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2008 SOUTHEASTERN NATURALIST 7(4):595–606
Soil Region Effects on White-tailed Deer Forage
Phillip D. Jones1,*, Stephen Demarais1, Bronson K. Strickland1,
and Scott L. Edwards2
Abstract - Body mass and antler development of Odocoileus virginianus (Whitetailed
Deer) vary by soil resource region in Mississippi, but a causative link from
soil to deer morphology has not been established. We investigated crude protein (CP)
content of 8 important deer forages in 5 soil-resource regions to determine if regional
differences in available nutrition could potentially explain some variation in morphometrics.
Crude protein levels of a species composite and all but 1 individual forage
species decreased from spring to summer. Every species exhibited regional differences
in spring, and only 1 species did not vary by region in summer. Composite CP
also differed by region. Differences in potential nutritional planes among regions
may be substantial enough to impact lactation success, fawn recruitment, and body
growth. Directly sampling seasonal diet quality among regions and comparing nutritional
planes among deer herds of different densities may further explain regional
Mississippi is commonly divided into 10 soil-resource regions based on
differences in soils, topography, and climate (Pettry 1977). Odocoileus virginianus
Zimmerman (White-tailed Deer; hereafter, “deer”) differ in body
mass and antler development among soil-resource regions (Jacobson 1984;
Strickland and Demarais 2000, 2006), but a direct nutritional link via forage
quality has not been established. Soil fertility is correlated with plant
biomass production (Biere 1995, Chapin 1980, Fraser and Grime 1998),
and mineral soil richness is correlated with deer body mass (Jacobson 1984,
Smith et al. 1975a). Factors other than soil fertility, however, can impact nutrition
levels and thus development in deer. Higher herd densities may lead
to reduced diet quality as deer are forced to consume less nutritious forage
(Kie et al. 1980); management actions which favor more nutritious plants
may increase the nutritional plane without altering the quality of individual
plant species (Jones 2008).
Different soils infl uence protein levels of deer forages (Hundley 1959,
Kreuger and Donart 1974, Pettorelli et al. 2001, Thorsland 1966). The
purpose of our study was to determine whether several common deer forages
differed in crude protein (CP) content among sites in 5 soil regions
in Mississippi differing in fertility (Jacobson 1984, Pettry 1977) and deer
1Forest and Wildlife Research Center, Box 9690, Mississippi State, MS 39762. 2Mississippi
Department of Wildlife, Fisheries and Parks, Box 9690, Mississippi State,
MS 39762. *Corresponding author - firstname.lastname@example.org.
596 Southeastern Naturalist Vol. 7, No. 4
morphometrics (Strickland and Demarais 2000). If basic differences in
forage quality do occur among regions, they might partly account for differences
in body size and antler development associated with these regions.
We collected forage plant samples from 4 state Wildlife Management
Areas (WMAs), and 3 private properties representing 5 soil regions
(Fig. 1). We chose these regions to correspond with Strickland and Demarais’
(2000) study of regional deer morphometrics in Mississippi. Delta
samples were taken from Sunflower WMA in Sharkey County, where the
Sharkey-Alligator-Dowling soil association was ubiquitous in coverage.
The Upper Thick Loess (Thick Loess) samples were collected on
Figure 1. Sites () sampled for deer forage plant quality in 5 soil resource regions of
Mississippi during spring and summer 2006.
2008 P.D. Jones, S. Demarais, B.K. Strickland, and S.L. Edwards 597
Malmaison WMA in Grenada and Carroll counties on areas dominated
by Memphis association soils. Upper Thin Loess (Thin Loess) samples
were collected on private property in Attala County; Gillsburg soils were
prominent on lower slopes, Hills-Providence soils on uplands. The Upper
Coastal Plain (UCP) samples were collected on Choctaw WMA in Choctaw
and Winston counties. Soil associations included a variety of sandy, silt,
and clay loams, including Susquehanna, Ruston, Pheba, and Collins. The
Lower Coastal Plain (LCP) samples were collected on Wolf River WMA
and two industrial forest sites in Lamar, Perry, and George counties. Soils
included the McLaurin-Heidel-Prentiss, McLaurin-Savannah-Susquehenna,
and Prentiss-Rossella-Benndale associations.
