased seed survival but decreased growth of oak seedlings relative to a common competitor. Importantly, acorn additions also resulted in different local plant assemblages than those at non-masting parent trees, which suggests that beta diversity may increase with localized variation in herbivory pressure. Overall, our results indicate that oaks may promote coexistence in the plant community by modulating spatiotemporal variation in herbivory pressure. These results highlight the important role of facultative consumers in linking resource pulses to the broader community.
... L.) regeneration depends on sunlight provided by canopy gaps (Hannah 1987, Brose andVan Lear 1999), and growth rates of crop trees are greater following release from competition (Wendel and Lamson 1987, Lamson et al. 1990, Kochenderfer et al. 2001. Additionally, sunlight availability influences forest understory development, and silvicultural techniques that reduce canopy coverage can improve conditions for various wildlife species (Masters et al. 1993, Mixon et al. 2009, Lashley et al. 2011, McCord et al. 2014. ...
... If trees are not merchantable because of species composition, size class, or volume, noncommercial techniques may be used to reduce canopy coverage (Nyland 2002). These techniques collectively are referred to as forest stand improvement (FSI) and can be applied to manipulate both economic and ecological conditions in a forest (Wendel and Lamson 1987, Nyland 2002, Lashley et al. 2011, McCord et al. 2014. Commonly, FSI is conducted by killing trees using herbicide introduced into the cambium (Pariona et al. 2003, Ohlson-Kiehn et al. 2006, Lewis and McCarthy 2008, McCord et al. 2014). ...
Treatment of individual trees in hardwood stands typically is conducted with herbicides that have no soil activity, such as triclopyr. However, triclopyr is not effective on some tree species. Applying a mixture of triclopyr and imazapyr would broaden the spectrum of species controlled, but nontarget mortality may be problematic as imazapyr may affect other trees through soil activity. We applied herbicide via girdle-and-spray as part of a forest stand improvement treatment in four upland hardwood stands in the Upper Coastal Plain of Alabama. We compared effects of using triclopyr alone with a mixture of triclopyr and imazapyr 18 months posttreatment. Only one untreated sweetgum (Liquidambar styraciflua) out of 440 trees was killed in the stands treated with the herbicide mixture (0.5 percent nontarget mortality rate). Nontarget mortality did not differ between treatments. However, the herbicide mixture controlled hickory (Carya spp.) and sourwood (Oxydendrum arboreum) better than triclopyr alone, with 56 percent of hickory treated with triclopyr still alive 18 months later, compared with 0 percent of hickory treated with the mixture. Our results indicate a mixture of triclopyr and imazapyr provides better control than triclopyr alone, and there is minimal risk to nontarget tree species in hardwood stands when used according to label recommendations.
Study Implications
Forest stand improvement (FSI) is a noncommercial practice typically conducted by cutting and using herbicide to kill undesirable trees. Consideration must be given to herbicide selection, which is based on efficacy on target species while minimizing nontarget mortality of residual trees. We found that a mixture of triclopyr and imazapyr applied via girdle-and-spray was more effective for FSI than triclopyr alone, which failed to control a significant proportion of treated hickory, and resulted in essentially no nontarget mortality in mixed hardwood stands 18 months after application.
... Oaks (Quercus spp.) have been a foundational species in forests of eastern North America, but have been failing to regenerate in recent decades [1,2]. Forests dominated by oaks are ecologically and economically valuable as a source of timber, as a food source and habitat for wildlife, and for relatively open stand characteristics that promote diverse understory flora [3]. These forests were historically maintained by a frequent fire regime, and current oak dominance often reflects legacies of heavy logging, burning, and grazing by early European settlers [2]. ...
... Northern red oak is highly preferred as browse for deer, relative to co-occurring woody plants in Indiana [47], and seedlings likely lacked the resources necessary to recover from browse damage where the dense midstory layer was left intact. Another perspective on this interaction between treatments is that higher light availability may have reduced browse pressure on individual oak seedlings in the midstory removal and shelterwood treatments by increasing the abundance of alternative browse, thereby making deer fencing less beneficial [3,24,48,49]. Precise estimates of white-tailed deer populations across Indiana are not currently available, but our results show that while deer may impact seedling growth more strongly in some areas than others, browse pressure on unprotected seedlings was evident throughout the region encompassed by this study. We observed the weakest effect of deer fencing on seedling height growth at the Nelson-Stokes tract. ...
Establishing adequate advanced oak reproduction prior to final overstory removal is crucial for regenerating oak forests in the eastern U.S. Many management approaches exist to this end, but benefits associated with any individual technique can depend on the suite of techniques employed and the geographic location. At four mixed-hardwood upland forest sites in central and southern Indiana, we tested factorial combinations of deer fencing, controlled-release fertilization, and various silvicultural techniques (midstory removal, crown thinning, and a shelterwood establishment cut) for promoting the growth and survival of underplanted red oak seedlings. Crown thinning resulted in slow growth and low survival. Midstory removal and the shelterwood establishment cut were nearly equally effective for promoting seedling growth. Seedling survival was strongly influenced by fencing, and differences in survival between silvicultural treatments were minimal when fencing was employed. Fertilization had minimal effects overall, only increasing the probability that unfenced seedlings were in competitive positions relative to surrounding vegetation. We suggest that underplanting oak seedlings can augment natural reproduction, but the practice should be accompanied by a combination of midstory removal and fencing, at a minimum, for adequate growth and survival.
... Across this region, forested areas are largely homogenous in structure, where a relatively uniform and dense canopy cover with negligible ground cover exists (Lashley et al., 2011). ...
... Research that creates a link between wildlife abundance, particularly highlyvalued game species such as white-tailed deer, and forest management practices has the potential to show the importance of actively managing forested patches to create desirable wildlife habitat, and public education efforts to elucidate this relationship should be emphasized. For example, in oak-dominated systems management actions that are known perpetuate oak dominance such as prescribed fire, thinning, or overstory removal may also improve wildlife habitat for some species and should be further explored (Lashley et al., 2011). ...
In the greater Central Hardwood Region, advance regeneration of oak (Quercus spp.) and
hickory (Carya spp.) has been in decline for several decades. Facilitated in part by an abrupt
change in disturbance regime, coupled with an increase in herbivore density, the existing midsuccessional,
mast-producing species are being outcompeted by late-successional, mesophytic
species. Oak-hickory forests provide keystone resources for a diverse forest wildlife community,
and a decline in its dominance will likely impact habitat use and occupancy patterns in the
mammal community, but to what extent is unclear. During May-August 2015-2016, I deployed
150 remotely-triggered camera traps in Trail of Tears State Forest (TTSF), Union County,
Illinois to investigate single-season, site occupancy patterns and detection probabilities as a
function of forest composition and structure for 3 mammals (eastern gray squirrel [Sciurus
carolinensis], raccoon [Procyon lotor], and white-tailed deer [Odocoileus virginianus]). I
collected microhabitat data at each camera-site and utilized a GIS application to estimate spatial
relationships among anthropogenic features and camera-sites. I recorded 404 photographs of 11
endothermic species during 3927 days of survey effort, with white-tailed deer, raccoons, and
eastern gray squirrels as the most detected species, respectively. Detection probability of eastern
gray squirrels was best explained by the global detection model, indicating no covariate
measured explained the variation in detection rates. Raccoon detection probability was best
described by a negative relationship with the average temperature recorded during survey period.
The best-fitting detection model for white-tailed deer indicated detection probabilities declined
ii
throughout the sampling period and across seasons. Eastern gray squirrel site occupancy models
received little support, however, ecological land type phase was the most supported model. The
best fitting habitat model described a negative relationship between eastern gray squirrel site
occupancy probability and coarse woody debris volume. For raccoons, no model with habitat
covariates was better fitting than the null model. Raccoon occupancy probability increased with
maximum DBH at a site, ground cover, and beech-maple importance values, but decreased with
oak-hickory importance values. White-tailed deer occupancy was most positively influenced by
ground cover and oak-hickory importance values, but decreased with distance to forest edge,
number of understory stems, and beech-maple importance values. My research provides
empirical evidence to predictions made regarding the impact of a decline in oak dominance
across the Central Hardwood region on a portion of the region’s mammal community. Shifts to
late-successional conditions in the Central Hardwood region will likely continue and magnify if
forest management approaches continue to minimize the frequency and occurrence of large,
anthropogenic disturbances to the forest overstory. A mosaic of forest conditions will be needed
to best support a diverse and complete mammal community across the region.
... Previous research has also investigated the response of deer, elk and moose to forest harvesting, including the influence of harvest blocks on ungulate habitat use and reproductive success (Darlington et al., 2022;Dawe, 2011;Fisher et al., 2020;Laurent et al., 2020), harvest block age and size (Fisher & Wilkinson, 2005;Lyon & Jensen, 1980;Visscher & Merrill, 2009), selective or partial harvesting methods (Nadeau Fortin et al., 2016;Vanderwel et al., 2009), response to herbicide-treated stands (Raymond et al., 1996;Thompson et al., 2003) and natural regeneration versus treated stands (Boan et al., 2011). Relationships between silviculture methods and available ungulate forage in harvest blocks (but not ungulate use of harvest blocks) have also been evaluated (Edwards et al., 2004;Lacascio et al., 1990;Lashley et al., 2011). ...
Forest harvesting causes habitat loss and alteration and can change predator–prey dynamics. In Canada, forest harvesting has shifted the distribution and abundance of ungulates (deer, elk and moose) that prefer early seral forest, resulting in unsustainable caribou predation by shared predators (bears, cougars and wolves). Long‐term solutions for caribou recovery require management to reduce ungulate prey species within caribou ranges. Silviculture practices applied after forest harvesting directly affect the amount of forage available in harvested areas, and therefore influence ungulate distribution, but few studies have completed detailed assessments on how specific treatments of site preparation, planting and stand tending influence ungulate use of harvest blocks.
We used camera traps, silviculture data, GIS‐derived habitat and disturbance data, and detailed vegetation data collected at field sites to investigate ungulate occurrence in harvest blocks in west‐central Alberta, Canada. We compared seasonal ungulate occurrence and investigated how site‐specific characteristics, the surrounding habitat and disturbance density, and fine‐scale silviculture treatments influenced ungulate occurrence in blocks.
Deer, elk and moose occurrence was higher in summer compared to winter. Elk, moose and white‐tailed deer occurrence was higher in blocks with greater availability of specific forage species. Moose occurrence was higher in blocks with a lower road density in the surrounding area, and white‐tailed deer occurrence was higher in blocks further from seismic lines and with a lower proportion of harvest blocks in the surrounding area.
