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2011 SOUTHEASTERN NATURALIST 10(1):53–74
Seasonal Forage Availability and Diet for Reintroduced
Elk in the Cumberland Mountains, Tennessee
Jason L. Lupardus1,2, Lisa I. Muller1,*, and Jason L. Kindall1,3
Abstract - Cervus elaphus (Elk) were reintroduced into the Cumberland Mountains,
Tennessee over a 3-year period beginning in December 2000. We radio-collared 159
Elk and monitored them by aerial telemetry from February 2001 to June 2003. Locations
(n = 321) were used to develop a core herd home range (789-ha sampling area) to
assess Elk seasonal forage use and availability. We monitored diet and resource availability
from November 2003 to October 2004 by microhistological analysis of feces
and vegetation sampling, respectively. We compared the relative availability of individual
plant species (% cover) to the relative percentage of plant species found in fecal
samples. A positive significant mean difference indicated plant species used in greater
proportion to availability. Lolium arundinaceum (Tall Fescue) comprised 35.1% of the
winter diet, and graminoids (65.9%) were the dominant forage class overall. The most
selected graminoid was Andropogon gerardii (Big Bluestem). The diet shifted in the
spring to a mixture of woody plants (28.1%), forbs (19.4%), and graminoids (38.4%).
Carex spp. (sedges) and Juncus spp. (rushes) were the most selected graminoids. The
highest seasonal use of forbs (45%) and legumes (23%) occurred during summer, with
Impatiens spp. (jewelweed; 27%) as the dominant and most selected plant in the diet.
The dominant fall forage class was woody plants (37.4%). Quercus spp. (oaks; vegetation
and acorns 14.3%) were the most used woody plants with oak acorns comprising
9.7% of the Elk diet. We suggest that historic evidence, presence of native grasses, and
Elk diets indicate that oak savannas could be an ideal habitat type for Elk in the reintroduction
zone of Tennessee.
The eastern subspecies Cervus elaphus canadensis Erxleben (North American
Elk) once ranged throughout the southeastern US (Bryant and Maser 1982);
however, that subspecies is now considered extinct (O’Gara and Dundas 2002).
The last remaining North American Elk was reported in Tennessee in the 1860s
(Ganier 1928). Overharvesting and habitat loss/changes were the major contributing
factors for extinction of the subspecies (O’Gara and Dundas 2002). In the
early 1900s, Cervus elaphus L. (Elk) reintroduction programs began in many
eastern states from Louisiana to New Hampshire. Most Elk reintroductions failed
due to crop depredation, disease, and poaching (O’Gara and Dundas 2002). More
recently, Elk reintroductions have occurred in Arkansas, Michigan, Pennsylvania,
Wisconsin, North Carolina, and Kentucky.
1Department of Forestry, Wildlife and Fisheries, University of Tennessee, Knoxville, TN
37996. 2Current address - GA DNR Wildlife Resources Division, Alabany, GA 31721.
3Current address - Ozark Natural Science Center, Huntsville, AR 72740. *Corresponding
author - firstname.lastname@example.org.
54 Southeastern Naturalist Vol. 10, No. 1
In 2000, the Tennessee Wildlife Resources Agency established a 271,145-
ha area for Elk restoration in the Cumberland Mountains of eastern Tennessee
(Fig. 1). Large tracts of continuous forests (87% deciduous) with sparse openings
(12% primarily pasture and reclaimed coal strip mines), lack of production
agriculture (1% cropland), and low human population densities were the primary
factors considered when identifying a restoration zone suitable for Elk in Tennessee
(TWRA 2000). Subsequently, Elk (n = 167) were released onto the Royal
Blue Unit of the North Cumberland Wildlife Management Area (hereafter called
the RB), TN over a 3-year period beginning in December 2000. Elk were translocated
from Land Between the Lakes National Recreation Area, KY and Elk Island
National Park, AB, Canada. Elk from Kentucky were initially populated from the
source herd at Elk Island National Park (Larkin et al. 2004).
Elk behavior, population growth rates, and fecundity are important for determining
the success of reintroduction efforts and, in turn, are influenced by
diet and nutrition (Cook 2002). Diets of Elk have been extensively studied in
the western US, where vegetation primarily consists of coniferous forests, shrublands,
and grasslands (Johnson et al. 2000, Korfhage et al. 1980, Kufeld 1973).
The importance of deciduous forests in a landscape dominated by grasslands
has been described by Clutton-Brock et al. (1982) for C. elaphus (Red Deer) in
Europe; however, Elk diets in US eastern deciduous forests are poorly known.
Additionally, many floral and faunal changes have occurred since Elk were extirpated
from the eastern US. Past research conducted on the Elk diet from the
eastern US occurred in Virginia (Baldwin and Patton 1938), Missouri (Murphy
1963), Michigan (Buss 1967, Spiegel et al. 1963), and Kentucky (Schneider et
al. 2006); however, none assessed seasonal forage use and availability. Therefore,
there was a lack of understanding of the seasonal diet composition of Elk in
eastern forests essential to facilitate long-term sustainability of newly repatriated
populations. In this study, we examined seasonal availability and use of plant
species by Elk in Tennessee.