We selected 8 forage species important to deer in Mississippi (Warren
and Hurst 1981), representative of vines, forbs, and browse which we expected
to be available statewide. We collected 5 independent sets of samples
of each species found from each study site in spring (April) and summer
(18 July–15 August) 2006. Sites were sampled in order of average date of
last spring freeze. Each set of samples included enough plant material to
yield ≥50 g wet weight per species. Samples included all leaves and growing
stem tips from selected plants. We selected individual plants with little
or no evidence of depredation or disease. Samples were not collected from
locations where fertilizer may have been recently used, such as food plots
or agricultural fields. We dried samples in a forced-air oven for 72 hours at
60 ºC, then tested for CP on a dry-matter basis using the Kjeldahl procedure
We compared species individually, both by regions and between seasons,
and by season and among regions, using 2-way analysis of variance with
PROC MIXED (SAS Institute 2000). We also averaged CP across species successfully
collected at all 5 sites within each season to create a composite CP,
which we tested for overall effects of region and season using 2-way analysis
of variance with PROC MIXED (SAS Institute 2000). We tested assumptions
of homogeneity of variance before each analysis and used heterogeneous
variance models that accounted for differing variances among fixed effects
when necessary (Littell et al. 2006). We used LSMEANS SLICE to identify
region effects within seasons and season effects within regions following a
significant interaction (Littell et al. 2006). When differences were found, we
conducted pair-wise tests using Fisher’s protected LSD (Carmer and Swanson
1973, Peterson 1985). We considered differences significant if P ≤ 0.05.
We successfully collected 6 of 8 species on all sites and in both seasons;
1 species was collected on 4 sites, and 1 other on 2 sites (Table 1). Seven
598 Southeastern Naturalist Vol. 7, No. 4
Table 1. Crude protein content (SE) of selected White-tailed Deer forages in 5 soil regions of Mississippi collected in 2006.
Site Season x
Species Season Delta Thick Loess Thin Loess UCPB LCPB RegionC region
Campsis radicans Seemann Spring 24 ABD (2) 27 A (1) 16 C (1) 26 A (1) 22 B (1) <0.001 <0.001
(Trumpet Creeper) Summer 12 BE (0) 11 B (1) 9 C (1) 17 A (2) 8 C (0) <0.001
Lonicera japonica Thunberg Spring 14 A (0) 13 BC (0) 12 CD (1) 12 D (0) 14 AB (0) <0.001 <0.001
(Japanese Honeysuckle) Summer 15 AE (0) 8 BE (0) 8 BE (0) 9 BE (1) 8 BE (1) <0.001
Rubus trivialis Michaux Spring 18 A (0) 18 A (1) 15 B (1) 16 AB (1) 18 A (1) 0.002 0.001
(Southern Dewberry) Summer 10 AE (0) 10 ABE (1) 11 AE (1) 9 BCE (0) 8 CE (0) <0.001
Smilax glauca Walter Spring 21 A (1) 24 A (1) 15 B (2) 19 B (0) 11 C (0) <0.001 <0.001
(Sawbrier) Summer 10 BE (0) 10 AE (0) 9 BCE (0) 10 ABE (0) 9 CE (0) 0.011
Ambrosia artemisiifolia Linnaeus Spring 20 B (1) 21 B (1) 26 A (1) 22 B (2) 23 AB (1) 0.004 0.029
(Common Ragweed) Summer 20 (1) 21 (0) 20 e (2) 20 (1) 20E (1) 0.844
Desmodium ciliare Willdenow Spring 24 A (2) 18 B (1) <0.001 0.629
(Tickclover) Summer 20 AE (1) 14 BE (0)
Phytolacca americana Linnaeus Spring 30 C (1) 37 A (1) 28 C (1) 34 B (0) 24 D (1) <0.001 <0.001
(Pokeweed) Summer 21 BE (1) 26 AE (1) 27 A (2) 23 BE (1) 22 B (1) <0.001
Vaccinium arboreum Marshall Spring 21 A (1) 8 C (0) 13 B (1) 12 B (0) <0.001 <0.001
(Sparkleberry) Summer 8 AE (0) 7 AB (0) 7 BCE (0) 6 CE (0) <0.001
AP-values correspond to least-square means. Degrees of freedom for Trumpet Creeper, Japanese Honeysuckle, Dewberry, Sawbrier, Ragweed, and Pokeweed
were: Region = 4,40, Season = 1,40, Interaction = 4,40; for Tickclover: Region = 1,16, Season = 1,16, Interaction = 1,16; for Sparkleberry Region = 3,32,
Season = 1,32, Interaction = 3,32.
