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. A preliminary study of food habits of elk in
Virginia. Transactions of the North American Wildlife Conference 3:747–755.
Braun, E.L. 1950. Deciduous Forests of Eastern North America. Hafner, New
York, NY.
2006 J. Schneider et al. 545
Cook, J.G. 2002. Nutrition and food. Pp. 259–349, In D.E. Toweill and J.W. Thomas
(Eds.). North American Elk: Ecology and Management. Smithsonian Institution
Press, Washington, DC.
Cox, J.J. 2003. Community dynamics among reintroduced elk, white-tailed deer, and
coyote in southeastern Kentucky. Ph.D. Dissertation. University of Kentucky,
Lexington, KY. 292 pp.
Foppe, T.M. 1984. Microhistological technique training program. Composition
Analysis Laboratory. Range Science Department, Colorado State University, Ft.
Collins, CO. 28 pp.
Hansen, R.M., and R.C. Clark. 1977. Foods of elk and other ungulates at low
elevations in northwestern Colorado. Journal of Wildlife Management 41:76–81.
Hansen, R.M., T.M Foppe, M.B. Gilbert, R.C. Clark, and H.W. Reynolds. 1978. The
microhistological analyses of feces as an estimator of herbivore diet. Composition
Analysis Laboratory, University of Colorado. 8 pp.
Harper, J.A. 1971. Ecology of Roosevelt elk. PR W-59-R. Oregon State Game
Commission, Portland, OR. 44 pp.
Hill, J.D. 1976. Climate of Kentucky. University of Kentucky Agricultural Experiment
Station, Progress Report No. 221, Lexington, KY.
Hinnant, R.T., and M.M. Kothmann. 1988. Collecting, drying, and preserving feces
for chemical and microhistological analysis. Journal of Range Management
41:168–171.
Hofman, R.R. 1985. Digestive physiology of the deer: Their morphophysiological
specialization and adaptation (deer digestive system). Pp. 393–408, In P.F.
Fennessey and K.R. Drew (Eds.). Biology of Deer Production. Bulletin 22. Royal
Society of New Zealand, Dunedin, New Zealand.
Hoffman, R., and K. Kearns. 1997. Wisconsin Manual of Control Recommendations
for Ecologically Invasive Plants. Wisconsin Department of Natural Resources.
Madison, WI. 102 pp.
Holchek, J.L., and R. Valdez. 1985. Magnification and shrub stemmy material influences
on fecal analysis accuracy. Journal of Range Management 38:350–352.
Irwin, L.L. 2002. Migration. Pp. 493–513, In D.E. Toweill and J.W. Thomas (Eds.).
North American Elk: Ecology and Management. Smithsonian Institution Press,
Washington, DC.
Kufeld, R.C. 1973. Foods eaten by the Rocky Mountain elk. Journal of Range
Management 26:106–113.
Larkin, J.L. 2001. Demographic and spatial characteristics of a reintroduced elk
population. Ph.D. Dissertation. University of Kentucky, Lexington, KY. 146 pp.
Larkin, J.L., D.S. Maehr, J.J. Cox, M.W. Wichrowski, and D.R. Crank. 2002. Factors
affecting reproduction and population growth in a restored elk (Cervus elaphus
nelsoni) population. Wildlife Biology 8:9–14.
Larkin, J.L., J.J. Cox, M.W. Wichrowski, D. Bolin, and D.S. Maehr. 2003. Demographic
characteristics of a reintroduced elk population. Journal of Wildlife
Management. 67:467–476.
Maehr, D.S., R. Grimes, and J.L. Larkin. 1999. Initiating elk restoration: The
Kentucky case study. Proceedings of the Annual Conference of the Southeastern
Association of Fish and Wildlife Agencies 53:350–363.
Maehr, D.S., J.J. Cox, and J.L. Larkin. In press. Elk or Wapiti (Cervus elaphus). In
M. Trani-Griep (Ed.). The Land Manager’s Guide to Mammals of the South, The
Nature Conservancy, Arlington, VA.
546 Southeastern Naturalist Vol. 5, No. 3
McFarlan, A.C. 1943. Geology of Kentucky. Waverly Publishing, Baltimore, MD.
McInnis, M.L., M. Vavra, and W.C. Krueger. 1983. A comparison of four methods
used to determine the diets of large herbivores. Journal of Range Management
36:302–306.
Nelson, J.R., and T.A. Leege. 1982. Nutritional requirements and food habits. Pp.
323–367 In J.W. Thomas and D.E. Toweill (Eds.). Elk of North America: Ecology
and Management. Stackpole Books, Harrisburg, PA.
O’Gara, B.W., and R.G. Dundas. 2002. Distribution: Past and present. Pp. 67–119,
In D.E. Toweill and J.W. Thomas (Eds.). North American Elk: Ecology and
Management. Smithsonian Institution Press, Washington, DC
Overstreet, J.C. 1984. Robinson Forest Inventory: 1980–1982. University of Kentucky,
College of Agriculture, Department of Forestry, Lexington, KY.
Phillips, J. 1997. Technical proposal for free-ranging elk in Kentucky. Kentucky
Department of Fish and Wildlife Resources, Frankfort, KY.
Rehder, A. 1940. Manual of Cultivated Trees and Shrubs Hardy in North America
Exclusive of the Subtropical and Warmer Temperate Regions. 2nd Edition.
Macmillan, New York, NY. 996 pp.
Rentfleish, J.A., and H.H. Hansen. 2000. Microhistology in the ENRECA Project.
Pp. 201–206, In Proceedings of the Integrated Livestock-Crop Production Systems
in the Smallholder Farming Systems in Zimbabwe Workshop. Harare,
Zimbabwe, 10–13 January 2000.
Sites, R. 1995. Coal and Virginia. Virginia Department of Mines, Minerals and
Energy. Available online at http://www.mme.state.va.us/Dmr/PUB/Brochures/
coal.html. Accessed 18 January, 2006.
Sparks, D.R., and J.C. Malechek. 1968. Estimating percentage dry weight in diets
using a microscopic technique. Journal of Range Management 21:264–265.
Ter Beest, J. 2005. Effects of elk on forest soils and succession in eastern Kentucky.
M.Sc. Thesis. University of Kentucky, Lexington, KY.
Thill, R.E., H.F. Morris, Jr., and A.T. Harrel. 1990. Nutritional quality of deer
diets from southern pine-hardwood forests. American Midland Naturalist
124:413–417.
Ulack, R., K. Raitz, and G. Pauer (Eds.). 1998. Atlas of Kentucky. University of
Kentucky, Lexington, KY.