Physiological Condition of Female White-tailed Deer in a
Nutrient-deficient Habitat Type
M. Colter Chitwood, Christopher S. DePerno, James R. Flowers, and Suzanne Kennedy-Stoskopf
Southeastern Naturalist, Volume 12, Issue 2 (2013): 307–316
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2013 SOUTHEASTERN NATURALIST 12(2):307–316
Physiological Condition of Female White-tailed Deer in a
Nutrient-deficient Habitat Type
M. Colter Chitwood1,*, Christopher S. DePerno1, James R. Flowers2,
and Suzanne Kennedy-Stoskopf
Abstract - Physiological and morphological indices are useful for determining condition
of Odocoileus virginianus (White-tailed Deer; hereafter deer) and are important
for deer management. However, information about deer condition in nutrient-deficient
habitat types is sparse. Pocosins have a low nutritional plane and are characterized by
deep, acidic, peat soils with a dense shrub layer that provides little or no hard and soft
mast. In July 2008 and March 2009, we collected a total of 60 female deer (30 from
each period) from a 31,565-ha pocosin forest managed intensively for Pinus taeda
(Loblolly Pine) in coastal North Carolina. We recorded whole weight, eviscerated
weight, spleen and adrenal gland weights, and kidney fat index (KFI). Abomasal parasite
counts (APC) and femur marrow fat index (MFI) were determined post-collection
in the laboratory, and blood samples were analyzed for packed cell volume and standard
serum chemistries. Serum chemistries were within expected ranges, with the
exception of elevated potassium concentrations. The KFI and MFI were within levels
reported in the literature, and APC levels did not indicate heavy parasite loads. Spleen
(t58 = 0.69, P = 0.492) and adrenal gland weights (t58 = 1.46, P = 0.151) were similar
between periods. Our results provide baseline physiological data for deer in a nutrientdeficient
habitat type. Though managers need to consider nutritional plane of particular
habitat types, our results indicate that deer can achieve normal body weights and
maintain body condition in nutrient-deficient sites.
Physiological analyses of Odocoileus virginianus Zimmermann (White-tailed
Deer; hereafter deer) based on blood-serum parameters and body-condition indicators
(e.g., kidney fat, femur marrow fat) have been used to evaluate health
and condition. Serum chemistry results have been reported from South Dakota
(Hippensteel 2000, Osborn 1994), Minnesota (Seal and Erickson 1969, Seal et al.
1978), Michigan (Johnson et al. 1968), Oklahoma (DeLiberto et al. 1989), Kansas
(Klinger et al. 1986), Missouri (Tumbleson et al. 1968), Texas (Blankenship
and Varner 1977, Kie et al. 1983, Waid and Warren 1984, White and Cook 1974),
and Maryland (Wilber and Robinson 1958), and body condition results have
been reported from Manitoba (Ransom 1965), South Dakota (Hippensteel 2000,
Osborn 1994), Oklahoma (DeLiberto et al. 1989), Texas (Kie et al. 1983, Waid
1Fisheries, Wildlife, and Conservation Biology Program, Department of Forestry and
Environmental Resources, North Carolina State University, Raleigh, NC 27695. 2Department
of Population Health and Pathobiology, College of Veterinary Medicine, North
Carolina State University, Raleigh, NC 27606. 3Department of Clinical Sciences, College
of Veterinary Medicine, North Carolina State University, Raleigh, NC 27606. *Corresponding
author - email@example.com.
308 Southeastern Naturalist Vol. 12, No. 2
and Warren 1984), and South Carolina (Finger et al. 1981, Johns et al. 1984).
However, physiological data for deer in the Southeast are lacking.
As Quality Deer Management (QDM) has grown in popularity, interest
in individual-level health parameters (e.g., kidney fat, body weight, parasite
load) has increased. Although deer management has focused typically on
population-level parameters (e.g., relative density, sex ratios), individual-level
health parameters can be used by state agencies, private managers, and hunters
to assess the success of management strategies. Thus, an understanding
of physiological condition of deer across their range is warranted. Further,
nutritionally deficient habitat types are of interest because few studies have
examined deer health at sites where nutritional plane is low. Hence, the objectives
of our study were to establish baseline physiological values and
determine the relative health of female deer in a nutritionally deficient habitat
type during the 2 most stressful time periods for deer.
