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Food Habits of Reintroduced Elk in Southeastern Kentucky
Jennifer Schneider, David S. Maehr, Karen J. Alexy, John J. Cox, Jeffery L. Larkin, and Brian C. Reeder

Southeastern Naturalist, Volume 5, Number 3 (2006): 535–546

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2006 SOUTHEASTERN NATURALIST 5(3):535–546 Food Habits of Reintroduced Elk in Southeastern Kentucky JENNIFER SCHNEIDER1, DAVID S. MAEHR2,*, KAREN J. ALEXY3, JOHN J. COX4, JEFFERY L. LARKIN2,5, AND BRIAN C. REEDER1 Abstract – Based on microhistological examinations of feces, Cervus elaphus (elk) from a reintroduced herd on the Cumberland Plateau in southeastern Kentucky exhibited an annual diet of grasses (24%), forbs (27%), and browse (32%). Diets shifted seasonally, possibly in response to availability and palatability. Forbs dominated the summer diet (34%), whereas grasses, forbs, and woody browse accounted for approximately equal portions of the fall diet. Grasses (40%), and browse (46%) dominated the diet during winter and spring, respectively. Grasses were eaten less during spring (10%) than during any other month. Nutritional quality does not appear to be limiting in this growing population. Introduction Although the ecology, demographics, and behavior of Cervus elaphus L. (elk) introduced into Kentucky have been examined, their local food habits remain largely unknown here and elsewhere in eastern reintroduction sites (Maehr et al., in press). A small-scale examination in Virginia is the sole exception (Baldwin and Patton 1938). Since 1997, the Kentucky elk herd has grown steadily—initially from annual releases of hundreds of animals transported from western herds, followed by steady reproduction and recruitment (Larkin et al. 2003). The current population is estimated at about 4000 (J.Gassett, Kentucky Department of Fish and Wildlife Resources, Frankfort, KY, pers.comm.), and most animals have remained associated with reclaimed surface mines where 7 of 8 release sites were located. The success of achieving the goal of 7500 elk in southeastern Kentucky (Phillips 1997) will depend in part on the availability and quality of forage. Poor nutrition in elk herds can negatively impact pregnancy rates, age at first breeding, fetal survival, birth weight, juvenile growth, juvenile survival, and adult survival (Cook 2002). Recent reproductive patterns and population growth suggest that elk in southeastern Kentucky exist on a high nutritional plane (Larkin 2001). However, landscape changes related to surface mining, subsequent reclamation, and altered successional patterns make elk habitat 1Morehead State University, Department of Biological and Environmental Sciences, Lappin Hall, Morehead, KY 40351. 2University of Kentucky, Department of Forestry, 205 Cooper Building, Lexington, KY. 3Kentucky Department of Fish and Wildlife Resources, 215 Cooper Building, Lexington, KY 40546-0073; 4University of Kentucky, Department of Forestry, 208 Cooper Building, Lexington, KY 40546- 0073. 5Current address - Department of Biology, Indiana University of Pennsylvania, Indiana, PA 15705. *Corresponding author - dmaehr@uky.edu. 536 Southeastern Naturalist Vol. 5, No. 3 management an inexact science in the southern Appalachians. This is especially true when the most preferred and nutritious foods are unknown. As Nelson and Leege (1982) observed, “effective elk habitat management requires skillful application of integrated knowledge of the animals’ diet, their need for nutriment, cover, and space, and how to manipulate habitat to provide for these needs.” Whereas the importance of diet to overwinter health may be less in the relatively snow-free, low-elevation southern Appalachian Mountains than in the western United States, where winter movements and diet are often related to snow depth (Cook 2002), the maintenance of a high nutritional plane will be essential for population growth and good herd health of Kentucky elk. It