United States

Department of

Agriculture

Forest Service

Southern Forest

Experiment Station

New Orleans,

Louisiana

Research Paper

so-259

Deer Habitat in the

Ozark Forests

of Arkansas

Mitchell J. Rogers, Lowell K. Halls, and James G. Dickson
SUMMARY

Two enclosures of 590 and 675 acres were constructed and stocked with

white-tailed deer (Odocoileus virginianus) to determine the deer carrying

capacity of an Ozark mountain forest and to evaluate the impact of winter food

plots on deer survival and productivity. Deer diets varied considerably within

and among years, and they were closely related to habitat type and availability

of acorns. Before food plots were established, the deer carrying capacity averaged only one deer per 45 to 100 acres, mainly because of a lack of high-quality,

native forage during winter. Capacity was somewhat higher in the enclosure

where cedar glades were more prevalent. Fawn production and winter survival

of adult deer fluctuated widely and were positively correlated with acorn yields.

After openings were established in the forest and planted with elbon rye (Secale

cereale), ladino clover (Trifolium repens), and Japanese honeysuckle (Lonicera

japonica), the carrying capacity increased to one deer per 21 acres, and population levels remained fairly stable from year to year. Food plots seemed beneficial

only during years of low mast yields. Even with access to high-quality forage,

the deer populations were limited by a low fawn survival rate due to predation,

disease, parasites, and other unknown factors.

November 1990
Deer Habitat in the Ozark Forests of Arkansas

Mitchell J. Rogers, Lowell K. Halls, and James G. Dickson

INTRODUCTION

The Ozark mountain range comprises approximately 5.2 million acres in northern Arkansas. The

wooded valleys and hills and the rivers and lakes have

long provided a haven for sportsmen and recreationists. White-tailed deer (Odocoikus virginianus) have

historically been a popular attraction, but their numbers have fluctuated widely due to the impacts of

farming, timber cutting, hunting, disease, parasites,

and poor nutrition.

In 1926, it was estimated that there were only 35

deer on the 173,000-acre Sylamore Ranger District of

the Ozark National Forest (Donaldson and others

1951). In 1927 and 1928, two Federal refuges were

established on the district. Intensive protection and

predator control helped to reestablish the deer herd. In

1944, 463 deer were harvested on the Sylamore

Ranger District (Segelquist and others 1969), and in

1945 the area was considered seriously overpopulated

with deer (Alexander 1954). The dense overhead canopy of the young developing forest limited the production of understory vegetation (Halls and Crawford

1960). The exclusion of fire from planted pines in old

fields further reduced range productivity. By 1950, the

declining forage supply and high deer populations

caused heavy overbrowsing and die-offs (Donaldson

and others 1951). Deer populations remained

depressed into the 1960’s; for example, only 49 deer

were harvested in 1967.

In an effort to resolve the situation, the Arkansas

Game and Fish Commission; the Fish and Wildlife

Service (Pittman-Robertson project W-53-R), U.S.

Department of the Interior; and the Southern Forest

Experiment Station, Forest Service, U.S. Department

of Agriculture initiated a long-term investigation in

north Arkansas. The study objectives were: (1) to

determine the deer carrying capacity of Ozark highland forests under the present management system,

(2) to explore the possibility of increasing deer numbers by improved management practices, and (3) to

determine factors influencing deer condition and

productivity.

The objective of the first 8 years of the study, 1959 to

1967, was to determine deer carrying capacity under

existing forest management practices. The second

phase, from 1968 to 1977, sought to determine if deer

numbers could be increased by establishing improved

food plots in the Caney enclosure.

METHODS

Area Description

The area was considered typical of Ozark highland

forests in topography, soils, and timber stand conditions. In general, the area was deeply dissected by

small streams, narrow valleys, and narrow, rounded

ridges. Elevations ranged from 400 to 1,020 ft. Slopes

ranged up to 60 percent but usually averaged from 30

to 40 percent.

The soil on ridges and upper slopes was extremely

rocky. Chert fragments made up 30 to 75 percent of the

total volume. Soils on lower slopes, benches, and floodplains were generally sandy to silty clay loams with a

few rock fragments. Water-holding capacity decreased

from streambottoms to the ridges.

On the average, annual precipitation was about 44

inches, with approximately 24 inches of rainfall during the warm months. Dry periods were frequent, and

the mean temperature was 37 “F in January and 81 “F

in July.