The study area (789 ha) within the RB (20,235 ha) is located in Scott and
Campbell counties, TN in the southern part of the Cumberland Mountain region
(Smalley 1984). Jackson (2003) reported that the RB was formerly known as the
Koppers Coal Reserve before the 1992 TWRA purchase. Strip, bench, and deep
mining occurred in the 1930s–1950s, 1970s, and 1990s till present as the predominant
means of coal extraction, which has left the area with a latticework of shelves
or benches across the landscape (Jackson 2003). The study area consists of 90%
deciduous forest and 10% grasslands (wildlife openings and mine reclamation
sites; TWRA 2000). Cabrera (1969) described the major community types found
within the deciduous forest as Acer saccharum Marsh (Sugar Maple) - Tulipifera
liriodendron L. (Yellow Poplar) - Tilia Americana L (Basswood) - Aesculus flava
Ait (Buckeye) in north-facing coves, Sugar Maple - Quercus rubra L. (Northern
Red Oak) - Yellow Poplar - Robinia psuedoacacia L. (Black Locust) on north2011
J.L. Lupardus, L.. Muller, and J.L. Kindall 55
and west-facing ridges and coves, and Quercus prinus L. (Chestnut Oak) - Black
Locust on west- and southwest-facing ridges and coves. Most grasslands created
by mining have been reclaimed with Lolium arundinaceum S.J. Darbyshire (Tall
Fescue) and Lespedeza cuneata G. Don (Serecia Lespedeza). Other grasslands
are seasonally planted with mixtures of Lolium spp. (rye grass), Triticum spp.
(wheat), and Trifolium spp. and Melilotus spp. (clovers).
Elevations on the study area range from 450 to 980 m within the RB (400
to 980 m). The 3–5 cm deep topsoils (TVA 1981) were developed from bedrock,
shale, and siltstone, and are classified as acidic, loamy, and well-drained
(Conner 2002). The climate is temperate with mean daily temperatures ranging
from 0.8 °C in January to 24.2 °C in August (NOAA 2003). The annual
mean temperature was 13.1 °C in 2003. The mean annual precipitation for
2003 was 172 cm, approximately 40 cm over the average for the area (NOAA
2003). Flooding and landslides are common in the area due to mining activities,
steep gradients, long slopes, soil-water saturation, and frozen ground that
increases subsurface water flow rates. Most slopes are 40% to 60%, but range
from 10% to 100% (Smalley 1984).
Techniques described in Schemnitz (1994) were used for collecting Elk at
Elk Island National Park by baiting with hay, capturing in corral traps, and moving
animals to a holding facility for processing. A system of gates and squeeze
chutes were used for disease testing, medication (Kreeger 1996), and radio
transmitter application. Elk (n = 94 females and 65 males) were fitted with Lotek
(Lotek Wireless, Inc., Newmarket, ON, Canada) and ATS (Advanced Telemetry
Systems, Isanti, MN) VHF radio transmitters prior to release. We used aerial
telemetry techniques described by White and Garrot (1990) to determine Elk
locations at various times during the day and on different days of the week from
February 2001 to June 2003. Fixed-wing aircraft were equipped with dual, wingstrut
mounted, 2-element, H-antennas for aerial telemetry. A Lotek Suretrack
1000 receiver was used to detect each Elk transmitter. A handheld Garmin GPS
unit (Garmin International, Olathe, KS) was used to record geographic coordinates
when the aircraft was directly over each Elk. Telemetry error was estimated
by locating 30 collars (transmitters) that were placed throughout the RB in locations
unknown to the data collector. Elk handling techniques were approved by
the University of Tennessee Institutional Animal Care and Use Committee (UTIACUC#
Study area delineation
We delineated our study area (Fig. 1) within the RB by developing a 50% kernel
herd home range using 1450 Elk locations from February 2001 to June 2003 (Edge
et al. 1987, Lupardus 2005). All location points for individual animals found exclusively
within the RB were censored from analyses if radio contact was lost, the
56 Southeastern Naturalist Vol. 10, No. 1
collar was found dropped, or if the collar was found on a dead Elk. Therefore, Elk
located exclusively within the RB boundary (n = 321) that remained alive and in
Figure 1. The 789-ha study area (50% kernel home range) and 95% kernel home range
(7100 ha) for reintroduced Elk exclusively using the Royal Blue Unit of the North Cumberland
Wildlife Management Area, TN in the 271,145-ha Elk restoration zone (based
on 321 Elk location points collected from February 2001–June 2003). The 1.6-km-wide
buffer zone was developed by the Tennessee Wildlife Resources Agency where nuisance
Elk will be tolerated.
2011 J.L. Lupardus, L.. Muller, and J.L. Kindall 57
radio contact for the duration of the monitoring were analyzed to estimate 95% and
50% kernel group home ranges using the Animal Movement Extension in Arc View
3.2® (ESRI, Redlands, CA; Hooge and Eichenlaub 1997) for Elk in the RB. The
50% kernel home range was chosen as the study area for sampling because it is considered
as a core center of animal activity; furthermore, the biological significance
of the 95% kernel home range can be unreliable (Dickson and Beier 2002, Seaman
et al. 1999). Additionally, we were restricted to the RB for sampling due to lack of
access in surrounding private lands. Sawyer et al. (2006) also used the distribution
Figure 2. Locations of 150 vegetation sampling points throughout the 3 major cover types
within the 789-ha Elk study area (50% kernel home range), November 2003 to October
2004, Royal Blue Unit of the North Cumberland Wildlife Management Area, TN. Sample
points were developed in Random Point Generator Extension for Arc View 3.2®.
58 Southeastern Naturalist Vol. 10, No. 1
of radio-collared animals to determine the available habitat at the study-area level
(McClean et al. 1998). Our study area included 70% (n = 63; 17 males and 46 females)
of the Elk herd (n = 114).