Deer, elk and moose occurrence was higher in younger harvest blocks. Mule deer and white‐tailed deer occurrence was lower in blocks with higher planting densities of lodgepole pine, and mule deer occurrence was also lower in blocks that had been stand tended.
Our study provides detailed information on ungulate response to fine‐scale silviculture methods used in Alberta, directly linking wildlife occurrence to forestry practices, and providing practical scientific information to inform sustainable forestry. Translating this research into practical landscape management decisions could benefit boreal biodiversity, including threatened species like caribou, and culturally and economically important species like deer, elk and moose.
... While roe deer in forests often rest in sites with closed canopies (Mysterud, 1996), they forage frequently on sites with little canopy cover (Ewald et al., 2014). This is probably due to higher forage abundance with little canopy closure (Gill et al., 1996, Kuijper et al., 2009, Lashley et al., 2011. Thus, it appears that thermal cover provided by the canopy is of low relevance in spring and autumn, but might be important in winter (Massé and Côté, 2009). ...
Browsing damages to young trees can have lasting impacts on forest structure. Roe deer (Capreolus capreolus), the most common and widespread large herbivore in central Europe, create a vast majority of this damage. To lessen the impact, it is important to understand the relationship between roe deer and the landscape matrix, and which factors such as food availability and cover will drive the use of habitat by roe deer. In this study, we explored how small scale-food availability (5 × 5 m 2), forest structure (100 × 100 m 2) and landscape heterogeneity (500 m radius) influenced the use of habitat by roe deer in an intensively managed temperate mountainous mixed forest with implemented retention forestry practices. Using camera-trap detections of roe deer from 130 study plots in the southern Black Forest, monitored for 2.5 years, we found that local forest structure had the strongest influence on roe deer habitat use. Contrary to our expectations, landscape features, such as edge density between forest and non-forest, did not affect roe deer detections, probably because overall anthropogenic pressure is high and homogenous throughout our study system. Small-scale food availability also had little influence, which is likely due to widespread availability throughout the study area. Roe deer were also detected less where there were higher amounts of lying deadwood in autumn, indicating that retention forestry methods may have a negative impact on roe deer habitat use. Since forest structure was the strongest driver of roe deer habitat use, this study supports earlier claims that forests may be managed by affecting roe deer habitat use, thereby browsing damage intensity, through manipulation of food availability and cover.
... Deer can thrive in disturbed landscapes, which explains their recovery once hunting pressure was relieved (Michigan Department of Natural Resources, 2016). Forest-clearing is widely used today to boost populations of deer and other game species (Lashley et al., 2011;Dechen Quinn et al., 2013;Michigan Department of Natural Resources, 2017). However, high deer population densities can have significant negative effects on forest regeneration, native herbaceous plants-especially charismatic floristic groups such as orchids-and songbirds and their habitats (Alverson et al., 1988;deCalesta, 1994;Rooney and Waller, 2003;Knapp and Wiegand, 2014;Jirinec et al., 2017). ...
A campaign is underway to clear established forests and expand early-successional habitats—also called young forest, pre-forest, early seral, or open habitats—with the intention of benefitting specific species. Coordinated by federal and state wildlife agencies, and funded with public money, public land managers work closely with hunting and forestry interests, conservation organizations, land trusts, and private landowners toward this goal. While forest-clearing has become a major focus in the Northeast and Upper Great Lakes regions of the U.S., far less attention is given to protecting and recovering old-forest ecosystems, the dominant land cover in these regions before European settlement. Herein we provide a discussion of early-successional habitat programs and policies in terms of their origins, in the context of historical baselines, with respect to species’ ranges and abundance, and as they relate to carbon accumulation and ecosystem integrity. Taken together, and in the face of urgent global crises in climate, biodiversity, and human health, we conclude that public land forest and wildlife management programs must be reevaluated to balance the prioritization and funding of early-successional habitat with strong and lasting protection for old-growth and mature forests, and, going forward, must ensure far more robust, unbiased, and ongoing monitoring and evaluation.
... Controlling tree regeneration was ranked 9th out of the 9 management activities listed. Managing invasive species and making stand improvements were ranked highest and were likely perceived as important for achieving priority management objectives, including enhanced wildlife habitat and recreational hunting opportunities (DeCalesta and Stout, 1997;Lashley et al., 2011). Landowner education programs may consider using a curriculum that links wildlife habitat with oak regeneration, as a way to encourage the strategic application of fire on private lands to benefit oaks. ...
Prescribed burning is important for the ecological health of fire-dependent forests, however, there is little economic research examining landowner preferences for living with fire in the age of the Anthropocene. To understand the value of reintroducing fire on the landscape we assessed forest owner willingness to pay (WTP) for various prescribed fire programs in Pennsylvania, where natural fire occurs infrequently. Survey responses were collected from 243 forest owners using Likert scales and choice experiment questions resulting in a 44% response rate. Most respondents were classified as having limited experience with prescribed fire, but many also had low risk perceptions about prescribed fire and high trust in prescribed fire implementors. A majority (66%) elected to enroll in at least one of 16 proposed burn programs and almost a quarter of landowners were willing to pay up to $200 per acre. Using mixed logistic regression methods, mean WTP was estimated to range from $11 to $19 per acre, but varied significantly under different program alternatives. Respondents overall preferred programs that helped maintain ecosystem health and biodiversity, and offered cost-share, reduced liability, and access to burn bosses. Demographic characteristics were also important predictors of enrollment (i.e., income level, age, and involvement in assistance programs). We conclude that forest owners in Pennsylvania see prescribed fire as potentially helping them meet priority management objectives and supporting cultural values about forest stewardship. Technical and financial assistance for forest owners will be important for expanding the use of prescribed fire in Pennsylvania.
... While roe deer in forests often rest in sites with closed canopies (Mysterud, 1996), they forage frequently on sites with little canopy cover (Ewald et al., 2014). This is probably due to higher forage abundance with little canopy closure (Gill et al., 1996, Kuijper et al., 2009, Lashley et al., 2011. Thus, it appears that thermal cover provided by the canopy is of low relevance in spring and autumn, but might be important in winter (Massé and Côté, 2009). ...
... Others have reported similar findings for males (Byrne et al., 2014;Karns et al., 2012), females (Larson et al., 1978), or both sexes (Montgomery, 1963). Deer use food plots because they often provide significantly greater forage density than the surrounding forest (Edwards et al., 2004;Lashley et al., 2011). Avoidance of food plots during the day may have been related to the hunting pressure they received. ...
Resource selection in sexually dimorphic ungulates is at least partially explained by sex-specific resource requirements and risk aversion strategies. Females generally spend more time in areas with less risk and abundant, high-quality forage due to their smaller body size. However, demographically variable responses to risk are context dependent, and few have concurrently quantified male and female behavior within areas with the same resource base. We captured 111 (54 males, 57 females) adult white-tailed deer (Odocoileus virginianus) from 2009 to 2018 on a site in South Carolina, USA, where hunters were the primary source of adult mortality. We fit each deer with a GPS collar programmed to collect locations at 30-min intervals. Upon collar recovery, we analyzed the data to estimate sex- and time-specific selection for, and distance to, various cover types. While both sexes generally avoided risky areas (i.e., sites hunted more frequently) during the day, females (p = .41) were more likely than males (p = .16) to use risky areas containing abundant food resources during the day, where p = probability of selection. Our findings indicate that female white-tailed deer may be forced to utilize high risk areas during high risk periods due to their smaller body size and increased nutritional demands, whereas larger males are better able to forgo foraging opportunities during risky periods to mitigate risk; however, our study design left room for the possibility that our observations were driven by innate sex-specific patterns in white-tailed deer. Nonetheless, our study contributes information to the literature by describing sex-specific resource selection by diel period on a site where sexes shared the same resources and were presented with the same landscape of risk.
... Because fire causes a temporary nutrient pulse by increasing plant quality and biomass (Christensen 1977, Boerner 1982, Batmanian and Haridasan 1985, Singh 1993, Van de Vijver et al. 1999, Lashley et al. 2011, Eby et al. 2014, fire phenology should affect nutrient availability for herbivores which may influence plant-herbivore interactions. For example, a mismatch between nutrient availability and herbivore demands is a near certainty if fires occur before the growing season because plant resprouting co-occurs with spring green-up when the fire occurs before the onset of growing season. ...
Trophic mismatches are commonly reported across a wide array of taxa and can have important implications for species participating in the interaction. However, to date, examples of trophic mismatch have centrally focused on those induced by shifts in climate. Here we report on the potential for humans to induce trophic mismatch by shifting the phenology of fire. Globally, anthropogenic fire ignitions are phenologically mismatched to that of historic lightning ignitions but the effects of this phenological mismatch on trophic interactions are poorly understood. Using fire records from 1980 to 2016 from the southeastern USA, a hotspot of anthropogenic fire, we demonstrate that there is a temporal mismatch between anthropogenic and lightning lit fires in this region. The peak of anthropogenic ignitions (i.e. 45% during March and April) occurred 3 months earlier than the peak in lightning‐ignited fires (i.e. 44% occurred during June and July), a pattern consistent with reports from several other regions and continents. We demonstrate with a field experiment conducted at a nutrient‐poor site in the southeastern U.S., that anthropogenic fire phenology shifts nutrient pulses in resprouting plants so that they mismatch herbivore reproductive demands. Consequently, plant nutrient quality in four commonly consumed forages was below the threshold to meet lactation requirements. Neonates subsequently were more likely to starve when born far from areas burned during the peak month of lightning fire phenology. Our data indicate that human activities may be an additional causative agent of trophic mismatch.
... Nutritional carrying capacity.-We calculated estimates of nutritional carrying capacity for deer using a mixed-diet approach with nutritional constraints according to Hobbs and Swift (1985). We used a nutritional constraint of 14% crude protein with a 2.4 kg/day intake rate to represent nutritional needs of a 50-kg doe at peak lactation with twin fawns (National Research Council 2007, Hewitt 2011, Lashley et al. 2011, Nanney et al. 2018. ...