For this study, two 9-footrhigh fenced enclosures

were constructed at the Sylamore Experimental Forest, a part of the Ozark-St. Francis National Forest on

the Springfield Plateau in northern Arkansas. The

Caney enclosure contained 590 acres, and the nearby

Big Spring enclosure contained 675 acres. Fence construction was completed in 1962.

Each enclosure contained four main forest types:

upland hardwood, upland pine-hardwood, cedar glade,

and streambottom hardwood. In the Big Spring enclosure there was also a transitional pine-hardwood forest type. The principal hardwood trees were white oak

Mitchell J. Rogers was previously biologist at the Arkansas Game and Fish Commission, Little Rock, AR, and is presently at Ducks Unlimited,

Fifty Six, AR; Lowell K. Halls is wildlife biologist (retired); and James G. Dickson is research wildlife biologist at the U.S. Department of Agriculture, Forest Service, Southern Forest Experiment Station, Nacogdoches, TX.

1
(Quercus alba), black oak (Q. uelutina), hickories

(Curya spp.), and black cherry (Prunus serotina). Shortleaf pine (Pinus e&in&u) was dominant in the pinehardwood stands. There had been no timber cutting in

recent years, and many timber stands were largely

mixtures of various size and age classes, with only a

few acres of even-aged hardwood pole stands. Overstory canopies were heavy and continuous in all types

except the cedar glade, where small noncommercial

trees predominated.

Procedures

Deer Data

All the deer in both enclosures (36 in Caney and 30

in Big Spring) were removed by hunting and trapping

in 1962. Between April and November 1963, the

Caney enclosure was restocked with 8 does and 2

bucks, and the Big Spring enclosure was restocked

with 10 does and 1 buck.

Deer were counted each March and December by a

line-drive census using 70 to 130 drivers spaced 60 to

120 feet apart (Segelquist and others 1969). Additional census data were obtained from 1968 to 1977 in

the Caneyenclosure by direct observation from raised

platforms.

During the course of the study, deer were added or

removed from the enclosures to maintain desired population levels and to collect data on animal condition,

productivity, and internal parasites. Several adult

deer and young fawns were fitted with motion-sensitive, lithium-powered transmitters in an effort to identify specific mortality factors.

Necropsies were performed by personnel from the

Southeastern Cooperative Wildlife Disease Study, College of Veterinary Medicine, University of Georgia, on

7 of the original deer population in 1962 and on a total

of 10 additional deer in 1967, 1971, and 1978. A helminth parasite analysis was performed on 11 deer by

personnel from the University of Arkansas, Department of Zoology, in 1975.

Vegetation Measurements

Each August, current-year annual growth to a

height of 5 ft was estimated on about 125 permanent

and 30 temporary 6.2-ft2 plots systematically spaced

throughout each enclosure using double-sampling procedures (Wilm and others 1944). Forage production on

the temporary plots was estimated and then clipped

and weighed for use as a correction factor for forage

estimates on the permanent plots. Data are expressed

on an ovendry basis.

Winter forage availability was estimated on plots

each March from 1965 through 1972 with the same

technique used to estimate summer forage. Winter

browse included the previous season’s twig growth of

deciduous species and twigs and leaves of evergreens.

Forage utilization was estimated each August and

March in conjunction with forage yield measurements.

Summer utilization data were estimates of the weight

of forage removed, and the winter utilization data were

estimates of the percentage of growing stems browsed.

Rumen contents of deer killed for population control

and necropsy were also examined to determine food

habits. Fecal pellets were collected regularly from January 1966 through January 1968 and examined for

mast. Transects used to sample pellet groups were

randomly located in each of the main forest types, and

all pellet groups were counted each season from the

fall of 1966 through the summer of 1969 to determine

seasonal habitat preference. Because utilization is governed to some extent by relative abundance of plants,

an importance value was calculated by multiplying

the frequency of occurrence of plants along specified

transects by their frequency of utilization.

Mast yields were sampled each fall by placing two

55-gal, open-top barrels at 99 randomly located points

throughout each enclosure. Acorns and other fruits

that fell into the barrels were counted and converted

to an ovendry yield.

In February 1968 four food plots, ranging in size

from 1.7 to 5.7 acres, were established in the Caney

enclosure by bulldozing and disking. Approximately

half of each plot was planted with Japanese honeysuckle (Lo&era juponicu), and the other half was seeded

annually with a mixture of elbon rye (Se&e cereule)

and ladino clover (Trifolium repens). Food plots in 1968

consisted of 3.7 acres of rye and clover and 6.0 acres of

honeysuckle. The acreage of rye and clover was

increased to 7.6 acres in 1969 and maintained at that

level for the duration of the study. Production and utilization of rye and clover were estimated in midwinter

and late winter 1968 through 1977 using 10 caged and

10 uncaged 3.1-ft2 quadrats. Honeysuckle data were

collected in August and March 1969 through 1977 by

clipping a portion of randomly located plants and by

comparing weights of caged and uncaged plants.