remains “unknown to what degree the modern plant life of southeastern Kentucky will provide satisfactory nutrition for elk” (Maehr et al. 1999). Thus, the objective of this study was to describe what introduced elk are eating in eastern Kentucky. Study Area We studied elk food habits at the Addington Wildlife Management Area (WMA) (formerly Cyprus-Amax WMA), a 7400-ha surface coal mine that consists of 4400 ha of forest, 2000 ha of reclaimed grassland, and 1000 ha of active mining (Larkin et al. 2002). It is centrally located within the 14-county, 1.06 million-ha elk restoration zone in southeastern Kentucky. The larger restoration zone is approximately 80% second and third growth, mixedmesophytic forest (Braun 1950), 10% active and reclaimed surface mines, 9% agriculture or cleared lands, and 1% urban (Cox 2003). Forests occur on narrow winding ridges, steep side slopes, and deep dendritic drainages (McFarlan 1943). Elevations ranged from 244 to 488 m above mean sea level (Overstreet 1984). Reclaimed surface mines are dominated by a variety of exotic grasses and forbs, including Lolium arundinaceum Schreb. (Kentucky- 31 tall-fescue), Lespedeza spp., Michx. (bush clover), Coronilla varia L. (crown vetch), and Lotus corniculatus L. (birds-foot trefoil). Both planted and colonizing woody species on the mines include Robinia pseudoacacia L. (black locust), Alnus glutinosa L. (black alder), Elaeagnus umbellata Thunb. (autumn-olive), and Pinus strobus L. (white pine). The climate is temperate humid continental with warm to hot summers and cool winters (Overstreet 1984), with a mean minimum January temperature of -4° C, a mean maximum July temperature of 30° C and an overall mean temperature of 13° C (Ulack et al. 1998). Annual precipitation averages 117 cm (Hill 1976), with the highest rainfall occurring in March and July (Ulack et al. 1998). Snow rarely accumulates for more than a few days during winter. Methods We used microhistological examination of fecal material (McInnis et al. 1983) to identify the components of elk diets in southeastern Kentucky. We used binoculars while in a vehicle to watch elk on alternating 2006 J. Schneider et al. 537 weekends from July 2002 to April 2003. Defecating elk were identified to gender and fresh fecal pellet samples were placed in plastic bags and sealed, with time and location recorded, then frozen. All feces from observed elk were collected within 1 hour of defecation. We did this to allow foraging elk to move away from the foraging area and reduce disturbance to the herd. In addition, feces from unidentified animals were collected when specimens appeared fresh, and elk were absent. The inclusion of soil, rocks, and vegetation were avoided to prevent sample contamination. We collected reference plants from 100 of the common species in the region (Overstreet 1984). Reference slides of stems and leaves were made (Nelson and Leege 1982), and cell features from collected plants were compared to fecal samples. We combined fecal samples based on week of collection to make composite samples using 2 pellets from each individual sample. We then grouped these composite samples based on season. Seasons were summer (July–August), fall (September–November), winter (December–February), and spring (March–April). We ground composite samples in a coffee grinder then forced them through a #18 mesh screen (1-mm opening; Hansen et al. 1978). Screened fragments were collected in a 15-ml test tube. Two ml of bleach were added to the test tube, which was then capped and shaken for 30 seconds (Hinnant and Kothmann 1988). One drop of 17.5% hydrochloric acid then was added to neutralize the bleach. This mixture was shaken by hand then centrifuged at 3400 rpm (relative gravitational force = 1380) for 1 min. The sample was then washed with 10 ml of distilled water, and centrifuged 3 times. This sample was mixed with 1–2 ml of distilled water and 5 drops of safranin-O solution and homogenized for 30 sec. This mixture