Restoration of early successional plant communities dominated by nonnative plant species is a central focus of many state and federal agencies to improve habitat for wildlife associated with these communities. Restoration efforts largely have concentrated on controlling nonnative species followed by planting native grasses and forbs. However, there are numerous establishment problems associated with planting that warrant evaluation of alternative approaches for restoration. We conducted a field experiment to compare vegetation composition and structure as related to habitat for focal wildlife among plant communities established by planting (Planted) native grasses and forbs and revegetation from the seedbank (Seedbank) without planting following control of tall fescue (Schedonorus arundinaceus) at 15 replicated sites in Tennessee and Alabama, USA. Planted and Seedbank treatments produced similar plant communities. Vegetation structure providing cover for nesting and brooding northern bobwhite (Colinus virginianus) and wild turkey (Meleagris gallopavo) was similar between Seedbank and Planted treatments except native grass cover was greatest in Planted, and we recorded greater openness at ground level in Seedbank than Planted or tall fescue control (Control). Abundance of northern bobwhite food plants and selected white‐tailed deer (Odocoileus virginianus) forage were similar between Planted and Seedbank treatments, but nutritional carrying capacity for deer was greatest in Seedbank. Despite similarities in food abundance, and even though all forbs included in the planting mixtures were food plants, the majority of food plants in Planted were from the seedbank. The compositional and structural characteristics deemed most influential in previous studies to selection of breeding sites by dickcissel (Spiza americana), field sparrow (Spizella pusilla), grasshopper sparrow (Ammodramus savannarum), Henslow's sparrow (Ammodramus henslowii), and northern bobwhite were similar in Planted and Seedbank. Tall fescue Control was most similar to characteristics of eastern meadowlark (Sturnella magna) breeding sites. Revegetation following Seedbank produced a plant community that provided habitat for many wildlife species equal to or better than Planted and was 3.7 times less expensive than Planted. © 2021 The Wildlife Society. Planting a native seed mixture following tall fescue control was not necessary in Tennessee and Alabama to establish a native early successional plant community beneficial to conservation‐priority grassland birds, northern bobwhite, wild turkey, or white‐tailed deer. Land managers should consider managing the natural seedbank response following eradication of nonnative species and spot‐treat undesirable plant species rather than routinely planting native grasses and forbs when restoring native early successional plant communities for wildlife.
... For example, implementation of dormant season prescribed fire in closed-canopy stands where most oak regeneration is suppressed has little positive impact, but these are the conditions under which most fires in the region are conducted (Brose et al. 2013). In contrast, growing season fires, especially when conducted in relatively open stands, have shown promise for reducing mesophyte competition, releasing oak reproduction, promoting an herbaceous fuel bed (Brose and Van Lear 1998, Gruchy et al. 2009, Brose et al. 2013, and providing a pulse of high-quality forage when nutritional demands of herbivores such as whitetailed deer are high (Lashley et al. 2011(Lashley et al. , 2015, which could be important for curtailing browse impacts on oak regeneration, particularly if mesophytes and oaks differ in their nutritional status. However, the potential critical importance of fire phenology and this vegetation-fire-herbivory interaction have been largely overlooked, likely because growing season fires are often avoided because of narrow burn windows (Chiodi et al. 2018) and high fuel moisture under shading from leaf cover (Harper et al. 2016). ...
Pyrophytic oak landscapes across the central and eastern United States are losing dominance as shade-tolerant, fire-sensitive, or opportunistic tree species encroach into these ecosystems in the absence of periodic, low-intensity surface fires. Mesophication, a hypothesized process initiated by intentional fire exclusion by which these encroaching species progressively create conditions favorable for their own persistence at the expense of pyrophytic species, is commonly cited as causing this structural and compositional transition. However, many questions remain regarding mesophication and its role in declining oak dominance. In the present article, we review support and key knowledge gaps for the mesophication hypothesis. We then pose avenues for future research that consider which tree species and tree traits create self-perpetuating conditions and under what conditions tree-level processes might affect forest flammability at broader scales. Our goal is to promote research that can better inform restoration and conservation of oak ecosystems experiencing structural and compositional shifts across the region.
... However, studies often consider too few nutrients to adequately test the nutrient balance hypothesis of diet selection (e.g., in white-tailed deer studies; 4 in Nelms, 1996;2 in Berteaux et al., 1998;2 in Dostaler, 2011). Moreover, even when many nutrients are considered, quantifying diet selection in wild populations is notoriously difficult and often researchers utilize indirect measures of selection such as bite counts (Lashley et al., 2011;Lashley and Harper, 2012), microhistological surveys (Alipayo et al., 1992;Marrero and Nogales, 2005;Lashley et al., 2015;Jung et al., 2015;Lashley et al., 2016), estimates of plant biomass removal (Lunceford, 1986), or captive animal observations (Nelms, 1996;Ceacero, 2009;Ceacero, 2015). However, each of those measurements have associated biases making accurate estimates difficult (Lashley et al., 2016). ...
Herbivores must navigate a heterogeneous matrix of nutrients in plant communities to meet physiological requirements. Given that the only difference between an essential nutrient and a toxin is the concentration in the herbivores diet, heterogeneity of nutrient concentrations in plant communities likely force wild herbivores to balance intake of abundant nutrients that may reach toxic levels with the need to meet nutritional demands of rare nutrients (i.e., nutrient balance hypothesis). While this hypothesis has been demonstrated in controlled studies with captive herbivores, experiments testing the nutrient balance hypothesis with wild herbivores are rare. We designed a cafeteria-style experiment to measure use of forages with differing nutritional compositions by wild white-tailed deer (Odocoileus virginianus) to test the nutrient balance hypothesis. We predicted deer diet selection would be explained by attraction to some nutrients and avoidance of others. Deer selected forages with low sulfur concentrations, a nutrient that commonly reaches toxic levels in herbivores. However, deer secondarily selected forages with greater digestibility and crude protein. Thus, our data indicate that the nutrient balance hypothesis may explain diet selection in wild herbivores where they avoid reaching toxicity of abundant nutrients while secondarily maximizing intake of limiting nutrients.
... Species-specific research is needed because wildlife species respond differentially to fire based on their ecology. For example, some species in the Appalachian region such as Turkey (Meleagris gallopavo;McCord et al., 2014), White-Tailed Deer (Odocoileus virginianus; Lashley et al., 2011), and many reptiles (e.g. Keyser et al., 2004;Greenberg et al., 2018;Hromada et al., 2018) appear to benefit from prescribed fire, but negative impacts have been documented for other species such as the Eastern Box Turtle (Terrapene carolina; Howey and Roosenburg, 2013). ...
Prescribed fire is used in the central Appalachians to promote and maintain mixed-oak and pine forests, create open forest conditions, improve habitat for wildlife, and to reduce the risk of impact of higher intensity wildfires on human development. Few studies have investigated responses of terrestrial salamander populations to habitat management using fire, and estimated responses have been neutral, negative, and positive depending on geography, species, and fire-severity. We examined woodland salamander (genus Plethodon) population responses to habitat management using prescribed fire on Shenandoah Mountain in the George Washington National Forest in West Virginia and Virginia, USA. We focused on responses of the Cow Knob Salamander (P. punctatus), a talus specialist and species of high conservation concern, but also examined responses of the Eastern Red-Backed Salamander (P. cinereus), a widespread habitat generalist. Three burn units were subjected to two low-severity burns and one unit was burned five times with ca. 40% tree mortality. Using a combination of nighttime visual encounter surveys and coverboard surveys, we compared terrestrial salamander abundance and body condition in unburned and burned areas. We also measured habitat characteristics at sampling sites to determine if prescribed burn histories were correlated with habitat conditions important to woodland salamanders. Mean abundance for P. punctatus was lower at sites that were burned, but there was not a strong burn effect for P. cinereus. Abundance of both species was positively correlated with canopy cover. Mean and median body condition index (BCI) score was higher for P. punctatus and lower for P. cinereus on the West Virginia side of Shenandoah Mountain, and lower in burned areas for both species. However, the most parsimonious BCI models did not contain the burn predictor. Management using prescribed fire altered microhabitat conditions that are important for woodland salamanders, such as canopy cover, leaf litter depth, and vegetation groundcover. Our study suggests that woodland salamanders in the central Appalachians can persist in forests managed using prescribed fire, but also indicates that prescribed fire can result in reduced habitat quality for some woodland salamander species.
... Research that creates a link between wildlife abundance and diversity and forest management practices has the potential to show the importance of actively managing forested patches, and emphasizing public education efforts to elucidate this relationship should be included in land management strategies [65]. For example, in oak-dominated systems management actions that are known perpetuate oak dominance, such as prescribed fire, thinning, or overstory removal, may also improve wildlife habitat for some species and should be further explored [66]. Future research should work to provide year-round comparisons of habitat use among forest stands differing in composition, which may offer additional insight into the implications of shifting forest composition across the region [67]. ...
Oak-hickory (Quercus-Carya spp.) forest types are widespread across the midwestern United States, but changes in forest disturbance regimes are resulting in little to no oak recruitment and a compositional shift to shade-tolerant, mesophytic species, such as American beech (Fagus grandifolia) and sugar maple (Acer saccharum). We conducted camera trap surveys in a mature upland hardwood forest of southern Illinois, USA during May to August 2015–2016 to document mammal summer habitat use in relation to forest structure and composition to further understand how regional shifts in forests may affect mammal communities. With nearly 4000 camera days of effort, we modeled occupancy patterns for white-tailed deer (Odocoileus virginianus), raccoon (Procyon lotor), and eastern gray squirrel (Sciurus canadensis). Forest composition models outcompeted forest structure models for white-tailed deer, where we observed a statistically significant negative relationship between white-tailed deer habitat use and beech dominance. Further, we found a strong, positive association between deer and oak dominance. Model selection indicated little support for within-stand forest structure or composition characteristics influencing habitat use for raccoons. Eastern gray squirrel occurrence was best described by forest composition, revealing a positive relationship with beech–maple importance values. Our predictive models indicated that the impact of forest changes underway will have varying impacts on wildlife species. We can expect changes in habitat use patterns to be more pronounced with time barring revised forest management practices, and these changes are likely to be most influential at the landscape-scale. We conclude that a patchwork mosaic of forest conditions will likely best support a diverse and abundant mammal community across the region.
... All plots had two 1 m 2 exclosures constructed from 5 cm mesh poultry wire placed randomly within the plots to evaluate the impacts of herbivory on growth (Lashley et al. 2011 monthly and full-season dry biomass production, and stem density. All experimental plots were evaluated, with each sown plot evaluated for the species planted and control plots evaluated for naturally growing species. ...
Reservoirs throughout the country exhibit degraded shorelines and simplified
littoral habitats because of aging. This study evaluated the establishment of agricultural
plantings on reservoir mudflats and the effectiveness of supplemental brush pile
structures in providing recruitment habitat for juvenile fish. The mudflats of Enid
Reservoir, Mississippi were seeded during the winter drawdown in October of 2016 and
2017 with agricultural plantings. Monitoring of plantings found grasses performed best in
terms of establishment and providing potential fish habitat. During Summer 2017 brush
piles and control sites, without brush, were sampled in Enid Reservoir using rotenone to
evaluate juvenile fish use. Juvenile fish exhibited greater abundances and larger sizes, on
average, in brush pile sites. Larger brush piles placed in shallower water provided the
greatest benefit to juvenile fish.