BUCP = Upper Coastal Plain, LCP = Lower Coastal Plain.
CWhen Season×Region is significant, Region P-values are for within-region comparisons.
DMeans within a row followed by the same letter are not different (α = 0.05).
ESeasonal effect within region (α ≤ 0.05).
2008 P.D. Jones, S. Demarais, B.K. Strickland, and S.L. Edwards 599
species showed region x season interactions; Desmodium ciliare (Tickclover)
showed differences due to both region and season. In spring, CP differed
among regions for all species; in summer, all species except Ambrosia artemisiifolia
(Ragweed) showed regional differences. In spring, 5 of 7 species
collected from the Thick Loess were ranked in the group with the highest
CP, followed by 4 of 6 in the Delta, 4 of 8 in the UCP, 3 of 8 in the LCP, and
1 of 7 in the Thin Loess. In summer, the Thick Loess placed 5 of 7 species
in the highest grouping, followed by 4 of 7 in the Thin Loess, 3 of 6 in the
Delta, and 4 of 8 in UCP. The LCP placed only Ragweed in the highest CP
grouping, and had the numerically lowest CP in 6 of 8 species.
Most species decreased in CP from spring to summer, though not
necessarily across all sites (Table 1). Ragweed decreased at 2 of 5 sites;
Phytolacca americana (Pokeweed) 3 of 5, and Vaccinium arboreum (Sparkleberry)
at 3 of 4. Lonicera japonica (Japanese Honeysuckle) increased by
1% CP in the Delta and decreased in all other regions by a mean of 4.5%
CP. All other species showed consistent decreases from spring to summer on
every site from which they were collected.
Composite CP was affected by season (F1,290 = 77.60, P ≤ 0.001; Table 2).
Spring composite CP averaged 1.5 times greater than summer (range =
1.3–1.6), refl ecting an average decrease of 6.6% CP (range = 4.6–8.7% CP)
from spring to summer. Region affected composite CP (F4,290 = 2.94, P =
0.021) consistently across seasons (F4,290 = 0.62, P = 0.647). The Thick Loess
provided the highest CP level, the LCP the lowest, differing by 3.5% CP.
Although digestible energy (DE) is a possible limiting factor for deer
(Meyer et al. 1984, Parker et al. 1999), we elected to test CP because of
lower reported nutritional carrying capacity estimates for diets based on CP
requirements than on DE requirements in the Mississippi Lower Coastal
Plain (Jones 2008). Nitrogen requirements for growth (French et al. 1956,
Table 2. Composite crude protein levels of 6 deer forage species common to 5 soil resource
regions in Mississippi, 2006.
Season Delta LCPA UCPA Loess Loess All
Spring Mean 21 19 21 23 19 21
(SE) (1) (1) (1) (1) (1) (1)
Summer Mean 15 12 15 15 14 14
(SE) (1) (1) (1) (1) (1) (1)
Combined Mean 18 BCB 16 C 18 AB 19 A 16 BC
(SE) (1) (1) (1) (1) (1)
AUCP = Upper Coastal Plain, LCP = Lower Coastal Plain.
BMeans within a row followed by the same letter are not different (α = 0.05).
600 Southeastern Naturalist Vol. 7, No. 4
Holter et al. 1979, McEwen et al. 1957, Ullrey et al. 1967) and antler development
(Asleson 1996) are well documented in the literature. Additionally,
body mass of deer in Oklahoma was reported as greater in areas with greater
dietary nitrogen (Soper et al. 1993).