was washed with 10 ml of distilled water and centrifuged; this process was repeated 3 times. The final mixture was washed through a #200 mesh screen (0.074-mm openings; Hansen et al. 1978). The remains were then scraped off the screen and onto a microscope slide with a 6-mm diameter template hole. We used a similar volume on each slide, based on the diameter and depth of the holes in the template, to ensure uniformity among samples. The slurry was mixed with Hoyer’s mounting media (500 g chloral hydrate, 50 ml glycerine, 150 g gum Arabic, 125 ml hot water; Foppe 1984), covered with a glass slip, and sealed over an alcohol flame. Slides were dried for 72 h at 50° C. We viewed epidermal characteristics of material from collections and fecal samples using a Hoffman modulation contrast microscope. At least 2 characteristics, such as trichomes, stomates, silica cells, and cell walls, were present before a positive classification was made (Holchek and Valdez 1985). Forbs required an identifiable characteristic plus cell walls; whereas grass required 2 characteristics other than cell walls, such as stomates and couples, stomates and silica cells, or stomates and crystals (Foppe 1984). A valid field had 2 identifiable fragments or a single fragment that covered 20% of the field (Foppe 1984). Five slides for each composite group were made. These were further subdivided into 20 fields per slide to determine the 538 Southeastern Naturalist Vol. 5, No. 3 density of each plant species. For each composite sample we tallied the number of fields (5 slides x 20 fields = 100) in which an identifiable plant fragment occurred, then converted this to relative density following Hansen et al. (1978) and Hansen and Clark (1977). The resulting value is highly correlated to dry weight consumed and can be used as an estimate of ingested vegetative mass (Rentfleish and Hansen 2000, Sparks and Malechek 1968). Temporal variation in the diet was determined by comparing relative density for plant species across seasons. Finally, we recorded additional plants that were consumed by elk while we watched them during routine fieldwork from 1997–2005 throughout occupied range in eastern Kentucky, and that were identified in the rumen of a dead animal in the Red Bird Unit of the Daniel Boone National Forest. Results We opportunistically collected 149 fecal pellet groups (48 from males, 49 from females, and 52 of unknown gender) and organized them into 41 composite slides (winter = 7, spring = 10, summer = 12, fall = 12). Collections came from at least 20 marked elk and an unknown number of unmarked elk. The annual diet was composed of grasses (24%), forbs (27%), and woody browse (32%: 16% shrubs and 16% primarily deciduous trees) (Table 1). Forbs dominated the summer diet (34.4%), whereas grasses were second (27.0%). During fall, grasses, forbs, and woody browse accounted for approximately equal portions of the diet (Table 1). Grasses dominated the winter diet (40%). Forbs and browse accounted for less than 24% of the diet at this time. Woody browse dominated the spring diet followed by forbs. Grasses were eaten less during spring than other seasons (Table 1). In general, when grass use was high, elk consumption of woody browse was low (Fig. 1). Of the 100 reference samples collected in eastern Kentucky, 30 were not found in our composite samples (Table 2). Relative density of grass dry matter varied among seasons. Although Bromus spp., L. (brome) dominated the summer diet, the other species consumed during this season were eaten fairly evenly by elk (Table 3). The fall diet was dominated by Dactylis glomerata L. (orchard grass), Lolium perenne L. (perennial ryegrass), and Lolium arundinaceum. Lolium perenne dominated the winter diet, followed by Lolium arundinaceum and Dactylis glomerata. Grasses in the spring diet were not as prominent; however, Lolium arundinaceum was the main choice. Table 1. Seasonal percent frequencies of foods found in fecal samples of elk in southeastern Kentucky during 2002 and 2003. Category Summer Fall Winter Spring Annual Grass 27.0 17.7 40.0 9.7 23.6 Forbs 34.4 21.8 23.7 26.9 26.7 Browse 23.2 41.9 17.8 46.1 32.2 Unknown 15.3 18.6 18.5 17.3 17.4 2006 J. Schneider et al. 539 Among forbs, Trifolium pratense L. (red clover) and Lespedeza cuneata Dum.