... Climatic variation can also induce cohort-specific changes to phenotype and reproductive success through its effect on plant productivity and phenology (Behrenfeld et al. 2001, Stenseth et al. 2003, Parker et al. 2009). Forage production and quality has been associated with growing season rainfall in moist, temperate ecosystems (Shiflet and Dietz 1974, Lashley et al. 2011, Lashley and Harper 2012. Summer precipitation in the southeastern United States has become more variable in recent years (Wang et al. 2010), with even greater variability projected as anthropogenic forces alter the behavior of the North Atlantic subtropical high (Li et al. 2011). ...
Effective deer management requires managers to distinguish between the density‐dependent influence of harvest and local environmental factors. The Batture region of the Lower Mississippi River Valley comprises land adjacent to the river that is not protected by the levee system, and is therefore subject to seasonal flooding with potential to influence the morphology and demographics of white‐tailed deer (Odocoileus virginianus). Using harvest records of 42,954 females and 3,588 males from 61 Batture properties in Mississippi and Louisiana, we created linear regression models of deer body development and recruitment indices to compare the influence of seasonal flooding, harvest rate, growing season weather, and agronomic forage availability during 1988–2016. Overall, deer in the Batture appeared to be more influenced by extrinsic factors than by harvest. Seasonal flooding appeared in every model and generally had stronger effects than weather or harvest variables. Flooding from 1 to 2 years prior, regardless of season, was correlated with greater female body mass, lactation rates, and antler mass of trophy males, possibly reflecting silt deposition effects on soil fertility and promotion of new understory forages. Conversely, current‐year flooding effects were invariably negative, implying direct effects of displacement. Summer flooding was concentrated during late gestation and peak parturition periods, and exhibited the potential to reduce fall lactation rates by 18%. Harvest rates correlated negatively with female body mass and had no correlation with lactation or antler mass. We detected contrasting long‐term trends of decreasing body mass and increasing harvest rate that may reflect deteriorating habitat. Similar to flooding, increased temperatures and rainfall had negative effects for the current year, whereas increased temperatures had positive effects when occurring in the previous year. Surprisingly, annual variation in the amount of soybeans planted appeared in one model only, exhibiting a small positive effect on antler mass. We hypothesize that extensive planting of soybeans in levee‐protected lands just outside the Batture maintained substantial soybean availability despite variation in the amount planted. Given the dominating influence of flooding and weather on deer physical and reproductive parameters in the Batture, these extrinsic variables should be incorporated into the interpretation of harvest data. The common practice of curtailing harvest, particularly female harvest, following years with extensive flooding is likely counterproductive unless intensive flooding occurs during summer.
... Numerous factors constitute forage quality, including palatability and digestibility, but forage quality is typically most often referred to in terms of nutrient content. Although protein and energy are both important during antler growth, gestation, and lactation, protein is generally more limited in terrestrial ecosystems and is required to a greater degree than energy for these processes for deer in southern climates (White 1993, Asleson et al. 1996, Barboza and Parker 2008, Lashley et al. 2011. Crude protein requirements for antler growth and lactation are over 1.5 and 2.5 times that of maintenance requirements, respectively (Holter et al. 1979, Verme and Ullrey 1984, Asleson et al. 1996. ...
Prescribed fire is a cost-effective habitat management tool in pine stands to enhance the quantity and quality of forage available for white-tailed deer (Odocoileus virginianus). Management recommendations typically suggest a 3- to 5-year burn rotation in mixed pine–hardwood stands to increase quality forage production, but as fire frequency increases, forb and legume biomass increases, and woody browse decreases. A more frequent burn rotation may be a viable management option for deer managers, but there is still a lack of information regarding preferred forage and nutritional carrying capacity response to prescribed fire at these intervals. We measured the production and nutritional quality of forage within mature pine–hardwood stands after a 1- or 2-year fire-return interval during three nutritionally stressful periods for deer on a 640-acre (259-hectare) enclosure located in east-central Alabama during 2014 and 2015. These stands had previously been burned annually for over 15 years, resulting in an abundance of herbaceous vegetation. We then compared forage class biomass, nutritional carrying capacity estimates, and digestible protein between burn treatments. A 1-year fire return interval improved habitat quality to a greater degree than a 2-year fire return interval by increasing the production of forage able to support greater nutritional planes. An annual burn rotation is an effective option for managers to increase protein availability in pine–hardwood stands, but other factors such as decreased cover availability and soft mast production should also be considered.
... Despite the variability, our observed negative relation between the abundance of natives and non-natives indicates that a high abundance of established native plants may mechanistically decrease the invasibility of non-natives, in spite of species richness (Corbin and D'Antonio 2004, Davies and Johnson 2017). As reported in other studies in the Southeast, combining canopy reductions with fire can lead to large increases in native understory vegetation abundance (Masters et al. 1993, Phillips and Waldrop 2008, Lashley et al. 2011) and, thus, is likely to promote resistance. In contrast, herbicide applications were intended to kill existing understory woody vegetation, even if it was native, which may have increased the invasibility of non-native herbaceous species. ...
The biotic resistance hypothesis (BRH) was proposed to explain why intermediate disturbances lead to greater resistance to non-native invasions proposing communities that are more diverse provide greater resistance. However, several empirical data sets have rejected the BRH because native and non-native species richness often have a positive relation. We tested the BRH in a mature loblolly pine (Pinus taeda) forest with a gradient of disturbance intensities including canopy reduction, canopy reduction + fire, and canopy reduction + herbicide and fire. We analyzed data from the study using a combination of Pearson’s correlation and beta regressions. Using species richness, we too would reject BRH because of a positive correlation in species richness between native and non-native plants. However, native species abundance was greatest, and non-native species abundance was lowest following intermediate disturbances. Further, native and non-native species abundances were negatively correlated in a quadratic relation across disturbance intensities, suggesting that native species abundance, rather than richness, may be the mechanism of resistance to non-native invasions. We propose that native species abundance regulates resistance to non-native invasions and that intermediate disturbances provide the greatest resistance because they promote the greatest native species abundance.
... Releasing the crowns of white oak trees can be an effective method to increase acorn production as well as increase forage and cover for various wildlife species (Lashley et al., 2011, McCord et al., 2014. ...
Forest management practices that influence mast production in oaks (Quercus spp.) are ecologically and economically important for regeneration of future oak forests, timber products, and wildlife that consume acorns. We conducted a 10-year experiment in upland oak-hickory forests of eastern Tennessee to determine the influence of canopy release, fertilization (addition of nitrogen, phosphorus, and potassium), and their Combined influence on white oak (Quercus alba) acorn production, acorn size and quality, and acorn depredation. We used a robust before-after-control-impact design where we collected pre-treatment acorn production (acorns/m2 of crown) data from 120 white oaks for 5 years, applied canopy release and fertilizer treatments and then monitored
post-treatment acorn production on the same trees for an additional 5 years. Acorn production was temporally variable with 6 of 10 years being near complete mast failures (≤3.67 ± 8.52 acorns/m2 of crown). Also, production varied greatly among individual trees with 11% of trees classified as excellent producers accounting for 31% of all acorns produced, and 41% of trees classified as poor producers accounting for only 17%
of all acorns produced. Canopy-released and canopy-released-and-fertilized trees increased acorn production 65% and 47%, respectively, following treatment relative to control trees, with effects greatest in trees classified as poor producers. Fertilization did not influence acorn production or size and did not consistently influence acorn quality. Furthermore, acorn depredation rates did not differ among treatments. Our results indicate crown release is an important management practice when management objectives include increasing white oak acorn production in closed-canopy conditions, whereas fertilization does not influence acorn production.
... The primary purpose of this study was to maximize survival of seedlings, and thus browse protection was implemented on the most seedlings in anticipation of herbivory causing poor survival. Bud caps were recommended to protect buds from browse damage (Ward et al. 2000, Ward andMervosh 2008), whereas exclusion cages were used to exclude deer from plants within a small area (Lashley et al. 2011(Lashley et al. , 2015. ...
Seedling survival and growth in eastern white pines (Pinus strobus) may be limited by white-tailed deer (Odocoileus virginianus) browsing. However, most studies have occurred in areas central to the white pine range, making other factors such as seedling microenvironment unimportant. If microenvironment becomes a concern near the edge of the white pine range, then factors such as seedling placement in relation to forest openings could be important, especially given that deer herbivory tends to be most intense near forest edges. We evaluated the relative importance of deer browse and seedling position in openings on seedling survival and growth in central North Carolina at the southern edge of the white pine range. Further, we determined if bud caps and caging improved survival and growth. Seedlings <10m from the edge survived at a greater proportion than those >10m from the edge (83% and 73%, respectively). Initial height was the most important predictor of survival (R2= 0.55; p < 0.01). When controlling for initial seedling height, the location of the seedling (p < 0.01) within the opening was the only significant predictor of survival, despite the increase of browse near the edges of openings on unprotected seedlings. Caging and bud caps decreased seedling browse by 80% but had no effect on subsequent seedling survival (p = 0.28). A smaller proportion of seedlings with bud caps survived - an effect exacerbated by being internal to the opening. Our data indicate seedling microenvironment is an important consideration at the periphery of the white pine range.
... We based deer NCC estimates on a comparative forage quality and quantity index calculated via a constraints algorithm (Hobbs and Swift 1985). Therefore, we made no assumptions that the index obtained reflects absolute estimates of deer NCC but only relative estimates useful for comparing treatment effects (Hobbs and Swift 1985, Stewart et al. 2000, Lashley et al. 2011. Adult deer require 4À12% crude protein diets for general metabolic maintenance, whereas a lactating female nursing 1 fawn requires a 14% crude protein diet during late summer (Edwards et al. 2004, Jones et al. 2009a. ...