We determined major habitat types in the study area from 1995 land-cover
maps developed by TWRA (1997) in compliance with the National Gap Analysis
Program. Classification was performed on Landsat TM Imagery. The major
cover types in our study area were deciduous forest and grassland. The importance
of edge habitat to Elk is well documented (Harper 1971, Larkin et
al. 2004, Skovlin et al. 2002, Witmer and deCalesta 1983), and Kufeld (1973)
listed many important plants utilized by western Elk that were classified as disturbance
or edge species. Therefore, we added an additional cover type called
“edge” for the analysis.
Delineating edge locations and widths is difficult due to variation in vegetative
structure, topography, and connectivity, but edges can be functionally defined by
wildlife use (Lidicker 1999, Yahner 1988). Preliminary vegetation sampling and
presence of edge forage species observed during on-site plant surveys were used
to determine edge width. A buffer distance of 10 m around fields, both sides of
roads (unimproved and improved), and grasslands was considered the edge-cover
type, as plant communities differed within this area.
We conducted preliminary sampling in summer 2003 to determine the sample
size necessary for distinguishing differences in vegetation composition at P ≤
0.05. Square 1-m2 samples of percent cover were taken 5 m from the plot center
in the 4 cardinal directions at each randomly placed preliminary plot (n = 23).
We recorded percent cover of forbs, grasses, and seedlings within each 1-m2
sample, and we determined plant species variability from plots (n = 23; σ2 =
34.51). Preliminary data, logistical constraints, and sampling costs were used to
determine the number of vegetation plots sampled (n = 150). We generated random
points throughout the 3 major cover types using Random Point Generator
1.28 (Jenness Enterprises, Flagstaff, AZ) Extension for Arc View 3.2®. The proportional
area of each cover type was used to derive the distribution of sampling
points. Deciduous forest (n = 123), grassland (n = 14), and edge (n = 13) plots
were randomly located within our study area and sampled in proportion to their
areas (Fig. 2). We conducted vegetation sampling seasonally from November
2003 until October 2004 as described by Kufeld (1973): summer (June–August),
fall (September–November), winter (December–February), and spring (March–
May). We sampled each plot once per season.
Major habitat assessments were conducted at each plot as described by Nixon
et al. (1970) to identify all plants that could be consumed by Elk. All species were
identified and percent cover was determined for saplings and shrubs (1-m to 2-m
height) in a 10-m2 square plot surrounding the plot center. In the deciduous forest
and grassland areas, vegetation was sampled within 1-m2 quadrants located 5 m
from the plot center in the 4 cardinal directions. We recorded all forbs, grasses,
2011 J.L. Lupardus, L.. Muller, and J.L. Kindall 59
and seedlings within each 1-m2 sample. Edge sample plots (1 m2) were taken at
5 m and 10 m from the plot center in each linear direction parallel to adjacent
roads or fields (Fig. 3). Ocular estimates for percent cover of each plant species
was conducted by the same person for all samples.
We used a 10-factor prism at each plot center for measuring basal area for
all trees >10 cm dbh to determine the tree-canopy composition and estimate
Figure 3. Illustration of forest, grassland, and edge samples where vegetation measurements
were conducted at each plot within the 789-ha Elk study area (50% kernel home
range), November 2003 to October 2004, Royal Blue Unit of the North Cumberland
Wildlife Management Area, TN (plots and subplots not drawn to scale).
60 Southeastern Naturalist Vol. 10, No. 1
hardwood mast yields. We identified and recorded the number of tree species
at each plot. We used methods described by Whitehead (1969) to estimate the
available acorn crop based on basal area, data from the mast crop survey (TWRA
2005), and mean acorn mass compiled from 14 Quercus spp. (oaks) in the southeastern
US (Long and Jones 1996). Oak acorn availability was estimated from
mean oak basal area (7.21 m2/ha) and the mast crop survey (5.3 out of 10) rating
(Whitehead 1969). A conservative estimate of viable acorns (52% or 17 per
square 1-m2 sample) was determined from Whitehead (1969).
We collected fresh Elk scat when traveling between vegetation plots within
the study area from November 2003 to October 2004. Elk movements, food
mixing by rumination, and distribution of collected scat throughout the study
area provided relatively unbiased samples. Microhistological analysis of feces
provides an efficient and less biased method when compared to visual feeding
observations, pen or restricted feeding, browse surveys, or stomach contents
to determine diet composition. Dearden et al. (1975) developed correction factors
using rumen fluid from domestic ruminants for in vitro digestion to adjust
diet composition because minute plant fragments might be overestimated in
the actual diet. Forb estimates may be lower and grass and woody species
estimates may be higher when correction factors are not applied. However,
other research indicates that correction-factor application may introduce bias
because digestion rates differ among individuals, plants species, and plant
growth stages (Alipayo et al. 1992, Barker 1986, Bartolome et al. 1995, Gill
et al. 1983, Hanley et al. 1985). Therefore, we chose not to apply correction
factors in our study.
Scat was considered fresh on the basis of rich color, moist consistency, and
strong odor (Kirchhoff and Larsen 1998, Weckerly and Ricca 2000). Pellets were
not collected if they were touching surrounding plant tissues. Pellets were placed
in plastic bags and frozen. We analyzed 30 pellet groups during each season (120
samples per year). A minimum of 5 individual pellets from each scat pile was
needed for adequate analysis (Kirchoff and Larsen 1998).