-Cours. (bush clover) were used most in summer. During fall, bush clover remained dominant, whereas Trifolium pratense use declined. Polystichum acrostichoides Michx. (Christmas fern), Lespedeza cuneata, and Trifolium pratense were the main forb choices in winter. Spring forbs mainly consisted of Lespedeza cuneata, Trifolium pratense, and Polystichum acrostichoides (Table 3). Woody browse use was lowest during winter (Fig. 1). During fall, however, Cornus florida L. (flowering dogwood), and Elaeagnus umbellata Thunb. (autumn olive) were used most among browse species. Autumn olive was also an important food choice in winter and spring. Trees did not dominate the diet during any season, however, Robinia pseudoacacia was eaten heavily in all seasons except winter. Acer (likely rubrum, red maple) was the most utilized tree species during winter (Table 3). One species of conifer, Tsuga canadensis L. (eastern hemlock), was found as a trace amount during summer. We also observed elk consuming Arundinaria gigantea Walt. (switchcane), Triticum aestivum L. (wheat), Zea mays L. (corn), Typha spp., L. (cattail), Pinus strobus, Magnolia spp., L. (magnolia), Morus alba L. (white mulberry), Prunus spp., L. (a cultivar cherry), Pyrus calleryana Dcne. (Bradford pear), Malus coronaria L. (American crab-apple), Andropogon gerardii Vitman (big bluestem), Phytolacca americana L. (pokeweed), and Coronilla varia. The latter three species were represented in our reference collection, but did not appear in fecal samples. Figure 1. Seasonal variation in relative density among the 3 major diet components based on fecal analyses of southeastern Kentucky elk from 2002–2003. 540 Southeastern Naturalist Vol. 5, No. 3 Table 3. Relative densities of plant species eaten based on microhistological analyses of elk fecal samples in southeastern Kentucky, 2002–2003. Number of composite samples per season appear in parentheses. Su = summer (n = 12), F = fall (n = 12), W = winter (n = 7), and Sp = spring (n = 10). Species Common name Su F W Sp Grasses Agropyron spp. L. Wheatgrass 3.3 0.0 0.0 0.0 Andropogon virginicus L. Broom-sedge 4.9 1.5 3.5 1.9 Bromus spp. Brome 13.4 0.0 0.0 0.0 Carex spp. Sedge 0.7 0.0 0.0 0.0 Dactylis glomerata L. Orchard grass 2.6 12.6 13.9 2.8 Eragrostis spp. Lovegrass 0.6 0.0 0.0 0.3 Lolium arundinaceum (Shreb.) Kentucky fescue 7.0 7.2 14.1 7.0 Lolium perenne L. Perennial ryegrass 3.9 8.8 26.4 2.9 Miscanthus sinensis Anderss. Chinese silver grass 2.4 0.8 0.0 0.0 Panicum virgatum L. Panic grass 2.7 0.7 0.4 0.9 Phleum pretense L. Timothy 3.4 0.7 0.0 3.4 Poa spp. Bluegrass 1.6 1.1 0.7 0.8 Sorghastrum nutans L. Yellow indiangrass 2.7 0.0 0.0 0.0 Table 2. Reference species from southeastern Kentucky that were not found in elk fecal samples, 2002–2003. We observed elk consuming species indicated by an asterisk. Scientific name Common name Acalypha virginica L. Three-seeded mercury Achillea millefolium L. Common yarrow Ambrosia artemisiifolia L. Annual ragweed Andropogon spp.* Broomsedge* Asimina triloba L. Pawpaw Bidens aristosa Michx. Beggar tick Boehmeria cylindrical L. False nettle Cacalia atriplicifolia L. Pale Indian plantain Campanulastrum americanum L. Tall bellflower Cichorium intybus L. Chicory Cirsium discolor (Muhl. ex Willd.) Field thistle Clematis virginiana L. Virginia virgin bower Conyza Canadensis L. Canada horseweed Coronilla varia L.* Common crown vetch* Cyperus strigosus L. Straw-colored flatsedge Dentaria laciniata L. Pepperwort Echinochloa crus-galli L. Barnyard grass Eleocharis engelmannii Steud. Engelmann’s spike rush Eupatorium maculatum L. Spotted Joe-Pye weed Eupatorium serotinum Michx. Late-flowering thorough-wort Ilex opaca Ait. American holly Lactuca spp. Wild lettuce Phytolacca americana L.