Switchgrass (Panicum virgatum) is a cellulosic feedstock for alternative energy production that can be grown between rows of planted pines (Pinus spp.) within intensively managed forests. Southeastern planted pine occupies 15.8 million ha and thus, switchgrass intercropping could have far-ranging effects on plant communities and biomass production within these forests if broadly implemented. Such intercropping could lead to alterations to plant communities that may cause bottom-up ecological changes affecting ecologically, economically, and socially important wildlife, such as white-tailed deer (Odocoileus virginianus; deer). Therefore, we tested whether intercropping switchgrass in loblolly pine (P. taeda) plantations would cause switchgrass to dominate vegetative biomass thereby decreasing biomass of forages and reducing white-tailed deer nutritional carrying capacity (NCC), or whether disturbance associated with establishment and harvest of switchgrass would facilitate ruderal forbs, and thereby increase biomass of preferred deer forages and increase NCC. In a randomized complete block design, we assigned 2 treatments (intercropped switchgrass and a standard pine management control) to 4 replicates of 10-ha experimental units in Kemper County, Mississippi during summers of 2011-2015. We detected 323 plant species. Intercropping switchgrass had little effect on plant biomass production, and did not affect white-tailed deer NCC at a maintenance diet of 6% crude protein. Intercropping provided additional disturbance allowing high-protein content ruderal plants to colonize, and temporarily increased (3 yr) deer NCC at the 14% crude protein diet considered necessary to support lactation. However, concomitant with a sharp increase in switchgrass biomass in the third year of the study, NCC dropped to levels similar to traditionally managed pine stands. Switchgrass intercropping is not a reliable means of increasing deer NCC as a management strategy but does not appear to reduce carrying capacity in the short-term relative to standard intensive pine management.
... Increased herbaceous development following fire occurs as a result of release from woody competition, reductions in litter (Lashley et al., 2011), and increases in soil nutrient availability (Scharenbroch et al., 2012). In our study, fire increased herbaceous cover, richness, and diversity. ...
... Following prescribed burning, wildlife forage may be more prolific, nutritious, and palatable (Harris and Covington 1983;Masters et al. 1993). For example, post-fire nutrition in the diets of ungulates has been shown to be elevated for bighorn sheep and mule deer (Hobbs and Spowart 1984) and white-tailed deer (Lashley et al. 2011). Similarly, prescribed (Penman and Penman 2010) and encourage the growth of seed-producing herbaceous plants, which in turn may increase wildlife forage availability. ...
... Additionally, we sampled an uncaged plot at a randomly generated distance (10 m to 100 m) and bearing (0 degrees to 360 degrees) from the original location of the cage. The uncaged plot was kept in the same vegetation type with the same years-since-fire; we replaced the cages in a new area and repeated the sampling protocol in 2012 (i.e., 160 caged and 160 uncaged plots per year; Lashley et al. 2011). We separated samples by species, bagged them in small paper bags, and dried them in an air-flow dryer at 50 °C (Lashley et al. 2014b). ...
Prescribed fire is commonly used to
restore and maintain the longleaf pine
(Pinus palustris Mill.) ecosystem
(LLPE). A key function of the LLPE
is the provisioning of food for wildlife.
Despite the plethora of literature
evaluating the effects of fire season
and fire-return interval on plant community
dynamics, little attention has
been given to the response of wildlife
foods to fire season or fire-return interval.
We measured the availability
of key wildlife foods (fleshy fruit [i.e.,
seed containing a nutritious pericarp]
and understory plant biomass) in upland
pine forest following dormant-
season (December–February)
and growing-season (April–June) fires
in a chronosequential design. Also,
we quantified the relative contributions
of the upland hardwood and bottomland
hardwood forest types, which
often are intentionally suppressed in
the LLPE. In 2011 and 2012, we
measured understory leafy biomass,biomass of forages selected by whitetailed
deer (Odocoileus virginianus
Zimm.), and soft mast production
chronosequentially in relation to
years-since-fire, fire season, and vegetation
type in the LLPE at Fort Bragg
Military Installation, North Carolina,
USA. Understory leafy biomass increased
in upland pine and hardwood
forests as years-since-fire increased
until two years post fire. Selected forages
decreased in upland pine forest
and increased in upland hardwood
forest as time-since-fire increased. In
upland pine forests burned during the
growing season, 94 % of the fruit was
detected two years after fire, 6 % one
year after fire, and 0 % the same year
as fire. In June, fruit density was
greatest in bottomland hardwood forest;
in July, fruit density was greatest
in dormant-season burned upland pine
forest; in August, fruit density was
greatest in upland hardwood forest;
and in September, fruit density was
greatest in upland hardwood and bottomland
hardwood forest. Overall
summer fruit density (i.e., the sum of
fruit density detected each month)
was greatest in upland hardwood forest.
Understory leafy biomass and
deer-selected forages were stable in
bottomland hardwood forest because
they were not burned, thereby providing
a relatively high and stable availability
from year to year. Our data
demonstrate the importance of diversity
in fire season and frequency, and
diversity in vegetation types to promote
key wildlife foods in the LLPE.
... Alternatively, female deer may be positively affected by growingseason fire because young, regenerating plant growth stimulated by fire is more palatable and higher in nutrients than older plant tissue (Jones and Case, 1990;Leigh et al., 1991;Lewis et al., 1982;Wood, 1988). Furthermore, fire can increase forage availability for several growing seasons after the fire (Edwards et al., 2004;Lashley et al., 2011;Masters et al., 1993Masters et al., , 1996. ...
Fields dominated by nonnative grasses, such as tall fescue (Schedonorus arundinaceus), are being restored to native plant communities across the eastern U.S. Upon restoration, disturbance is necessary to maintain native communities in an early seral stage, and plant response to different management practices is of interest to managers to guide habitat enhancement for various wildlife species. We evaluated effects of burning and mowing following restoration of native plant communities via 2 methods (planting native grasses and forbs and seedbank response without planting), across 11 replicated sites in Tennessee and Alabama, 2019–2020. We compared vegetation composition and structure, openness at ground level, forage availability, and nutritional carrying capacity (NCC) following 4 treatments (Seedbank Burned, Seedbank Mowed, Planted Burned, Planted Mowed, and tall fescue Control), and we related these measurements to the food and cover requirements for 3 popular game species: white‐tailed deer (Odocoileus virginianus), wild turkey (Meleagris gallopavo), and northern bobwhite (Colinus virginianus). The combination of planting and mowing increased grass coverage, whereas units that were established via seedbank response and managed by burning had greater forb coverage. Visual obstruction above 25 cm was greater in all treatments than Control, which provided cover that has been described as selected nesting cover by wild turkey and bedding cover for deer. Openness at ground level, which is especially important for bobwhite and wild turkey broods, was 30% greater in Seedbank Burned than Planted Mowed where we recorded the least openness among treatments by the second year of treatment. Similarly, coverage of bobwhite food and deer forage plant species was greatest in Seedbank Burned. Biomass (kg/ha) of plant species known to be selected as forage by deer was greater in all treatments than in Control, and NCC (deer days/ha) was greatest in Seedbank Burned by year 2. Our results highlight differences in plant composition and structure following management of early successional communities. Where possible, we encourage managers to use fire instead of mowing to maintain plant communities, especially if food plants and enhanced structure at multiple levels for bobwhite, wild turkey, and deer are management objectives. Furthermore, our results illustrate planting native grasses and forbs is not necessary to restore native early successional plant communities on most sites. Following 2 years of evaluating the effects of burning and mowing on vegetation composition and structure, we recommend managers use fire over mowing to set back succession and maintain early seral‐stage communities, especially if increased food for bobwhite and deer and enhanced structure at multiple levels for deer, wild turkey, and bobwhite are management objectives. We recommend selective spot‐spray herbicide applications to reduce undesirable plant species and encourage colonization of additional desirable species, and we encourage managers to consider using seedbank response and natural colonization of plants instead of planting native grasses and forbs to more effectively and efficiently promote and enhance habitat for bobwhite, wild turkey, and deer.
Prescribed burns and understory thinnings are forest management practices aimed at reducing fuel loads to lessen wildfire threat in the Southern Appalachians, USA. Spiders play a critical role in forest ecosystems by controlling insect populations and providing an important food source for vertebrates. We used pitfall and colored pan traps to investigate how abundance, species richness, and diversity of spiders differed among three fuel reduction treatments administered repeatedly over a 15-year period and untreated controls. Additionally, we examined how spiders responded to one round (before and after) of fuel reduction treatments. We established treatments within the 15-year period as follows: mechanical understory removal (twice; M), prescribed burning (four times; B), mechanical understory removal followed one year later by high-severity prescribed burns and three subsequent burns (MB), and untreated controls (C). Our study period (2014–2016) occurred after multiple prescribed burns and two rounds of mechanical understory removal had occurred. Salticidae and Lycosidae were the two most commonly collected spider families in Southern Appalachian hardwood forests. Generally, we found increased spider abundances within all fuel-reduction treatments compared to controls. Individual spider families and species showed variable responses to treatments, but abundance of several spider families was greater in one or more fuel-reduction treatments than in controls. Additionally, abundance of several spider families and hunting/web building guilds (webs built for hunting purposes or defense) exhibited yearly differences to the last round of fuel-reduction treatments. Overall, our results suggest that changes in the overstory and understory of a forest are important drivers of regional spider abundance and assemblages, and forest management practices that modify forest structure can dramatically alter spider abundance and richness, usually in a positive manner.
The ecotone between the Great Plains and the Eastern Deciduous Forest region is characterized by transitional grassland-forest ecosystems with a robust history of frequent fire regimes and fire-adapted natural communities. Historically, fires created a mosaic of prairies, savannas, woodlands, and forests juxtaposed by landscape controls. Humans have been strong determinants of fire regimes, causing frequent fires in historical times and an extended period of fire exclusion for nearly the last century. In recent decades, interest has increased in understanding the region’s fire ecology and management. This interest is driven by management objectives to promote and maintain plant and animal diversity, restore ecological processes, and increase ecosystem resilience. Plant species in the region exhibit adaptation to frequent fire regimes and wildlife species are associated with habitats maintained by fire. However, exclusion of fire over the past century has left a long-lasting mark on ecosystems by changing ecosystem structures and compositions and, in some cases, by eliminating fire-adapted natural communities. In the future, the rise of campaigns that promote appropriate fire uses will be contingent upon science, demonstrated management successes, public perspectives, and the broader challenges associated with global changes.
The role of fire in the eastern broadleaf and Appalachian forest regions, until recently, was poorly understood or minimally examined, as this region was long overlooked as a flammable landscape and fire was seen primarily as a threat to the timber resource and wildlife. In the past few decades, a significant body of research has enhanced our understanding of fire and its effects. We now recognize that fire has strongly shaped many ecosystems of this region along complex geomorphological gradients, and that returning fire, or its absence, has significant consequences for forest structure, species composition, and ecosystem function. This chapter synthesizes the state of knowledge regarding the prehistoric and historical roles of fire in these ecosystems; presents research demonstrating the effects of contemporary prescribed fire and wildfire on forest structure, species composition, and consequences for wildlife; examines evidence for shifting flammability of these ecosystems; and discusses the implications for both fire management and ecosystem sustainability in the twenty-first century.