Pellet samples were sent to the Washington State Habitat Lab (Pullman, WA)
for microhistological analysis (Davitt and Nelson 1980). Plant material within
the pellets were described to species when possible. Plant identification and
percentage composition were determined by examining 1 slide of 50 field views
per fecal sample. All plants viewed microscopically were compared to reference
slides. The Washington State Habitat Lab prepared reference slides for this
geographical area from previous studies (Castleberry et al. 2002) and from plant
species collected in the RB.
Vegetation data from all plots were pooled by season to determine the
seasonal availability (% cover) of individual plant species. We used repeated
measures ANOVA to compare the mean availability of individual plant species
2011 J.L. Lupardus, L.. Muller, and J.L. Kindall 61
(independent variable) to the mean percentage of plant material found in fecal
samples (n = 30) to test for disproportionate use (P ≤ 0.05). Tests between forage
availability and diet composition were based on linearly independent pairwise
comparisons between means to determine significant difference each season.
Positive mean differences indicated plants were used in greater proportion than
their availability. Forages were not considered selected when the significant
mean difference (SMD) was negative. “Selection” is here used to describe disproportionate
use of plant species and is relevant only to available forages within
our study site. All data analyses were performed using SPSS version 12.0 (SPSS
Inc., Chicago, IL).
Mean telemetry error was 261 m (n = 30, SE = 24.5). The 95% and 50%
kernel home ranges covered 7100 ha and 789 ha, respectively (Fig. 1). The
50% kernel home range was used as our study area and represented a statistical
center of Elk activity. Our study area was chosen because of size, abundant
Elk activity, and concentration of data points needed for effective habitat sampling
Winter Elk diets had the lowest number of species of plants (n = 45;
Appendix 1). Graminoids (65.9% of plant matter) were the dominant forage
class (Fig. 4), with the most frequently consumed being Tall Fescue (35.1%).
Polystichum acrostichoides Schott (Christmas Fern) comprised 12% of the diet
composition. The largest positive SMD (7.82) found in the diet was Andropogon
gerardii Vitman (Big Bluestem). Other selected graminoids were Tall Fescue,
Schizachyrium scoparium Nash (Little Bluestem), Echinochloa crusgalli Beauv
(Barnyard Grass), Triticum spp. (wheat), and Dactylis glomerata L. (Orchard
Grass). Rubus spp. (briars) were used little when compared to availability (SMD
The spring diet had the highest number of forage items identified compared
to all other seasons (n = 57). Although the diet shifted in the spring to a mixture
of woody plants (28.1%) and forbs (19.4%), graminoids (38.4%) remained the
dominant forage class. Elaeagnus spp. (autumn olive) was the most highly selected
woody plant (SMD = 10.1). Other woody plants selected were Juniperus
virginiana L. (Eastern Red Cedar), oaks, and Pinus spp. (pines). Carex spp.
(sedges) and Juncus spp. (rushes) were the most frequently consumed (12.7 %)
and selected (SMD = 8.4) graminoids. Impatiens spp. (jewelweed) were the only
forbs selected (SMD = 4.6). Acer spp. (maples) were the least selected (SMD =
-5.5) among all forage items overall.
62 Southeastern Naturalist Vol. 10, No. 1
Forb use increased to 45% in the summer. Jewelweed constituted 27% of the
diet and was the most selected forage species (SMD = 24.2). Legumes comprised
23% of the diet, and clovers were highly selected (SMD = 12.2). Big Bluestem
was the only graminoid selected (SMD = 2.4). Autumn olive, oaks, and Lindera
benzoin L. (Spice Bush) were selected woody plants. Briars (SMD = -10.9) and
maples (SMD = -6.2) were the least selected of all used forages.
The dominant forage class in the fall diet was woody plants (37.4%). Oaks
(vegetation and acorns) were the most used woody plant (14.3%), with oak acorns
comprising 9.7% of the diet. Autumn olive was also a highly used (8.7%) and
selected woody plant (SMD = 7.7). Tall Fescue comprised 10.8% of the fall diet
Figure 4. Major seasonal forage classes for Elk diets determined from microhistological
analysis of plant material in feces within the 789-ha study area (50% kernel home range),
November 2003 to October 2004, Royal Blue Unit of the North Cumberland Wildlife
Management Area, TN.
2011 J.L. Lupardus, L.. Muller, and J.L. Kindall 63
and constituted nearly half of the overall graminoid forage class (24%); however,
Tall Fescue was not selected (SMD = -8.8). Big Bluestem was the only graminoid
selected during fall (SMD = 2.48). The combined legume diet composition was
19.3% for clovers and Lespedeza spp. (lespedezas). Briars were the least selected
forage overall (SMD = -10.2).
Elk diets have been extensively documented in the western US for a variety of
habitat types (Cook 2002, Kufeld 1973). The association of Elk with large grasslands
of the central plains and western states could inaccurately classify them
as grazers (Cook 2002). Elk have been classified as intermediate feeders due to
their body size, rumination, and morphology, and perhaps the most accurate description
of Elk food habits is opportunistic (Hofmann 1989). An opportunistic
foraging strategy may facilitate reintroduction.