* Common pokeweed* Sassafras albidum Nutt. Sassafras Smilax rotundifolia L. Common greenbriar Solidago juncea Ait. Early goldenrod Solidago nemoralis Ait. Field goldenrod Triodia flava L. Tall purple-top fluffgrass Vernonia angustifolia Michx. Tall ironweed 2006 J. Schneider et al. 541 Table 3, continued. Species Common name Su F W Sp Forbs Ambrosia artemisiifolia L. Annual ragweed 6.5 1.2 2.7 0.0 Artemisia spp. Wormwood 0.4 0.0 0.0 0.0 Aster spp. Aster 0.7 2.2 2.2 2.6 Lespedeza spp. Bush-clover 4.8 2.1 0.7 0.8 Lespedeza cuneata Dum.-Cours. Chinese bush-clover 15.2 18.6 8.8 22.8 Lespedeza striata Thunb. Korean clover 2.4 1.6 0.0 1.2 Lespedeza violacea L. Violet bush-clover 0.9 0.0 0.0 0.0 Lotus corniculatus L. Birds-foot trefoil 0.6 3.8 0.7 0.6 Melilotus spp. Sweetclover 0.8 1.0 2.2 0.0 Oxalis spp. Woodsorrel 0.0 0.0 0.0 0.9 Polygonum pensylvanicum L. Pennsylvania smartweed 1.0 1.7 3.3 3.4 Polystichum acrostichoides Michx. Christmas fern 0.7 1.4 9.6 14.5 Pycnanthemum incanum L. Hoary mountain-mint 1.1 1.0 0.0 0.0 Sabatia angularis L. Square-stemmed rose pink 4.0 1.8 0.0 0.0 Solidago canadensis L. Canada goldenrod 0.8 0.0 0.0 0.0 Trifolium pretense L. Red clover 22.3 4.7 4.9 9.4 Woody browse Acer spp. Maple 0.0 0.0 3.0 2.8 Acer rubrum L. Red maple 1.6 0.8 0.0 0.6 Acer saccharinum L. Silver maple 2.3 6.8 0.0 0.0 Acer saccharum Marsh Sugar maple 1.4 0.7 0.0 0.0 Aralia spinosa L. Hercules club 2.6 1.4 2.7 0.6 Amelanchier arborea Michx. f. Downy serviceberry 0.9 0.0 0.0 0.0 Carya ovata P. Mill. Shagbark hickory 1.9 0.7 0.7 6.0 Cercis Canadensis L. Eastern redbud 1.8 0.8 0.6 1.9 Cornus florida L. Flowering dogwood 1.6 19.5 0.0 0.0 Elaeagnus umbellate Thunb. Autumn olive 4.6 18.6 15.7 28.8 Fagus grandifolia Ehrh. American beech 1.5 2.9 0.0 0.0 Lindera benzoin L. Spicebush 1.4 1.8 0.0 0.0 Liriodendron tulipifera L. Tulip tree 3.0 5.6 0.0 0.0 Oxydendrum arboretum L. Sourwood 3.7 0.9 0.0 0.6 Platanus occidentalis L. Sycamore 1.0 0.8 0.8 2.2 Quercus alba L. White oak 0.0 0.6 0.0 0.0 Quercus coccinea Muenchh. Scarlet oak 0.0 0.6 0.0 0.0 Quercus prinus L. Chestnut oak 0.9 1.3 0.0 0.0 Quercus velutina Lam. Black oak 0.0 3.0 0.0 0.0 Robinia pseudoacacia L. Black locust 9.2 8.9 1.8 27.4 Rosa mulitflora Thunb. Multiflora rose 0.8 1.2 0.0 3.0 Rubus spp. Rose 0.6 0.9 0.0 22.2 Tilia spp. Basswood 0.6 1.5 0.8 0.0 Tsuga canadensis L. Eastern hemlock 0.6 0.0 0.0 0.0 Unknown Total unknown 27.9 35.2 27.3 36.2 Discussion Among woody plants, the invasive Elaeagnus umbellata (Hoffman and Kearns 1997), Robinia pseudoacacia, and Rubus spp. L. (blackberry) were utilized throughout most of the year by Kentucky elk. Along with Lespedeza cuneata, these were the most commonly consumed plants during spring. 542 Southeastern Naturalist Vol. 5, No. 3 These species are ubiquitous in reclaimed and abandoned surface mines where elk are common. Interestingly, Elaeagnus umbellata was not recorded as an elk food in Giles and Bland Counties, VA by Baldwin and Patton (1938), even though this plant was introduced in eastern North America in 1830 (Rehder 1940). We suspect that because this area is not in Virginia’s coal-mining region (Sites 1995), Elaeagnus umbellata had little opportunity to establish in the area inhabited by elk in the 1930s. Species recorded as common elk food in Virginia, but not in Kentucky fecal analyses included Galax aphylla L. (galax), Gaultheria procumbens L. (wintergreen), Zea mays, and Pyrus malus L. (apple) (Baldwin and Patton 1938). The presence in the diet of the two domesticated plants may be a reflection of Virginia elk living in regions where agriculture was more widespread. Indeed, subsequent crop depredations lead to the eventual extermination of elk in Virginia (O’Gara and Dundas 2002). The occurrence of Zea mays in our opportunistic list derived from the stomach contents of one animal that resided near a small farm. However, the use of the