Forests typically have limited high quality nutrition available to meet the seasonal needs of white-tailed deer (Odocoileus virginianus) during summer. Prescribed fire is used to enhance deer habitat in pyric-disclimax communities but is normally applied during the dormant season. Historically, lightning-ignited fires in pine systems occurred during summer in the southeastern US, but fire ignited by humans occurred at all times of the year and was most common in fall/winter. The different timing of prescribed fire may be important when trying to couple forage resources with nutritionally demanding periods for herbivores. We evaluated fire-timing effects (dormant season, March; growing season, June; and unburned control) on nutrient availability (crude protein, % phosphorus, and biomass) and nutritional carrying capacity (NCC) for moderate-high use forages of white-tailed deer during summer of 2018 and 2019 in nine mid-rotation loblolly pine stands. Fire increased nutrient content the year of the burn and the increase persisted into the second growing season following growing season fire. Dormant season fire increased protein- and phosphorus- based NCC for two growing seasons but growing season fire reduced biomass during the first year and delayed NCC improvement until the second year. Prescribed fire can be timed to manipulate forage quality, quantity, and improve NCC to recouple forage nutrient supply with herbivore seasonal nutrient demands. We recommend land managers consider diversifying fire timing across the landscape in pyric disclimax systems to support the changing nutrient requirements of large herbivores.
Commercial thinning and prescribed fire can improve habitat quality for white-tailed deer (Odocoileus virginianus) in lobolly pine (Pinus taeda) stands by increasing coverage of forage plants. However, the relationships among thinning intensity, prescribed fire, and deer forage have not been quantified. We estimated percent cover of deer forage plants in 5 loblolly pine stands thinned to 11 (low), 14 (medium), and 18 (high) m2/ha basal areas during 2017 in Georgia, USA. We applied prescribed fire during 2018. From years 1-2 post-treatment, cover of total deer forage increased 26% and 29% in the low and medium basal area treatments, respectively, compared to 19% in the high basal area treatment. Similarly, the increase in forb coverage was greater for the medium (13%) and low (11%) basal area treatments than the high (6%) basal area treatment. Increases in vine and bramble coverage were greater in unburned medium basal area units. Woody browse was not affected by any treatment. Our results suggest thinning loblolly pine stands to 14 m2/ha can increase coverage of deer forage plants during the first two growing seasons post-thin, but deer forage was not greater in stands thinned to <14 m2/ha two years post-thin.
Mammal distribution and diversity is quickly changing as humans modify the landscape. In particular, silviculture, which is the practice of controlling the growth, structure, and quality of forests to meet the needs of society and the landowner, influences the habitat usage of mammals. Utilizing camera traps, I monitored shifts in mammal communities across different silviculture treatments in the northern hardwood forests of the Great Lakes region in North America. I assessed the community composition across six canopy treatments and three understory treatments with a total of 2,018 active camera trap nights with 3,321 detections over the course of 147 days. For canopy treatments, high canopy cover shelterwood had the largest positive influence of mammal detection while clearcut showed a negative influence of mammal detection. For understory treatments, artificial tip-up and scarification had higher mammal detection compared to control. Within areas with a history of disturbance it may be beneficial to the mammal communities to include small disturbances, such as those created by silviculture treatments, as local species are likely disturbance-adapted. Camera traps alone may miss part of the mammal community. To monitor a full community, other techniques need to be considered, such as invertebrate derived DNA (iDNA). iDNA, is emerging as a novel tool which utilizes genomic technologies to monitor and assess mammal communities. Some invertebrates ingest their host’s DNA as they feed, which then allows researchers to extract the host’s DNA and sequence it. By doing so, the researchers can then create a more complete image of mammal community compositions. This technique has been widely used in tropical zones to monitor mammal community compositions; however, it can be adapted to be used in temperate zones by utilizing ticks and mosquitoes. To adapt this technique, one must understand the environmental influences on invertebrate collection. I investigated the environmental influences on mosquito collection success by running linear regression models. Through running the linear regression models, I found that the canopy cover and time of the month had the largest influence on the collection of female mosquitoes, while tick collection was possibly influenced by the harshness of the winter before.
Citizen science is increasingly being used by natural resource professionals to monitor the environment at large spatial or temporal extents. Developing, initiating, and sustaining an effective citizen‐science program requires careful planning, marketing, and engagement. We describe the development, implementation, and evaluation of a citizen‐science program for quantifying mast production (called MI‐MAST: Wildlife Food Tracker), and offer recommendations on marketing, recruitment of volunteers, and ways to sustain programs. In 2014, MI‐MAST was introduced to approximately 150 potential participants to test technology and capabilities of volunteers. In 2015, we worked with project partners to market MI‐MAST to approximately 500 additional potential participants throughout Michigan, USA. From 2014 to 2017, 510 people registered for a MI‐MAST account, but only 57 people submitted data. On average, each person that submitted data entered 4 instances of mast production. We randomly field‐validated 60 mast observations submitted by 15 different participants throughout the study region and found that 100% of the plant species were correctly identified and 97% of the records describing mast amounts were accurate. The majority (61%) of registered users listed hunting as a primary outdoor interest. Interactions with participants revealed that using MI‐MAST to evaluate mast production on their own property was a primary attraction for enrollment and participation. We recommend that effective citizen‐science programs are marketed to provide personal benefits and tangible outputs to individual users. We found the biggest challenges facing a natural resource agency implementing a citizen‐science program to be recruitment and retention of participants across spatial and temporal extents that are ecologically meaningful. © 2018 The Wildlife Society.
Since elk (Cervus elaphus) were extirpated from eastern North America, they have been successfully reintroduced in numerous eastern states and one Canadian province. In forest-dominated areas, reintroductions of large ungulates can create potential for conflict with humans if elk select agriculture or private property to meet nutritional needs. We determined the seasonal diet selection of recently reintroduced elk in the Missouri Ozarks by comparing diet composition with availability of 12 forage classes. We measured diet composition using microhistological analysis of feces collected from June 2011 through February 2013, and we determined forage availability through vegetation sampling. Elk selected grains and cool-season grasses over all other forage classes across all seasons and years except during summer 2012 and fall 2012. Legumes were the most highly consumed forage class for all seasons and years. Native forbs were selected during summer and fall 2012 but were not selected during 2011. Approximately half (44.6%) of the annual elk diet was comprised of species cultivated in managed forage openings. Clover (Trifolium spp.), alfalfa (Medicago sativa), and common lespedeza (Kummerowia spp.) were the most prevalent species in the elk diet, indicating the importance of managed forage openings for the Missouri elk herd. However, during a drought in 2012 elk diet shifted to native species, likely because elk became more selective of forages that met their nutritional needs. In forest-dominated landscapes, where conflict with humans on privately owned land is possible, we recommend the creation of forage openings composed of cool-season grasses and legumes to increase forage availability for elk. We recommend management for natural open lands that promote native forbs and grasses given they provide a valuable food source when environmental conditions affect availability, nutritional quality, or palatability of cultivated forages and because cultivated forage openings are not always an option in some areas or over long time periods.
Prescribed burning has the potential to improve habitat for species that depend on pyric ecosystems or other early successional vegetation types. For species that occupy diverse plant communities over the extent of their range, response to disturbances such as fire might vary based on post-disturbance vegetation dynamics among plant communities. Although responses of snowshoe hares (Lepus americanus) to fire have been studied in conifer-dominated forests in northern parts of the species’ range, there is a lack of information on snowshoe hare habitat use in fire-dependent communities in southern parts of their range. We used global positioning system (GPS) and very high frequency (VHF) radio-collars to monitor the habitat use of 32 snowshoe hares in a scrub-oak (Quercus ilicifolia)-pitch pine (Pinus rigida) barrens complex in northeastern Pennsylvania where prescribed fire has been used for habitat restoration. The area contained stands that underwent prescribed burning 1–6 years prior to our study. Also, we investigated fine-scale determinants of habitat use within stands. We found that regardless of season, hares did not select for areas that had been burned within 6 years prior. Hares primarily used stands of older scrub oak, conifer, or hardwoods, which contained dense understory vegetation and canopy cover. Hare habitat use also was positively associated with stand edges. Our results suggest that hares do not respond to prescribed burning of scrub oak in the short-term. In addition, by focusing on structural determinants of habitat use, rather than broad-scale characteristics such as stand type, management strategies for snowshoe hares can be adapted over the extent of their range despite the multitude of different land cover types across which the species occurs. © 2017 The Wildlife Society.
Outline: Deer in the eastern oak forests prior to and soon after European settlement. Understanding density and carrying capacity. Carrying capacity for diversity. Carrying capacity for tree regeneration. Carrying capacity for maximum sustained yield of deer. Crash. Is there a problem? Is it deer? Managing deer impact. Reducing deer density. Protecting understory vegetation. Counteracting effects of prolonged high deer density on understory vegetation. Selecting a management strategy. Monitoring: have you solved the problem? Further suggested reading.
Management of forested habitats for wild turkeys (Meleagris gallapavo) is critical to provide the proper
vegetative requirements to improve reproductive success and survival. We investigated the effects of prescribed fire, shelterwood harvest, and wildlife retention cuts on wild turkey habitat in a mixed hardwood forest in eastern Tennessee. Vegetative response, soft and hard mast production, invertebrate availability, and crown expansion were measured. Shelterwood harvest (61–80% density), wildlife retention cut with prescribed fire (61–80%), and wild-life retention cut (41–60%) increased the density of understory vegetation. White oak (Quercus alba) crown expansion was greater in the shelterwood harvest (25%) and wildlife retention cut (8%) than in the control. No effects of treatments were found on herbaceous cover, acorn production, soft mast production, or invertebrate densities. Shelterwood harvest, wildlife retention cut, and wildlife retention cut with prescribed burning enhanced the nesting and brooding cover for wild turkey in the short term; however, additional treatment is needed to increase herbaceous cover and reduce density of woody understory vegetation.
Offers an algorithm for estimating supportable densities of herbivores consuming diets at different levels of nutritional quality. This procedure is used to predict carrying capacity of burned and unburned mountain shrub habitat for mule deer Odocoileus hemionus and mountain sheep Ovis canadensis in Colorado. Unburned areas had higher carrying capacities for animals consuming low quality diets, but burned areas could support more animals on a high plane of nutrition. Traditional procedures for estimating carrying capacity failed to detect these interactions between carrying capacity and animal nutritional status. Forage amount and quality should be treated as integrated rather than distinct features of habitat.-from Authors
Forage production for white-tailed deer Odocoileus virginianus is often limited in closed-canopy forests. We measured browse production and nutritional carrying capacity after prescribed burning and understory fertilization in closed-canopy hardwood stands one growing season after treatment in two physiographic regions of Tennessee. Nutritional carrying capacity estimates for prescribed burning, fertilization, and prescribed burning with fertilization were greater than in controls on the Cumberland Plateau. However, the cost per pound of forage produced after fertilization exceeded US$26. In the Coastal Plain, fertilization did not affect nutritional carrying capacity, and prescribed burning and prescribed burning with fertilization lowered nutritional carrying capacity from controls. At both sites, prescribed fire had minimal effect on soil pH or soil phosphate and potash levels. Our results suggest prescribed fire and fertilization are of limited utility for increasing browse production in closed-canopy hardwood forests.