Grassland areas in our study area were created during reclamation of past
coal mining activities. Topsoil removal, storage activity, grading operations,
and hydroseeding reclamation efforts reduced the native seed bank dramatically
(TVA 1981). Tall Fescue, lespedezas, and autumn olives were the 3 most common
species groups used for reclamation of mined areas and roads in the RB (TVA
1981). Elk selected Tall Fescue during winter probably because it was the only
species available in large quantities; however, Schneider et al. (2006) found Elk
in Kentucky used Tall Fescue heavily during all seasons. Other species of fescue
have been considered valuable forage for Elk in other portions of the species’
range (Cook 2002, Kufeld 1973). Selection for Tall Fescue and other grasses
in the winter diet may be related to costs and benefits of travel to potentially
more nutritious forage (Wickstrom et al. 1984). Many of the graminoids (e.g.,
Big Bluestem, sedges/rushes, Orchard Grass, Little Bluestem) were available
in smaller quantities, but were found in juxtaposition to Tall Fescue. Christmas
Fern was a substantial component (12%) of the winter diet, but it was used less
(SMD = -7.9) than its widespread availability. Elk maximize digestible energy
from changing seasonal forage intake by optimizing time in the rumen to increase
completeness of digestion (Merrill 1994). As such, we believe that during winter
Elk used any available green forage in the area.
Jost et al. (1999) reported that Elk in the Burwash region of Ontario ate jewelweed
in relative proportions to what was available. Williams et al. (2000) found
that jewelweed was highly preferred by Odocoileus virginianus Zimmermann
(White-tailed Deer) in Pennsylvania. Jewelweed was a highly selected plant during
summer in the RB. Jewelweed plants are succulent, and they are primarily
found in cool, damp, closed-canopy areas. Cook (2002) stated that succulent and
nutritious vegetation was used to a great extent by Elk during calving and the subsequent
neonatal period, and Korfhage et al. (1980) suggested that Elk in Oregon
selected forages at their most succulent stage. Peak lactation for Elk yielded up
to 4 L of milk per day (Cook 2002), and water comprised 81% of milk (Robbins
1993). Thus, jewelweed is likely an important component of Elk diets during
64 Southeastern Naturalist Vol. 10, No. 1
summer, when physiological demands require nutritious forage with high water
contents. More research into the nutritional components of native plants will be
fundamental to understanding how Elk and other herbivores utilize plant species
for homeostatic regulation.
We classified clover and lespedeza spp. as legumes rather than forbs to better
understand the dynamics of artificially planted forages in the Elk diet. However,
these items were classified broadly as forbs in other diet studies (Cook, 2002,
Kufeld 1973, Schneider et al. 2006). The forb composition in the diet would have
been substantially higher in the summer (45% to 68%) and fall (10.0% to 29.3%)
if we had categorized legumes as forbs. Our finding of large amounts of clovers
in the summer and fall diets of Tennessee Elk is similar to those of studies conducted
elsewhere (Cook 2002, Kufeld 1973, Schneider et al. 2006). Legumes in
our study were common along roadsides, reclaimed mines, and artificially placed
food plots (TVA 1981).
Some forage items in the diet were not expected. Elk use of lichens and mosses
has been found in Alaska (Kirchoff and Larsen 1998); however, we believe that
the low levels of lichens, mosses, and insects found in the diet were arbitrarily
eaten as Elk foraged on more important foods. We identified Zea mays L. (Corn)
kernels, which was not planted on the study site or in nearby areas (≤8 km), in
the seasonal diets of Elk. The RB is a large area that receives hunting pressure
throughout the year, and we believe that Corn found in the Elk diet was from
baiting and artificial feeding of deer and Meleagris gallopavo L. (Eastern Wild
Turkey) in our study area.
Elk viability in deciduous forests may be somewhat dependent upon the
hard mast (Baldwin and Patton 1938, Murphy 1963). Acorns composed 9.7%
of the fall diet, which could be a substantial nutritional value due to their high
calories and fat content (Kirkpatrick and Pekins 2002). Documented mastdependent
species utilize acorns to increase body fat reserves for the winter
(Pelton 1989, McShea 2000, Wentworth et al. 1992). Hard-mast production
significantly affects natality, mortality, and overall population dynamics
for White-tailed Deer (Wentworth et al. 1990) and Ursus americanus Pallas
(Black Bears) in the southern Appalachians (Pelton 1989). Moreover, interspecific
competition for acorns may influence populations of other species
(McShea and Schwede 1993, Ostfeld et al. 1996). Forage consumption for 1
adult Elk is equivalent to 3 adult White-tailed Deer (Habitat Monitoring Committee
1996), and interspecific competition and dietary overlap has been noted
between Odocoileus spp. (deer) and Elk (Gogan and Barrett 1995, Hobbs et al.
1983, Kirchhoff and Larsen 1998).
Elk herbivory can cause plant community changes (Hobbs 1996, Ripple and
Beschta 2003). Large densities of Elk may reduce habitat quality for other species
(Lindzey et al. 1997). One plant species of concern is oaks. We found that
Elk used oaks in all seasons. Oak regeneration may be limited by the lack of fire
regimes (Van Lear and Waldrop 1990), deer herbivory (Buckley et al. 1998), and
acorn loss from Curculio spp. (acorn weavils), which can range from 10–100%
2011 J.L. Lupardus, L.. Muller, and J.L. Kindall 65
for Quercus alba L. (White Oak) and 2–92% for Quercus rubra L. (Southern Red
Oak) (Gibson 1972, 1982). Elk may function as another competitor that could
affect oak communities.