We conducted our study at Hofmann Forest, which is owned and managed
by the North Carolina State Natural Resources Foundation. Hofmann Forest is
a 31,565-ha tract of contiguous pocosin intensively managed for Pinus taeda
L. (Loblolly Pine) production in the Coastal Plain of North Carolina (Jones and
Onslow counties). Pocosins are characterized by deep, acidic, nutrient-deficient
sandy or peat soils (Richardson et al. 1981). Typical pocosins are fire adapted
(15- to 20-year disturbance interval), have temporary surface water (but may
flood for long periods), and maintain a high water table (Christensen et al. 1981,
Richardson et al. 1981). During the study, Hofmann Forest contained 28% natural
pocosin, 52% pine plantation, 10% clearcut, and 2% agriculture (e.g., corn, soybeans,
wheat). In the natural areas, dominant vegetation included Pinus serotina
Michaux (Pond Pine) and a dense shrub layer comprised of Cyrilla racemiflora
L. (Titi), Magnolia virginiana L. (Sweetbay), Persea borbonia (L.) Spreng.
(Redbay), Ilex glabra (L.) A. Gray (Inkberry), and Smilax spp. (Greenbriar)
(Christensen et al. 1981, Richardson et al. 1981). Also, the pine plantations contained
the dense shrub layer characteristic of the natural areas. Pocosins provide
little or no hard mast, and soft mast is limited. Thus, deer are largely dependent
upon browse (Hazel et al. 1978). At their natural climax stage, pocosins represent
a low browse resource with many plants unpalatable and containing low crude
protein and phosphorus, which could affect body maintenance of deer (Smith et
During the study, 9 hunt clubs were active on Hofmann Forest, and their
hunting areas ranged in size from about 445 to 5460 ha. Deer were hunted predominately
using dogs, and harvest records maintained by hunt clubs from 2001
through 2006 indicated a male-biased harvest. On average, hunters harvested
antlered males 74% of the time, and the total deer harvest averaged 430 deer/
year during this time period. Deer harvest was stable throughout and showed
no indication of decline. The North Carolina Wildlife Resources Commission
2013 M.C. Chitwood, C.S. DePerno, J.R. Flowers, and S. Kennedy-Stoskopf 309
(NCWRC) estimated the deer density in the two-county area including Hofmann
Forest was between 6 and 17 deer/km2, with the lower density in pocosins and the
higher density in the agricultural areas outside of Hofmann Forest (R. Norville,
NCWRC, Kinston, NC, pers. comm.).
We shot female deer in the head with high-powered rifles at night in July 2008
and March 2009. Collections corresponded with the 2 most stressful time periods
for female deer in the southern portion of the range (i.e., late summer, during
lactation and late winter, before spring green-up). Within minutes of collapse,
we collected blood via cardiac puncture and stored blood samples on ice until
centrifuged for serum separation, usually within 6 hours of collection. Serum
samples were placed on ice, frozen, and later analyzed by Antech Diagnostics (on
an Olympus AU5400, Melville, NY) for glucose, urea nitrogen (BUN), creatinine,
total protein, albumin, total bilirubin, alkaline phosphatase (ALP), alanine
aminotransferase (ALT), aspartate aminotransferase (AST), cholesterol, calcium,
phosphorus, sodium, potassium, chloride, albumin/globulin ratio, BUN/creatinine
ratio, globulin, and creatine kinase (CK). We measured packed cell volume
(PCV) of whole blood in the field using a hematocrit centrifuge.
We recorded total body weight and collected kidneys with all perirenal fat,
spleens, adrenal glands, fetuses (in March), and the right femur (Hippensteel
2000, Osborn 1994). Eviscerated weights were recorded after all internal organs,
the lower jaw, and the right femur were removed. For abomasal parasite counts
(APC), we randomly selected a deer from the first 6 processed in each time period,
then systematically sampled every 6th deer to obtain a total of 5 deer, which was
the number suggested by Eve and Kellogg (1977). Each abomasum was removed
from the digestive tract and stored on ice until processed (Eve and Kellogg 1977)
by the North Carolina State University College of Veterinary Medicine.
We determined fat reserves using total perirenal fat (KFI: relative to naked
kidney weight; Monson et al. 1974) and femur marrow fat (FMI) using a 2–3
gram sample of marrow from the center third of a cut femur combined with a
2:1 chloroform:methanol solution in the laboratory (Verme and Holland 1973).
We recorded spleen and paired adrenal gland weights (Hippensteel 2000, Osborn
1994) and estimated ages of collected deer by tooth replacement and wear
(Severinghaus 1949) to separate deer into 2 classes: <1.5 years and >1.5 years.