I do not have copies of Silviculture: Concepts and Applications to distribute. Waveland has exclusive rights to publish the book. It offers a traditional print copy, or you can get an electronic version at a reduced cost. You can secure either version of the book directly from Waveland Press ( Waveland.com ), or through any online or local book seller. Make certain to specify the 3rd edition, published in 2016.
Ralph D. Nyland
ABSTRACT Mid-rotation management practices for pine (Pinus spp.) plantations enrolled in cost-share programs have not been widely evaluated for wildlife. Mid-rotation pine plantations often have a substantial hardwood mid-story that limits growth of desirable understory forage species important to white-tailed deer (Odocoileus virginianus; deer). We treated with imazapyr herbicide and prescribed burning (HB) 11 thinned, 13–22-year-old loblolly pine (P. taeda) plantations in the Upper Coastal Plain (UCP; n = 5) and the Lower Coastal Plain (LCP; n = 6) of Mississippi, USA, enrolled in cost-share programs. We then sampled these plantations for production of important deer forages during July of 2003 and 2004, years 1 and 2 posttreatment. Deer foraging habitat was clearly improved by the HB treatment in both regions by year 2. Forb species of annual importance to deer increased in percent cover and biomass in the UCP and in biomass in the LCP. We estimated nutritional carrying capacity using a target diet quality of 14% crude protein; estimates in HB plots were 3 times greater than controls in the UCP and 19 times greater in the LCP. Although UCP sites had baseline carrying capacities nearly 8 times greater than LCP sites, the greater relative response to HB in the LCP eliminated the regional difference. Our results indicate that imazapyr herbicide treatment followed by prescribed fire is a beneficial tool for deer management in mid-rotation pine plantations.
Chronic herbivory by white-tailed deer (Odocoileus virginianus Zimmerman) can have profound impacts on the function and structure of forest ecosystems. We examined the combined influence of intense herbivory associated with a deer population eruption and chronic herbivory by the post eruption population on the spring flora of Cades Cove, Great Smoky Mountains National Park. During the 1970s the deer population reached a peak of 43 deer per km2, from which it has slowly declined in recent decades. To examine the influence of intense herbivory, we compared the abundance and flowering rates of early flowering plants in Cades Cove to a nearby reference site with similar bedrock geology, vegetation, and disturbance history but contrasting history of deer abundance. Our results suggest that significant changes (p ⩽ 0.05) in the diversity, evenness, and species richness of the spring flora occurred during the eruptive phase. Trillium spp. and other liliaceous species appeared to be disproportionately impacted. Comparisons between control and exclosure plots established after the deer population eruption indicate that recovery has been largely restricted to species that were able to persist under intense herbivory. These species have increased in number in exclosures, suggesting continued impacts by deer on the plant community outside the exclosures. Little to no recolonization by browse sensitive species was observed. Consequently, to restore the natural diversity of early flowering plants once present in Cades Cove, active restoration may be necessary in addition to maintaining deer densities below current levels.
White-tailed deer (Ododcoileus virginiana) can substantially affect the structure and species composition of a forest. The tolerance of a forest community to browsing may vary by type as a result of varying biotic and abiotic factors of the environment. To date, no studies have compared the effects of browsing among forest communities within a physiographic region. We investigated the effects of browsing on vegetation structure and woody seedling composition in three forest types (oak–hickory, Virginia pine–eastern red cedar, bottomland hardwood) in Manassas National Battlefield Park (MNBP), Virginia, USA. We compared forb cover, vertical plant cover (0–1.5 m tall), and survival of tagged seedlings in 10 exclosed (2 m × 6 m) and 10 unexclosed plots in each forest type during a 5-year period. No differential effects of browsing were found among forest types. In all forest types, deer (67 deer/km2) suppressed forb and vertical plant cover to levels less than would be expected in the absence of deer. Seedling survival rates of most species were significantly reduced by browsing. By the 4th year of the study, box elder (Acer negundo), hickory (Carya spp.), and red maple (Acer rubrum) had been eliminated from unexclosed plots, and red and white oaks (Quercus spp.) dramatically reduced. Ash (Fraxinus spp.), black cherry (Prunus serotina), and hackberry (Celtis occidentalis), although significantly impacted, remained the most abundant species throughout the study. These findings suggest that white-tailed deer may be modifying the structure of the forest interior to the extent that it adversely affects wildlife species dependent on a dense understory to thrive. We predict that the future composition of forests in MNBP will shift towards stands with fewer species and a greater dominance of ash, black cherry, and hackberry, particularly in the oak–hickory and bottomland hardwood forests, where the majority of current dominants are most affected.
Supplemental feeding of wintering white-tailed deer (Odocoileus virginianus) has increased in recent years, yet little data address its implications. During January-March 1993 and 1994, consumption of supplemental feed, provided ad libitum, was monitored daily at 2 recently established supplemental feeding (RSF) sites in northcentral Minnesota. We also monitored browse availability and use monthly at the 2 RSF sites and at 2 control (no supplemental feeding) sites by a sequential clipping method. Cumulative feed consumption and deer-visits were consistent between sites within years, but not between years; greater feed consumption and deer-visits occurred during the more severe Winter 1994. Browse pressure differed (P < 0.0001) by month during Winter 1993, and by month, twig density (sparse, moderate, dense), and site during 1994. A site x distance interaction (P < 0.0001) in 1993 was not significant in 1994 when browse pressure was higher. During 1994, mean browse pressure at control and RSF sites was different (P< 0.0001). A key finding in our study was that supplemental feeding effects differed annually. Furthermore, in both years, browse pressure means of both RSF sites differed, in the same manner, from mean browse pressure at the 2 control sites. The observed consistency of response among replicates, within treatments and years, suggests that supplemental feeding effects may apply generally to other local deer populations.
In loblolly pine Pinus taeda stands in the Francis Marion National Forest, South Carolina, prescribed burning led to higher Mg in total white-tailed deer Odocoileus virginianus forage in May, higher Mg in stems in total forage in May, higher Ca in stems on controls in July, higher P in palatable woody plant leaves in July, higher digestibility of palatable stems in July, higher crude protein in gallberry Ilex glabra and waxmyrtle Myrica cerifera leaves, and lower crude protein in waxmyrtle stems on burns in May the 1st yr after burning. -P.J.Jarvis
We tested the hypothesis that intensity of use of warm-season food plots by white-tailed deer (Odocoileus virginianus) increases with decreasing nutritional quality and quantity of vegetation in the surrounding habitat. Relative track density (%) in 3-m-wide avenues surrounding each food plot (n=4) and percent of food-plot forages in feces (determined by microhistological analysis) were determined during June, August, and October 1992 and 1993. Nutritional carrying capacity of the food plots (FCC) and habitat (HCC; deer days/ha) was estimated by multiplying available forage (kg/ha) by digestible energy of the forage (kcal/g) and dividing the product by the daily digestible energy requirement of a 60 kg deer (kcal/deer day). Relative track density (%) and percent of food-plot forages in feces collected within 13-ha sites consisting of food plots and the surrounding habitat were more strongly related to FCC (R-2 = 0.39, P = 0.006 and R-2 = 0.47, P < 0.001; n = 24 [6 sampling dates X 4 food plots]) than to HCC (R-2 = 0.24, P = 0.055 and R-2 = 0.10, P = 0.354). Deer feces collected in 13-ha sites (n = 4) 1 km from each food plot contained less food-plot forage (P < 0.05). more browse (P < 0.05) and cacti (P = 0.016), and similar (P = 0.456) native forbs than feces collected within 13-ha sites (n = 4) centered on each food plot. Intensity of use of food plots was related to standing crop and nutritional quality of food plot forages, however, variation in intensity). of use of food plots was explained only partly by variation in FCC and HCC. Because food plot forages did not replace native forbs in deer diets, the use of food plots to artificially increase carrying capacity might result in excessive use of native forbs.
A statistical technique evaluating preference or avoidance of a given habitat or forage species is presented, using moose (Alces alces) distribution patterns in an area including the Little Sioux Burn of northeastern Minnesota as an example. The technique is used in conjunction with a chi-square analysis, after the chi-square has led to the rejection of the null hypothesis that a set of observations follows an "expected" occurrence pattern. The technique involves the use of a Bonferroni z statistic which may be used in estimating whether a specific observation occurs more or less frequently than expected. The technique provides a refinement of quantitative methods which heretofore have not been used to test a multinomial distribution applicable to the example.
White-tailed deer (Odocoileus virginianus) populations have been maintained at high densities in Pennsylvania for several decades with unknown effects on songbirds and their habitats. I evaluated effects of white-tailed deer density on songbird species richness, abundance, and habitat. I simulated 4 deer densities (3.7, 7.9, 14.9, and 24.9 deer/km2) within individually fenced enclosures on 465-ha forest areas in northwestern Pennsylvania. Within all enclosures, 10% of the area was clear-cut and 30% was thinned. Enclosures were subjected to 10 years of deer browsing, 1980-90, at the 4 simulated densities. I conducted bird counts in 1991. Varying deer density had no effect (P > 0.1) on ground- or upper canopy-nesting songbirds or their habitat, but species richness of intermediate canopy-nesting songbirds declined 27% (P = 0.01) and abundance declined 37% (P = 0.002) between lowest and highest deer densities. I did not observe the eastern wood pewee (Contopus virens), indigo bunting (Passerina cyanea), least flycatcher (Empidonax minimus), yellow-billed cuckoo (Coccyzus americanus), or cerulean warbler (Dendroica cerulea) at densities >7.9 deer/km2, and the eastern phoebe (Sayornis phoebe), and American robin (Turdus migratorius) were not observed at 24.9 deer/km2. Threshold deer density for effect on habitat and songbirds within managed (100-yr rotation) forests was between 7.9 and 14.9 deer/km2.