The importance of maintaining a mature oak forest component was evident
from diet selection. Historically, native Elk in Tennessee foraged in a completely
different type of habitat than what is found today. Ramsey (1853) described portions
of the Cumberland Mountains as oak savannas enriched with deer, Elk, and
Bison bison L. (Bison) in 1783. We found native warm-season grasses throughout
the study site where an arson fire burned nearly 5 years previous. As evidence of
this bygone landscape, we believe that these grasses developed from the remnant
seed bank where past mining activity did not occur. We suggest that the historic
accounts, native grasses, and the Elk diet indicate that oak savannas could be
an ideal habitat type for reintroduced Elk in eastern Tennessee. Periodic fires,
grazing, or drought would maintain oak savannas (Guyette et al. 2002, Komarek
1965). Prescribed burning and other silvicultural techniques should be used to
promote oak regeneration and potentially enhance the quantity and quality of
native grasses and forbs (Van Lear and Waldrop 1990). Additionally, further
monitoring of Elk resource selection is needed to fully understand Elk foraging
dynamics in oak communities.
Our results indicated that Elk used many plant species disproportionately to
their availability. We believe some of the highly used plants in the diet could
be indicative of nutritional needs. Cook (2002) suggested that spring and fall
forages are more important nutritionally than winter forages for Elk. Elk in our
study appeared to opportunistically forage on plants that were different from their
source habitats. Forage opportunities associated with oak forest communities
may promote Elk population growth in eastern Tennessee. As such, we believe
that oak savannas would benefit the Tennessee Elk herd. Knowledge of the seasonal
Elk diet in eastern forests can be used to better assess potential Elk habitat
and understand how future Elk populations will disperse and inhabit new areas
in eastern deciduous forests.
We thank the Rocky Mountain Elk Foundation, Tennessee Wildlife Resources
Agency, and the University of Tennessee, Knoxville for funding and support. We are
grateful for pilots B. Roten (Tennessee Wildlife Resources Agency) and C. Proffit
(Knoxville Flight Training Academy). We want to thank D.S. Buckley, J.D. Clark,
M.A. O’Neil, and J.B. Wilkerson for their support and expertise on various aspects
of this project. We also thank volunteers from the student chapter of the University of
Tennessee Wildlife and Fisheries Society.
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Appendix 1. Mean percentage (x̅ %) of Elk diet and mean difference (MD) between availability (% cover) and percentage of Elk diet. Positive mean differences
represent plants used in greater proportion to availability, and negative mean difference represent plants used in lesser proportion to availability. Diet
composition determined from microhistological analysis of plant material in feces in 789-ha core area (50% kernel home range), and availability determined
from plant composition and cover collected from November 2003 to October 2004, Royal Blue Unit of the North Cumberland Wildlife Management Area,
TN. NA = not applicable because items were not in the diet, unknown, or considered to be arbitrarily eaten (“other” category). *Significance (P < 0.05).
Winter (n = 30) Spring (n = 30) Summer (n = 30) Fall (n = 30)
Plant taxa x̅ % SE MD x̅ % SE MD x̅ % SE MD x̅ % SE MD