planted Prunus spp., Triticum aestivum, and Pyrus calleryana hint at the potential for more widespread use of cultivars in areas near human settlements. Anecdotal information relating to complaints from home owners suggests that the use of crops such as Zea mays is not unusual in southeastern Kentucky. Although digestibility of woody browse is higher than grasses during summer (Cook 2002), it is likely that increased availability and digestibility of forbs such as Trifolium pratense, Lespedeza cuneata, and Ambrosia artemisiifolia L. (ragweed) explain reduced browsing after the spring leafout. In Colorado, percentage crude protein and dry-matter digestibility are highest among forbs during early summer (Baker and Hobbs 1982), a pattern that likely exists in Kentucky as well. Dry-matter digestibility remains relatively constant among grasses throughout the year; thus, with reduced digestibility of woody browse and the disappearance of many forbs, elk grazing, especially on Dactylis glomerata and Lolium perenne, increases in fall and peaks in winter. During fall, the relatively even use of Cornus florida, Lespedeza cuneata, and Dactylis glomerata made this the season with the least differentiation among plant types. Among all species, the plants used most consistently throughout the year were Elaeagnus umbellata, Robinia pseudoacacia, Lolium perenne, Lolium arundinaceum, Lespedeza cuneata, and Trifolium pratense. In general, seasonal dietary patterns reflect the vegetative characteristics of occupied range (Cook 2002). For example, grasses accounted for 50–75% of elk diets in northern California (Cook 2002), whereas woody browse made up 50–70% of the diet in forest-dominated western Oregon (Harper 1971). In Kentucky, elk behaved as intermediate feeders (Hofman 1985) with an annual diet evenly divided among grasses, forbs, and browse. The considerable variation among the components of the elk diet in southeastern Kentucky is likely due to a combination of seasonal changes in forage quality and the many different kinds of food that occur in 2006 J. Schneider et al. 543 a landscape that does not experience dramatic changes in availability due to snow accumulation and temperature extremes. Thus, although woody browse is available throughout the year, it is the primary component of the diet only during spring when new growth is more palatable. At other times of the year, woody browse provides fewer nutritional benefits. Grasses are used most in winter when tree leaves are mostly absent, and when forbs are least available. The low use of grasses in spring suggests a preference for woody browse at this time of year because both food groups offer rapidly growing and lush vegetation. The relatively high use of forbs during summer reflects the apparent widespread availability of this group in both grassland and forested settings. After the last of 1543 elk were released in southeastern Kentucky in 2002, the herd required only 3 years to reach an estimate of > 4000 (J. Gassett, pers. comm.)—a possible mean annual increase of nearly 70%. This increase has occurred despite up to 2–5% annual mortality due to Parelaphostrongylus tenuis Dougherty (meningeal worm); Alexy 2004, Larkin et al. 2003), and 5 consecutive fall hunts with harvests ranging from 12 to 40 from 2000–2004. Reproduction, including occasional twinning (Larkin 2001), successful breeding of yearling females, and rapid individual growth rates (K. Alexy, unpubl. data) are strong indicators of an almost unlimited food supply with adequate to superior nutrition, even though ungulate forage on southern upland sites tends to be nutritionally suboptimal (Thill et al. 1990). The determination of food preferences is complicated by the availability of forage and the density of the population of interest (Kufeld 1973). In our study, forage availability was not measured, thus preferences could not be determined with confidence. In addition, the population is believed well below the target