Year-round foraging of white-tailed deer (Odocoileus virginianus) on the national forests of the Southern Appalachian mountains in Georgia, North Carolina and Tennessee was determined by analysis of rumen samples from 581 deer taken over 5 years and by sampling forage availability and evidence of browsing in selected habitat types. Spring and summer diets consisted mainly of green leaves and succulent twig tips of deciduous woody plants (58% in spring, 54% in summer). These forages were more abundant and used more intensively by deer in and around clearcuts than in adjacent older forests. Most other important foods in these seasons (i.e., fruits, forbs and fungi) also were much more abundant in the clearcuts. Flowers of yellow-poplar (Liriodendron tulipifera), which dominated the diet in late May and early June, and squawroot (Conopholis americana), a parasitic herb associated with red oaks (Quercus spp.) and an important secondary food throughout most of the spring and summer, were more available in older forests. Fall and winter diets were influenced greatly by the availability of mast, especially acorns. When mast was abundant, it dominated the diet. When mast was scarce, deer fed mainly on the leaves of broadleaf evergreen woody plants, especially rosebay rhododendron (Rhododendron maximum). Twig ends were used little by deer during fall and winter. Older forests generally provided best foraging conditions for deer in fall and winter, being the primary source of mast and also producing about the same quantity of broadleaf evergreen foliage as clearcuts. Clearcuts and older forests provide complementary benefits to deer. Forests of intermediate age (after crown closure reduces browse availability and before onset of significant mast production) contribute much less to the deer forage resource than either clearcuts or older forests. Specific contributions of different forest age classes to the forage base vary with site characteristics. Implications of changing forest management policy are discussed.
White-tailed deer (Odocoileus virginianus Zimm.) appear to concentrate foraging activity along the perimeters of warm-season food plots. Because of this, we tested the hypothesis that (1) providing travel lanes (i.e., rows not planted) free of vegetation within food plots will increase deer use of the plots and result in an equal spatial distribution of forage use within the plots, and (2) skip-row planting will result in increased yield and survival of lablab (Dolichos lablab L.), an annual legume. During 1994 and 1995, lablab was established by planting (1) every row spaced 0.9 m apart (solid), (2) 2 rows and not planting 1 row (skip 1), and (3) 2 rows and not planting 2 rows (skip 2) in two 5-ha food plots. Planting scheme did not affect spatial patterns of food plot use by deer. Utilization was concentrated at food plot perimeters on 9 of 15 sampling dates. Food plot utilization by deer was greater in skip 2 treatments only during August 1995, possibly as a result of greater forage availability resulting from greater plant survival than solid rows. Deer foraging in food plots apparently shifted foraging activities to an area of greater forage availability as the resource supply was depleted. Skip-row planting had lower overall planting costs/ha than solid planting but maintained similar forage production per hectare.
In each of 2 experiments, 6 different yearling white-tailed deer (Odocoileus virginianus) were fed 6 equicaloric concentrate diets with varying crude protein content. Maintenance requirement for protein was computed to be met by a diet containing 5.8 ± 2.0% crude protein (moisture-free basis). Each additional unit of dietary nitrogen resulted in 0.23 ± 0.01 units increase in retained body nitrogen and 0.66 ± 0.03 units increase in urinary nitrogen loss. Metabolic fecal nitrogen component was estimated to be 0.53 g/100 g dry matter consumed.
Mixed pine (Pinus spp.)-hardwood forests are common in the southern United States (U.S.), but little quantitative information exists on the response of understory forage to reductions in basal area from thinning. We determined understory forage characteristics before thinning and 2 and 4 years after thinning a 35-year-old natural loblolly pine (P. taeda)-hardwood stand (initially $27\ {\rm m}^{2}/{\rm ha}$ of pine and $8\ {\rm m}^{2}/{\rm ha}$ of hardwood basal area). A combination of 3 loblolly pine (15, 18, and $21\ {\rm m}^{2}/{\rm ha}$ ) and 3 hardwood (0, 3.5, and $7\ {\rm m}^{2}/{\rm ha}$ ) basal areas was replicated 3 times, resulting in 27 0.08-ha plots. Understory coverage and forage biomass were determined on 25 understory plots systematically located within each plot, with data analyzed using analysis of variance and regression. Herbaceous forage biomass and coverage and light intensity were correlated negatively (P<0.05) with retained pine and hardwood basal areas, with hardwood basal area being the more important factor. Stand thinning improved herbaceous forage availability for wildlife, but the response was time-dependent. Forage from woody browse and vines also increased following stand thinning, although responses were not as time-dependent as herbaceous forages. Results of our study indicate that managers can manipulate forage production by thinning stands to prescribed basal areas and compositions.
Drought frequently limits alfalfa (Medicago sativa L.) herbage productivity during mid-summer in the North Central USA, but the productivity of other legumes during drought is unknown. We determined the effect of drought on the herbage yield and quality and stand persistence of birdsfoot trefoil (Lotus corniculatus L.), cicer milkvetch (Astragalus cicer L.), red clover (Trifolium pratense L.) and alfalfa. Legumes were established on a sandy, mixed, Udorthentic Haploborolls and subjected to two soil water regimes promoting ‘droughted’ and ‘well-watered’ (control) plant growth. Mid-day stem pressure potentials for droughted and control plants ranged from −1.3 to −3.8 MPa and −0.1 to −1.4 MPa, respectively. When drought occurred throughout growth, legume herbage yields of droughted alfalfa, birdsfoot trefoil, cicer milkvetch, and red clover averaged 33, 21, 19, and 13% of their respective controls. Average herbage yield of droughted alfalfa was 120% greater than yields of birdsfoot trefoil and cicer milkvetch, and 165% greater than red clover yield. Averaged for all harvests and species, whole herbage acid detergent fiber (ADF), neutral detergent fiber (NDF), and acid detergent lignin (ADL) concentrations were reduced by 30 to 4870, 26 to 46%, and 0 to 49%, respectively, when drought occurred throughout growth. Effects of drought on lignin in NDF and crude protein concentration were not consistent. Droughted alternative legumes produced herbage with lower ADF, NDF, and ADL concentrations than alfalfa. Improved quality in droughted legumes was related to greater leaf:stem weight ratio, delayed maturity, and often higher quality in both the leaf and stem fractions compared to the control treatment. Although drought reduced the herbage yield of all legumes, alfalfa has the greatest yield potential in drought.
Contribution of the Minnesota Agric. Exp. Stn. Publ. no. 18796.
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Vegetation responses were compared following an array of timber harvest and fire regimes on oak-pine sites in the Ouachita Highlands of eastern Oklahoma during 8 yrs. Nine treatments were replicated 1-3 times in a completely randomized design on 23 units. Little and big bluestem dominated harvested and winter-burned treatments. September total standing crop was up to 25 times greater on harvested and burned than on control treatments. In general, response was related to overstory canopy cover, basal area, litter accumulation, and burn interval. Details for specific plant groups are presented and treatment recommendations are given.
Although many researchers have reported yield increases due to alfalfa irrigation when soil water has been limited, little information is available on alfalfa plant and soil water relationships which might be used to improve water management strategies. Our objective was to determine effects of plant water deficits on alfalfa seasonal growth patterns, forage quality, forage yield, water use, and water‐use efficiency. During 1981, established alfalfa grown on a sandy soil was subjected to four water supply levels designated: high irrigation (H), medium high irrigation (MH), medium low irrigation (ML), and unirrigated, rainfall only (U). Soil water depletions occurred for all treatments to at least 1.9 m. Alfalfa midday plant water potentials in H and MH treatments ranged from −0.7 to −1.3 MPa throughout the season. For alfalfa subjected to ML and U treatments, ψ mp reached −2.7 and −4.0 MPa, respectively. Dry weight (DW) accumulations for alfalfa in H and MH treatments increased with successive harvests during the summer while harvest DW for alfalfa in ML and U treatments declined. Relative growth rate (RGR) declined sharply as ψ mp decreased. At moderate plant water stress (ψ mp of −1.5 to −2.0 MPa), little growth occurred and under severe water stress, (ψ mp < −2.5 MPa), RGR were negative due to leaf loss. Reduced soil water availability resulted in increased leaf:stem weight ratios and in vitro dry matter disappearance (IVDMD), but only under severe and long‐term plant water stress. Crude protein (CP) concentration was unaffected by plant water status. It appeared that irrigation water could be used efficiently on coarse‐textured soils by moderate water application to alfalfa at 50% depletion of extractable soil water, an apparent threshold for maintenance of favorable plant water status. A close association between forage yield of individual harvests and cumulative indicates potential for initiation of irrigations when plant measurements indicate soil water depletions to a threshold level.
Browsing by white-tailed deer (Odocoileus virginianus) is a major cause of regeneration failure in Allegheny hardwood forests of northwestern Pennsylvania. I examined the impact of deer at 5 different densities (0, 10, 20, 40, and 80 deer/259 ha) on tree seedlings, woody shrubs, and herbaceous plants in large enclosures over 5 years. I examined 3 silvicultural treatments (clearcut, thinning, and uncut) at each density. After 5 years, tree seedlings in the clearcuts at the lowest deer densities were nearly twice as tall as those at the highest deer densities. Browsing at high deer densities also reduced the diversity of tree seedlings, resulting in nearly pure black cherry (Prunus serotina) regeneration. Fern cover ipereased with increasing deer densities and blackberry cover (Rubus spp.) decreased. I suggest deer populations should be maintained at : � 18 deer/ 259 ha to ensure tree regeneration and desired tree species composition.
A study was initiated in 1994 to evaluate the degree of bole damage and crown decline residual overstory trees would experience because of prescribed burning of shelterwood stands. Three oak-dominated shelterwood stands, partially harvested 2 to 4 yr earlier, were divided into four treatments (unburned control, spring burn, summer burn, and winter burn). Fifteen permanent sampling points were systematically located in each 5 to 12 ac treatment area, and overstory trees were selected from these points with a 10 BAF prism. Before burning, each tree was evaluated for lower bole and crown condition and reevaluated two growing seasons after the fires. Hickory, oak, and yellow-poplar were largely unaffected by the winter and summer prescribed fires but displayed bole damage and crown decline following spring burning. American beech and red maple declined after all fire treatments. Fire damage to oak, hickory, and yellow-poplar was strongly associated to presence of logging slash near a tree's base. Directional felling or moving slash should minimize injury to these trees. This research will aid resource managers wishing to use prescribed fire in shelterwood stands to favor oak regeneration while minimizing damage to residual overstory oaks. South. J. Appl. For. 23(2):88-93.
Understory response was studied for six years after thinning cove hardwood stands in the mountains of western North Carolina and northern Georgia. Understory vegetation increased from an average 294 lb/acre before thinning to 512 lb/acre the first year after thinning. Average peak production of 692 lb/acre was reached in the third year, more than twice the prethinning production and began to decline in the fourth year. Both the level and duration of the increase were related to degree of thinning. The increased production included a variety of woody and herbaceous vegetation known to be heavily utilized by both deer and grouse. The thinnings resulted in significant, if temporary, improvement in habitat for these and perhaps other species.