Tilia americana L. (American Basswood) 0.0 NA NA 0.7 0.3 0.7 0.0 NA NA 0.0 NA NA
Fagus grandifolia Ehrhart (American Beech) 0.0 NA NA 0.1 0.1 0.0 0.0 NA NA 0.1 0.1 -0.1
Ilex opaca Aiton (American Holly) 0.0 NA NA 0.0 NA NA 0.0 NA NA 0.1 0.1 0.0
Vaccinium spp. (blueberry) 0.5 0.1 -1.2 2.5 0.5 1.2 0.5 0.2 -0.6 0.7 0.2 -0.7
Prunus spp. (cherry) 0.4 0.1 0.2 0.2 0.2 -0.1 0.1 0.1 -0.1 0.1 0.1 0.8
Sambucus canadensis L. (Common Elderberry) 0.1 0.1 -0.1 0.0 NA NA 0.0 NA NA 0.5 0.2 0.5
Aralia spinosa L. (Devil’s Walking Stick) 0.1 0.1 7.8* 0.2 0.2 0.2 0.0 NA NA 0.0 NA NA
Tsuga canadensis L. (Eastern Hemlock) 0.0 NA NA 0.1 0.1 0.0 0.0 NA NA 0.0 NA NA
Cercis canadensis L. (Eastern Redbud) 0.2 0.2 0.6 0.4 0.1 -0.5 0.3 0.3 -0.3 0.7 0.3 0.5
Juniperus virginiana L. (Eastern Redcedar) 0.0 NA NA 3.2 1.1 3.5* 0.0 NA NA 0.3 0.2 0.3
Cornus florida L. (Flowering Dogwood) 0.1 0 0.1 0.0 NA NA 0.8 0.5 0.8 0.6 0.3 0.5
Celtis spp. (hackberry) 0.1 0 0.3 0.0 NA NA 0.0 NA NA 0.0 NA NA
Carya spp. (hickory) 0.5 0.3 1.9* 0.3 0.1 -0.3 0.2 0.1 -0.4 0.1 0.1 -1.3
Magnolia spp. (magnolia) 0.0 NA NA 0.1 0.1 -0.1 0.1 0.1 -0.1 0.5 0.2 0.2
Viburnum acerifolium L. (Maple Leaf Viburnum) 1.0 0.2 0.6 0.1 0.1 -0.3 0.0 NA NA 1.0 0.4 1.0
Acer spp. (maple) 0.2 0.1 -3.3* 0.4 0.1 -5.5* 0.2 0.0 -6.2* 0.7 0.5 -5.3*
Ceanothus americanus L. (New Jersey Tea) 0.3 0.1 0.3 0.6 0.3 0.2 0.0 NA NA 0.2 0.1 -0.4
Quercus spp. (oak) 2.1 0.4 0.8 3.4 0.9 2.5* 3.1 0.5 1.6* 4.6 0.9 3.3*
Quercus spp. (oak acorns) 0.0 NA NA 0.0 NA NA 0.0 NA NA 9.7 2.6 6.7*
Elaeagnus spp. (autumn olive) 1.7 0.7 1.8* 9.7 1.8 10.1* 2.3 0.5 2.4* 8.7 2.5 7.7*
Pinus spp. (pine) 0.9 0.4 1.0 2.0 0.7 2.1* 0.1 0.1 0.1 0.3 0.2 0.4
Rhododendron spp. (rhododendron) 0.1 0.1 0.4 0.9 0.4 0.6 0.0 NA NA 0.7 0.4 -0.5
Rosa spp. (rose) 1.5 0.3 1.2 1.8 0.7 1.7 0.0 NA NA 0.1 0.0 0.0
Sassafras albidum Nutt. (Sassafras) 0.1 0.1 -0.5 0.0 NA NA 0.5 0.3 -1.2 0.0 NA NA
72 Southeastern Naturalist Vol. 10, No. 1
Winter (n = 30) Spring (n = 30) Summer (n = 30) Fall (n = 30)
Plant taxa x̅ % SE MD x̅ % SE MD x̅ % SE MD x̅ % SE MD
Oxydendrum arboreum L. (Sourwood) 0.0 NA NA 0.0 NA NA 0.5 0.3 -0.8 0.0 NA NA
Lindera benzoin L. (Spice Bush) 1.3 0.7 1.5 0.3 0.2 0.4 1.5 0.5 1.6* 3.9 1.0 4.3
Rhus spp. (sumac) 2.6 0.8 1.3 0.1 0.1 -3.0* 0.0 NA NA 3.5 1.0 4.0*
Betula lenta L. (Sweet Birch) 0.0 NA NA 1.0 0.4 1.0 1.0 0.7 0.7 0.0 NA NA
Fraxinus americana L. (White Ash) 0.0 NA NA 0.0 NA NA 0.0 NA NA 0.3 0.3 0.1
Liriodendron tulipifera L. (Yellow Poplar) 0.0 NA NA 0.0 NA NA 0.1 0.1 -2.6* 0.0 NA NA
Unknown woody spp. 1.3 0.2 NA 2.2 0.3 NA 2.0 0.3 NA 3.1 0.3 NA
Total woody spp. 15.1 28.1 13.3 37.4
Heuchera spp. (alumroot) 0.0 NA NA 0.0 NA NA 0.1 0.1 -1.1 0.0 NA NA
Phytolacca americana L. (American Pokeweed) 0.2 0.2 0.0 0.0 NA NA 0.5 0.3 -0.7 0.0 NA NA
Aster spp. (aster) 0.0 NA NA 1.2 0.4 0.6 0.9 0.2 0.4 0.3 0.1 -0.6
Galium spp. (bedstraw) 0.0 NA NA 1.0 0.4 -1.3 0.9 0.3 0.3 0.1 0.1 -0.2
Monarda spp. (bee balm) 0.0 NA NA 0.0 NA NA 0.2 0.2 0.0 0.0 NA NA
Rubus spp. (briar) 0.5 0.3 -21.2* 1.0 0.2 -3.0* 1.3 0.4 -10.9* 1.4 0.2 -10.2*
Centrosema virginianum L. (Butterfly Pea) 1.1 0.3 1.0 1.1 0.5 0.2 2.6 0.7 0.6 1.4 0.4 0.1
Potentilla spp. (cinquefoil) 0.0 NA NA 0.5 0.2 -0.8 1.0 0.5 0.0 0.2 0.1 -0.6
Erigeron annuus L. (Eastern Daisy Fleabane) 0.0 NA NA 0.2 0.1 0.1 0.6 0.2 0.4 0.0 NA NA
Smilacina racemosa L. (False Solomon's Seal) 0.0 NA NA 0.6 0.3 -1.3 0.1 0.1 -0.9 0.0 NA NA
Solidago spp. (goldenrod) 0.1 0.0 -0.1 0.2 0.1 -2.3* 0.6 0.2 -4.0* 0.8 0.3 -3.6*