The younger class represented deer that were fawns during the previous breeding
season (relative to our collection periods) and were not likely to have bred,
while the older class represented deer that would have been of breeding age in
the previous fall breeding season.
We evaluated fecundity and breeding season dates by noting lactation status
in July and recording the number and lengths of fetuses collected in March.
We determined reproductive rate from fetal counts (Hesselton and Sauer 1973)
and estimated conception date using a commercially available fetal-aging scale
(Quality Deer Management Association, Bogart, GA; based upon Hamilton et al.
310 Southeastern Naturalist Vol. 12, No. 2
1985). We determined lactation rate by dividing the number of lactating females
by the number of females collected. All research activities were approved by the
NCWRC and the North Carolina State University Institutional Animal Care and
Use Committee (08-082-O).
We defined population health as the combination of the overall condition
of each individual deer (including body weight, fat levels, and serum chemistries)
and the reproductive data from the herd. We used the combination of
physiological and reproductive metrics to avoid basing our health assessment
on just 1 health parameter. Analyses were primarily descriptive, which facilitated
qualitative comparison to the literature and veterinary reference values.
Values falling within reported ranges (e.g., DeLiberto et al. 1989, Johns et
al. 1984, Kie et al. 1983) were accepted as normal. In such cases, we did not
test for seasonal differences because we did not want to confuse statistical
significance with biological significance. However, for metrics with potential
biological significance and few reference values (i.e., spleen and paired
adrenal gland weights), we compared seasonal means with t-tests (α = .05) in
SYSTAT 10 (Systat Software, Chicago, IL).
We collected 30 female deer in July 2008 and 30 in March 2009 with ten
<1.5 years old and three <1.5 years old in July and March, respectively. Serum
chemistry results were obtained for all deer except 1 in the March collection;
chemistries were normal in both seasons, with the exception of high potassium
Table 1. Serum chemistries of female White-tailed Deer collected at Hofmann Forest, NC, July
2008 and March 2009.
July (n = 30) March (n = 29)
Chemistry (units) x̅ SD Range x̅ SD Range
Total protein (g/dL) 7.3 0.44 6.4–8.2 6.3 0.44 5.3–7.0
Albumin (g/dL) 2.5 0.22 2.1–3.0 2.8 0.24 2.3–3.3
Globulin (g/dL) 4.8 0.36 3.9–5.3 3.5 0.38 2.7–4.4
Albumin:globulin ratio 0.5 0.07 0.4–0.6 0.8 0.11 0.5–1.0
Aspartate aminotransferase (U/L) 90 21.5 62–157 87 31.1 47–166
Alanine aminotransferase (U/L) 42 7.6 30–64 29 6.9 11–43
Alkaline phosphatase (U/L) 119 53.5 47–267 71 31.4 24–152
Total bilirubin (mg/dL) 0.2 0.06 0.1–0.3 0.3 0.15 0.1–1.0
Urea nitrogen (mg/dL) 17 6.4 6–33 19 6.3 7–35
Creatinine (mg/dL) 1.1 0.20 0.7–1.6 1.3 0.21 1.0–1.9
Blood urea nitrogen:creatinine ratio 17 6.7 5–31 14 5.3 5–29
Phosphorus (mg/dL) 12.6 1.85 8.9–15.5 9.7 1.65 5.6–13.2
Glucose (mg/dL) 194 75.9 85–333 200 90.2 74–409
Calcium (mg/dL) 10.1 0.65 8.8–11.6 9.6 0.54 8.7–10.9
Sodium (mEq/L) 151 6.2 142–171 144 4.5 139–158
Potassium (mEq/L) 9.7 1.65 5.8–12.0 8.9 1.34 6.5–11.9
Chloride (mEq/L) 107 3.8 101–119 101 2.4 97–105
Cholesterol (mg/dL) 45 7.1 31–58 45 8.6 29–65
Creatine kinase (U/L) 327 335 80–1883 194 142 63–739
2013 M.C. Chitwood, C.S. DePerno, J.R. Flowers, and S. Kennedy-Stoskopf 311
values (Table 1). Total KFI and MFI were appropriately low given the nutritionally
stressful periods in which we collected deer (Table 2). Total and eviscerated
body weights were comparable to deer in the region (K. Huffman, Hofmann Forest
Wildlife Manager, Deppe, NC, pers. comm.; Table 2). Mean spleen (t58 = 0.77,
P = 0.444) and paired adrenal gland (t58 = 1.85, P = 0.070) weights (standardized
by eviscerated body weight) were similar between sampling periods (Table 2).