We assumed that composite CP value represented potential forage quality
within each region. Deer are selective foragers (Cote et al. 2004, Crawford
1982, Weckerly and Kennedy 1992) capable of discriminating among forages
to meet nutritional requirements (Berteaux et al. 1998, Vangilder et
al. 1982). While our samples did not account for all possible forages, we
believe they are indicative of real differences among soil regions, especially
since the composites used identical species across all regions. The range
from 16–19% composite CP could potentially impact nutritional plane and
habitat quality for White-tailed Deer and may partly explain the variation in
deer morphometrics among soil-resource regions reported by Strickland and
Deer may select forest clearings with greater biomass of high-quality forage
(Beckwith 1964) even if overall forage biomass is less than other areas
(Stewart et al. 2000). Greater availability of high-quality forage combined
with selective foraging could potentially overcome reduced composite CP
where deer herds are well below carrying capacity. However, access to alternative
forages would be limited in herds at or near carrying capacity (Kie
et al. 1980).
The Thick Loess site appeared to produce the highest potential nutritional
plane, followed by the UCP, Delta, Thin Loess, and LCP. Strickland
and Demarais (2000) reported greater body weights and antler development
in the Delta, followed by the loess regions, the UCP, and the LCP. They
assumed this pattern was related to regional variation in soil fertility. The
discrepancy between their rankings of morphometrics and ours of potential
nutritional plane might be explained in that models explaining deer growth
and antler size in these regions often contained a positive correlation with the
nearby acreage of agricultural fields (Strickland 2005), an abundant source
of high-quality forage. The Delta is the most heavily agricultural region in
Mississippi, containing about 44% of the state’s total cropland (National
Agricultural Statistics Service 2004), so it is likely deer diets in the Delta are
infl uenced by agricultural crops. Comparisons of deer body measurements
and soils in Missouri found deer from prairie soil regions were heavier and
consumed greater amounts of cultivated crops than deer from other regions
(Murphy and Porath 1969), and that antler characteristics were positively
correlated with area of harvested cropland (Kissell et al. 2002). Additionally,
Strickland and Demarais (2000) combined the 2 loess regions into a single
entity; averaging our results for those regions might reasonably result in the
same order as theirs.
Seasonal differences in CP are common among deer forages (Fuller
1980, Meyer and Brown 1985, Smith et al. 1956, Thorsland 1966), and our
2008 P.D. Jones, S. Demarais, B.K. Strickland, and S.L. Edwards 601
results reflect predictable seasonal growth cycles (Chapin 1980, Mattson
1980). Protein levels fell in nearly all forages from spring to summer, reducing
but not eliminating regional differences in composite CP. Because
our samples were limited to 1 spring and 1 summer sample, it is possible
that species were not always sampled at either their greatest or lowest CP
within each region. If not, it is possible that seasonal differences may be
either greater or lesser than what we detected. However, we sampled in accordance
with long-term weather data such that spring samples were taken
in order of average latest freeze date, and followed the same order for summer
sampling. Thus, species were sampled at similar phenological stages
Comparing seasonal protein needs with seasonal protein levels in forage
reveals potential deficiencies, especially in the LCP. Fawning dates in
Mississippi range from late June to early September (Jacobson et al. 1979),
making does dependent on summertime forage to provide nutritional requirements
for lactation. A diet level of 14% CP is minimal for lactating does
(Murphy and Coates 1966, Verme and Ullrey 1984). On the LCP site, this
need could only be met by foraging more selectively on higher quality plants
which exhibited summer CP values ≥14%, such as Pokeweed, Tickclover,
and Common Ragweed; does on sites in the other 4 soil regions could afford
to forage more generally and still maintain sufficiently high diet quality.
Average fetal rates of does ≥2.5 years old do not differ among regions (B.K.
Strickland, unpubl. data); however, data from 4 Mississippi WMAs showed
fawn recruitment on the LCP site to be less than half that in the Delta and
Upper Thick Loess sites (McDonald 2003). Deficiencies in summertime
nutrition for lactating does may impact fawn survival and limit their recruitment
in the LCP.