density (Phillips 1997) as it continues to expand into vacant range. Future changes in the nutritional condition of the herd may result from shifts in diets that are related to higher elk density. For now, however, we believe that the elk population in southeastern Kentucky is not limited by forage quality or availability. Conclusions and Management Recommendations Although eastern Kentucky remains a mostly forested landscape, large expanses of artificial grassland allow elk to consume high-quality foods throughout the year. This, combined with the absence of severe winters and the lack of natural predators such as Canis lupus L. (wolf) and Puma concolor L. (cougar), may discourage migratory movements (Irwin 2002) and encourage continued herd growth. Further, continued coal mining will result in more grassland habitat than currently exists in the region. Maintaining a landscape matrix that maintains the array of habitats that provide elk with an exceptional diet should not be problematic in the next few decades. The challenge to elk managers in Kentucky will be to maintain and improve existing habitat, while restoring or maintaining 544 Southeastern Naturalist Vol. 5, No. 3 post-reclamation forests of various age classes and related biodiversity. The conditions that appear to promote elk colonization, good nutrition, and herd growth are the same that encourage symptoms of forest decline such as habitat loss, the invasion of exotic species, the creation of edge, and the loss of forest interior conditions and associated interior obligate species. Already, elk appear to be changing forest succession and soils in ways that will exacerbate the conditions resulting from surface mining and reclamation (Ter Beest 2005). Therefore,we encourage managers to reduce the negative consequences of expanding post-reclamation grasslands on native biodiversity by maintaining a population of elk that does not compromise forest connectivity or the potential to restore it. Certainly, elk should not be used as a rationale for expanded surface mining, but simply as a grassland-compatible ungulate that can prosper in a highly altered and disturbed landscape. Given the relatively high proportion of unknown food components in our study, we encourage future studies to help detail this aspect of elk ecology so that a complete profile of its diet can be obtained. Finally, we suggest that future studies of elk nutrition include coupled examinations of diet and dietary quality from Kentucky and other eastern states such as Arkansas, North Carolina, and Pennsylvania where elk herds have been successfully established. This will allow managers to target the plant species that contribute most to maintaining herd health and productivity. Acknowledgments This work was supported by the University of Kentucky, College of Agriculture. The paper is contribution # 05-09-084 of the Kentucky Agricultural Experiment Station and is published with approval of the director. J. Schneider was supported by the graduate program of the Department of Biological and Environmental Sciences, and the Institute for Regional Analysis and Public Policy, Morehead State University. We appreciate the assistance of J. Gassett, J. Day, D. Crank, and C. Logsdon of the Kentucky Department of Fish and Wildlife Resources in facilitating aspects of project administration and field work. We are especially appreciative for the assistance of the staff of the Composition Analysis Laboratory at Colorado State University in helping with the laboratory analyses. Literature Cited Alexy, K.J. 2004. Meningeal worm (Parelaphostrongylus tenuis) and ectoparasite issues associated with elk restoration in southeastern Kentucky. Ph.D. Dissertation. Clemson University, Clemson, SC. 161 pp. Baker, D.L., and N.T. Hobbs. 1982. Composition and quality of elk summer diets in Colorado. Journal of Wildlife Management 46:694–703. Baldwin, W.P., and C.P. Patton. 1938. 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