Mean PCV was 45% (SD = 5.6; range = 38–63; n = 28) in July and 53%
(SD = 6.6; range = 39–70; n = 30) in March. Mean abomasal parasite counts were
low in both seasons, with 440 and 580 worms/L of abomasal content in July and
March, respectively. Three genera were identified (Ostertagia, Trichostrongylus,
Skrjabinagia), though most worms were Ostertagia spp. (possibly O. mossi).
In July, lactation rate was 50% (75% excluding the 10 deer <1.5 years old).
In March, reproductive rate was 1.5 fetuses/female (1.7 fetuses/female excluding
the 3 deer <1.5 years old), and we collected 7 singletons, 16 sets of twins, and 2
sets of triplets. Of the 5 non-gestating females, 3 were <1.5 years old, 1 was old
(estimated age >8 years), and the other was estimated at 2 years of age. Estimated
conception dates ranged from mid-October to mid-December.
Serum chemistry results were consistent with the published literature, with the
exception of high potassium values. High potassium values have been reported
in free-ranging cervids (e.g., Kie et al. 1983, White and Cook 1974, Wilber
and Robinson 1958). However, why deer experience higher than expected potassium
concentrations and how they tolerate levels that would have adverse
consequences in other mammals is not clear (Stringer et al. 2011).
Seasonal variation in fat indices has been described for deer in the southeastern
United States (Johns et al. 1984), with peak fat reserves occurring in
winter (DeLiberto et al. 1989, Finger et al. 1981, Stockle et al. 1978, Waid and
Warren 1984). However, our sampling periods were not designed to capture
peak reserves. Rather, our sampling periods reflect the most stressful periods
encountered by female deer in pocosins. Forage limitations before spring greenup
(March) and during the late-summer stress period (July), coupled with the
physiological demands of reproduction (gestation in March and lactation in July)
Table 2. Body parameters of female White-tailed Deer collected at Hofmann Forest, NC, July 2008
and March 2009.
July (n = 30) March (n = 30)
Parameter (units) x̅ SD Range x̅ SD Range
Kidney fat index (%) 25.2 25.79 3.0–116.6 32.4 26.62 2.8–110.6
Marrow fat index (%) 33.8 27.00 2.1–85.4 78.5 53.44 10.0–204.2
Spleen weight (g/kg)A 8.22 1.78 5.18–13.72 7.87 1.73 5.2–14.43
Adrenals weight (g/kg)A 0.15 0.048 0.04–0.26 0.13 0.037 0.07–0.25
Whole body weight (kg) 39.8 6.75 27.2–54.5 39.4 7.99 22.2–50.8
Eviscerated weight (kg) 27.9 4.49 20.4–40.9 28.1 4.92 17.3–36.8
AGram weights standardized by kg of eviscerated body weight.
312 Southeastern Naturalist Vol. 12, No. 2
could require mobilization of stored fat. Further, in the Southeast, hard mast provides
a critical energy and fat source in fall, which yields highest fat deposition
in winter. However, it is possible that deer in pocosins simply cannot deposit
copious amounts of fat due to unavailability of hard mast. Our total KFI and MFI
were comparable to values reported from South Carolina (Johns et al. 1984) and
Texas (Kie et al. 1983, Waid and Warren 1984) but lower than those reported
from northern regions (DeLiberto et al. 1989, Hippensteel 2000, Osborn 1994).
We believe the KFI and MFI values we observed appear to confirm that large fat
reserves might not be necessary for White-tailed Deer survival in the southern
range (Finger et al. 1981).