Growth potential for young deer may also be infl uenced by regional forage
quality. Fawns require from 15–25% CP for optimal growth (French et
al. 1956, Smith et al. 1975b, Ullrey et al. 1967). All sites had springtime CP
levels within this range (Table 3); however, mean values differed by up to
4.7% CP, suggesting biologically meaningful differences in spring forage
quality among regions. Lambert (1998) documented greater mass gain in
fawns receiving higher protein diets during the 6 months following weaning,
a period equivalent to winter and spring in Mississippi. Newly weaned
male fawns gained mass faster on a 20.2% CP diet than on a 12.7% CP diet
(Ullrey et al. 1967). Holter et al. (1979) tested weight gain in yearlings
given experimental diets from 7.9–24.0% CP and found the percentage of retained
body nitrogen to be constant across that range, indicating that growth
was directly correlated with CP during May–October. We might therefore
reasonably expect faster growth in regions with higher forage CP levels.
Growth-rate curves for these regions coincide with composite CP, with the
exception of the Delta (Strickland and Demarais 2000), which may be affected
by enhanced diet quality from cultivated crops.
602 Southeastern Naturalist Vol. 7, No. 4
Forage protein digestibility is reduced by tannins (Robbins et al.
1987). Plant species may respond to changes in soil fertility by altering
growth patterns and levels of tannins. Species growing in lower fertility
soils may produce more tannins in their leaves than when grown in
more fertile soils, and these differences may be significant over relatively
subtle gradients (Kraus et al. 2004, Muller et al. 1987). Plants on more
fertile soils increase their total biomass (Biere 1995, Chapin 1980, Fraser
and Grime 1998, Kraus et al. 2004) and often divert fewer resources to
herbivore defense (Coley et al. 1985), though this response varies by species
(Almeida-Cortez et al. 1999). If this general relationship between
soil fertility and tannin production holds across the areas we sampled, it
is possible that differences in forage protein availability among these sites
would be increased.
Average precipitation is similar across all sampled regions, with weather
stations nearest each sampling area reporting long-term annual means of
142–159 cm and similar patterns of monthly accumulation (National Climate
Data Center 2008). Cumulative precipitation during January–April
2006 at the LCP site was 38 cm, 38% below the long-term mean. Sites in
other regions ranged from 9% below to 16% above normal for this period.
All sites suffered rainfall deficits for the period of May–July, ranging from
32–61%. Moderate moisture deficits are unlikely to have significant effects
on CP (Peterson and Scheaffer 1992, Seguin et al. 2002); because the
LCP sites received an average 37 cm rainfall in January–April, we believe
it unlikely that CP was reduced. Forage plants under moisture stress in this
region may actually increase their CP and total dietary nutrients as plant
growth slows (R. Lemus, Mississippi State University Extension Forage
Specialist, pers. comm.). Because rainfall deficits during late spring–early
summer were similar across all sites, we do not believe they were a biasing
The next logical step in understanding the relationship of soil regions
with deer morphometrics is to explore whether differences in forage quality
find their way into actual diet quality. We suggest direct measurement of diet
quality through ruminal sampling to determine seasonal CP levels in deer
diets among regions for comparison with morphometric data. Because diet
quality can be impacted by population density, nutritional plane should be
compared within regions among populations at different densities relative
to carrying capacity. We need to quantify the effects of habitat management
activities on plant quality, deer diet quality, deer morphometrics, and population
dynamics, especially fawn recruitment.
The authors are grateful to 3 anonymous reviewers for helpful comments on the
manuscript, and to P. Hanberry for sample collection. Support for this research was
provided through the McEntire-Stennis Fund and the Mississippi Department of
2008 P.D. Jones, S. Demarais, B.K. Strickland, and S.L. Edwards 603
Wildlife, Fisheries, and Parks through project W-48-57 of the Federal Aid in Wildlife
Restoration Program. This manuscript is contribution number WF257 of the Mississippi
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