Smilax spp. (greenbrier) 0.1 0.1 -6.6 0.3 0.2 -3.1* 1.0 0.4 -1.4 0.3 0.2 -3.1*
Impatiens spp. (jewelweed) 0.0 NA NA 7.9 2.6 4.6* 27 3.7 24.2* 0.0 NA NA
Arisaema triphyllum L. (Jack in the Pulpit) 0.0 NA NA 0.0 NA NA 0.7 0.2 -0.1 0.0 NA NA
Chenopodium album L. (Lambs Quarters) 0.0 NA NA 0.1 0.1 0.1 0.0 NA NA 0.2 0.2 -0.1
Ipomoea spp. (morning glory) 0.2 0.2 0.4 0.1 0.1 0.1 0.0 NA NA 0.1 0.1 0.1
Allium spp. (wild onion) 0.0 NA NA 0.1 0.1 0.0 0.3 0.2 0.3 0.0 NA NA
Mitchella repens L. (Partridge Berry) 0.9 0.3 1.0 0.0 NA NA 0.2 0.1 0.1 0.9 0.4 3.3*
Chamaecrista fasciculate Michx. (Partridge Pea) 0.0 NA NA 0.8 0.3 -0.1 0.0 NA NA 0.0 NA NA
Lathyrus spp. (pea) 0.6 0.2 0.5 0.0 NA NA 0.5 0.2 -0.1 0.1 0.1 -0.3
Antennaria spp. (pussytoes) 0.0 NA NA 0.2 0.1 0.1 0.6 0.2 0.6 0.1 0.1 0.0
2011 J.L. Lupardus, L.. Muller, and J.L. Kindall 73
Winter (n = 30) Spring (n = 30) Summer (n = 30) Fall (n = 30)
Plant taxa x̅ % SE MD x̅ % SE MD x̅ % SE MD x̅ % SE MD
Ambrosia spp. (ragweed) 0.0 NA NA 0.1 0.1 0.1 0.3 0.1 -1.3 0.1 0.0 -0.1
Euonymus americana L. (Strawberry Bush) 0.2 0.1 -0.7 0.7 0.4 0.0 0.7 0.2 0.0 0.5 0.3 -0.2
Helianthus spp. (sunflower) 0.0 NA NA 0 NA NA 0.8 0.2 -4.3* 0.0 NA NA
Vicia spp. (vetch) 0.1 0.1 0.1 0.3 0.1 0.2 0.4 0.2 0.4 0.0 NA NA
Geranium maculatum L. (Wild Geranium) 0.0 NA NA 0.9 0.3 -0.2 1.5 0.3 1.5 0.1 0.1 0.1
Unknown forbs 0.6 0.2 NA 2.1 0.4 NA 2.5 0.3 NA 3.4 0.4 NA
Total forbs 4.6 19.4 45 10.0
Polystichum acrostichoides Schott. (Christmas Fern) 12.0 2.1 -7.9* 2.5 1.0 -4.4* 0.2 0.2 -5.1* 0.6 0.4 -7.6*
Athyrium filix L. (Lady Fern) 0.5 0.2 1.1 0.4 0.2 -2.9* 0.3 0.2 -2.0* 0.0 NA NA
Unknown ferns 0.6 0.2 NA 4.3 2.0 NA 0.6 0.1 NA 0.4 0.2 NA
Total ferns 13.1 7.2 1.1 1.0
Echinochloa crusgalli Beauv. (Barnyard Grass) 0.2 0.2 3.8* 0.0 NA NA 0.4 0.4 0.1 0.0 NA NA
Andropogon gerardii Vitman (Big Bluestem) 8.5 1.2 7.8* 4.6 1.1 4.9* 2.3 0.6 2.4* 2.4 0.7 2.5*
Setaria spp. (foxtail) 0.5 0.2 0.7 0.2 0.1 0.2 0.0 NA NA 0.3 0.2 0.3
Microstegium japonicum Nees (Japan Grass) 0.0 NA 0.4 0.0 NA NA 0.0 NA NA 0.1 0.1 -3.3*
Schizachyrium scoparium Nash (Little Bluestem) 4.9 0.8 4.9* 1.4 0.4 1.3 0.2 0.1 0.1 1.1 0.4 1.0
Dactylis glomerata L. (Orchard Grass) 3.3 0.6 3.0* 2.5 0.6 2.6* 1.1 0.4 1.2 1.2 0.5 1.3
Panicum spp. (panic grass) 3.9 0.5 1.5 1.0 0.3 -1.6 0.5 0.2 -2.7* 1.0 0.3 -2.4*
Carex spp. (sedge) and Juncus spp. (rush) 1.4 0.2 -2.8* 12.7 3.1 8.4* 1.3 0.4 -1.3 2.0 0.4 -1.3
Sorghum spp. (sorghum) 0.7 0.3 0.9 0.9 0.2 0.7 0.1 0.1 -0.1 0.8 0.4 0.9
Lolium arundinacea S.J. Darbyshire (Tall Fescue) 35.1 2.8 5.6* 10.7 1.9 -2.8* 5.0 1.2 -4.7* 10.8 2.3 -8.8*
Phleum pratense L. (Timothy Grass) 2.3 0.6 0.4 1.8 0.9 1.8 0.5 0.3 -1.1 0.9 0.3 -0.4
Triticum spp. (wheat) 3.5 0.7 3.0* 1.0 0.4 0.6 0.9 0.4 0.7 1.5 0.6 1.3
Unknown grasses 1.6 0.2 NA 1.6 0.2 NA 1.7 0.3 NA 1.9 0.3 NA
Total Graminoids 65.9 38.4 14.0 24.0
Zea mays L. (Corn) 0.0 NA NA 0.2 0.1 0.2 0.6 0.3 0.7 0.6 0.3 0.7
Total crops 0.0 0.2 0.6 0.6
74 Southeastern Naturalist Vol. 10, No. 1
Winter (n = 30) Spring (n = 30) Summer (n = 30) Fall (n = 30)
Plant taxa x̅ % SE MD x̅ % SE MD x̅ % SE MD x̅ % SE MD
Trifolium and Melilotus spp. (clovers) 0.3 0.1 -4.0* 2.4 0.8 -0.7 15 2.7 12.2* 6.5 1.3 5.0*
Lespedeza spp. (lespedeza) 0.2 0.1 0.2 1.6 0.7 -2.6* 7.5 1.4 0.9 12.8 3.5 5.7*
Total legumes 0.5 4 23 19.3
Lichen/moss 0.1 0.1 NA 0.2 0.1 NA 0.2 0.1 NA 0.4 0.1 NA
Insect 0.1 0.1 NA 0.1 0.1 NA 0.7 0.2 NA 0.2 0.1 NA
Unknown 0.6 0.2 NA 0.2 0.1 NA 2.5 0.3 NA 4.0 0.1 NA
Number of plants 45 57 55 54