Spleen (Aiton 1938, Hippensteel 2000, Osborn 1994) and paired adrenal gland
weights (Hippensteel 2000, Osborn 1994, Welch 1962) were greater (≈60–110%
and ≈7–75%, respectively) than those reported in other studies. Increased spleen
and adrenal gland weights have been linked to increased stress levels resulting
from social stress caused by high population density (Aiton 1938, Christian 1959,
Christian and Davis 1964, Christian et al. 1960). However, few studies of Whitetailed
Deer physiology (e.g., Aiton 1938, Hippensteel 2000, Osborn 1994, Welch
1962) have reported spleen or adrenal glands weights, so regional comparisons
are difficult. Without data from other seasons and hormone concentrations, we
cannot determine if the weights we observed in July and March were normal (but
simply higher than other published values) or elevated due to stress. Thus, our
results establish baseline values for these metrics, but further study is required
to adequately determine the seasonal relationship of these values to deer physiological
Mean PCV from both seasons did not indicate anemia, supporting our assertion
that APC levels in our study were not pathogenic. According to Eve and
Kellogg (1977), our APC values indicate a low probability of deer overpopulation
relative to habitat quality. Although seasonal variation has been documented
(Baker and Anderson 1975, Eve and Kellogg 1977, Moore and Garner 1980), our
APC values are consistent with data available from the Southeast (Demarais et
al. 1983, Monschein 1977), suggesting our parasite numbers are unlikely to have
an adverse impact on overall health. Additionally, we did not detect Haemonchus
contortus, a large stomach worm, which has been implicated as a major pathogen
for White-tailed Deer, particularly in the Coastal Plain of the Southeast (Davidson
et al. 1980, Prestwood et al. 1973).
Lactation and reproductive rates determined in this study indicate adequate
productivity spanning 2 separate breeding seasons. The high reproductive rate
and prevalence of twins and triplets suggest the productivity of the population is
higher than might be expected from a nutritionally deficient habitat type. Conception
dates spanned 2 months, which could be explained by hunter bias toward
harvesting males. Fewer breeding males could contribute to females not being
bred in their first estrus of the breeding season. The lactation rate in July was 75%
(excluding the 10 deer less than 1.5 years old), but some females may have lost fawns due
to predation or malnutrition. Canis latrans Say (Coyote) have been increasingly
implicated in fawn mortality in the Southeast (Kilgo et al. 2010), and Coyotes
2013 M.C. Chitwood, C.S. DePerno, J.R. Flowers, and S. Kennedy-Stoskopf 313
were present at Hofmann Forest and seen or heard commonly. However, our
study did not evaluate predation mortality on fawns. Further, though our study
did not quantify recruitment, there was no local evidence to suggest inadequate
reproduction or recruitment.
Considering that pocosin soil and vegetation are nutrient-deficient and our
collections occurred during the 2 most nutritionally stressful time periods for
female deer, we believe that deer from pocosins are finding adequate nutrition
and the habitat type is not necessarily deficient from a deer health perspective. It
is possible that deer benefit from herbaceous forages that become more available
after silvicultural practices (e.g., clearcutting and thinning). Herbaceous forages
typically are more nutritious and digestible than browse, but the extent of this
benefit is unknown. Furthermore, agriculture is prevalent around the boundary
of Hofmann Forest, but many deer do not have access to agricultural areas to
supplement their diet. Though our reproductive results, combined with the other
physiological parameters, do not suggest that deer are nutritionally constrained
by overpopulation, we speculate that deer condition could be dramatically lowered
if population density was allowed to increase, thereby limiting access to
Though pocosins are characteristically described as nutrient-deficient, our results
indicate that deer are obtaining adequate nutrition, which implies that land
managers must consider deer health on the natural nutritional plane of any habitat
type before establishing management strategies that are meant to improve deer
health. We are not suggesting managers need complete necropsies to evaluate
the condition of every individual deer at a site, but continued use of informative
metrics (e.g., kidney fat, body weight, parasite load) are easily available from
hunter-harvested deer. Combined with population-level metrics (e.g., sex ratio,
density), individual-level metrics can be important indicators of deer condition
even on low productivity sites with poor soils. Thus, state agencies and deer
managers should benefit from deer physiological data as hunters and private land
managers become increasingly focused on evaluating the health of deer under
QDM-type programs in varying habitat conditions throughout the White-tailed
Funding was provided by the North Carolina State Natural Resources Foundation,
the North Carolina State University (NCSU) Department of Forestry and Environmental
Resources, and the NCSU Fisheries, Wildlife, and Conservation Biology Program. We
thank the North Carolina Wildlife Resources Commission for help with deer collections
and the NCSU College of Veterinary Medicine for lab space and consultation. We thank
E. Stanford and R. Norville for organizational and field support. We thank technicians
J.H. Harrelson and A. Partin for help in the field and the lab. In addition, we thank the
undergraduate and graduate students of the NCSU Fisheries, Wildlife, and Conservation
Biology Program for help in the field. We thank K.V. Miller, R.G. Osborn, R.A. Lancia,
M.A. Lashley, and all anonymous referees for providing helpful comments on earlier
drafts of the manuscript.
314 Southeastern Naturalist Vol. 12, No. 2
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