Honeysuckle plants were mowed to about a 30-inch

height to control woody invaders. All plots were limed

at 1.5 tons/acre in April 1968. The rye-clover plots

were fertilized annually with 10-20-20 (N-P-K) fertilizer, and the honeysuckle plots were fertilized with

approximately 100 lb/acre of 30-0-O (N-P-K) fertilizer.

Some commonly used native species and forages

grown on food plots were collected for nutritive analyses in the summer of 1969 and the spring of 1971 and

analyzed for crude proteins, calcium, and phosphorus

(Association of Official Agricultural Chemists 1960).

2
Cell wall contents, acid detergent fiber, and acid detergent liquor were determined (Van Soest 1967), and in

vitro digestibility was determined using bovine rumen

liquor and pepsin (Tilley and Terry 1963). Native forage samples consisted of leaves and stems of cool season grasses, sedges, and forbs, as well as the terminal

end of browse twigs with attached leaves.

Woody measurement plots were established 6 feet

from the center of each forage plot. The basal area of

each tree species was recorded by l-inch diameter

classes. Cover of understory woody vegetation up to a

height of 5 feet was recorded from eight clusters of

pointrloop transects. ’

RESULTS

Caney Enclosure

Woody Plants

Hardwood saplings predominated in the upland

hardwood and upland pine-hardwood forest types

(table 1). Pole stocking (trees in the 5.0- to 9.0-inch

d.b.h. class) was densest in upland hardwoods. Sawtimber stocking (trees >9.0-inch d.b.h.1 was greatest

‘Parker, Kenneth W. 1953. A method for measuring trend in

range condition on national forest ranges. Forest Service, USDA,

Administrative Studies (Condition & Trend). 26 p. Plus form.

Unpublished mimeo.

in the streambottom hardwood forest type. Pines were

represented mainly in the upland pine-hardwood type

and to some extent in the cedar glade type.

The understory cover estimate for the enclosure

from plot samples was less than 21 percent. Flowering

dogwood (Cornus florid4 made up 16 percent of the

total browse cover; common deerberry (Vaccinium

stumineum) and lowbush blueberry (V. uaduns), 13

percent; several species of oaks, 14 percent; hickory, 6

percent; lindera (Linderu benzoin), 8 percent; tree huckleberry (V. arboreum), 5 percent; and blackgum

(Nyssu syluatica), 5 percent. Twenty-two other species

were recorded, but none comprised more than 4 percent of the total cover.

Forage and Mast Yields

Growing season yield of native vegetation averaged

115 lb/acre, ranging from 78 to 139 lb/acre (table 2).

Preferred browse comprised 48 percent of total yields;

nonpreferred browse, 38 percent; forbs and ferns, 18

percent; and grass and grasslike plants, 11 percent.

The yields were very consistent among years, except in

1974 and 1976, when yields increased as a result of

tree canopy openings caused by a tornado in 1974.

Cedar glade was the most productive habitat (192

lb/acre) followed by streambottom hardwood (138

lb/acre), upland pine-hardwood (96 lb/acre), and

upland hardwood (73 lb/acre). Yields of preferred

browse were highest in the upland pine-hardwood, and

the herbaceous plants were most productive in the relatively open cedar glade.

Table L-Tree basal area in the Canny deer enclosure, located on the Syhmore Ranger District of t?w

Ozark Natimnl Forest

Forest types

Upland Upland Cedar Streambottom All types

hardwood pine-hardwood glade hardwood combined

______________________________ Feet2/acre______________________________

Hardwood

Sapling* 52 43 29 28 45

Polet 36 13 21 19 26

SawtimberS 18 15 16 44 19

Total hardwood 106 71 66 91 90

Pine

Sapling 0 5 0 0 2

Pole 0 5 0 0 2

Sawtimber 1 17 10 0 7

Total pine 1 27 10 0 11

Total timber 107 98 76 91 101

* Trees 1.0 to 4.9 inches in d.b.h.

’ Trees 5.0 to 9.0 inches in d.b.h.

* Trees >9.0 inches in d.b.h.

3
Table a.--Summer yields and winter availability of native forage in the Canq enclosure, located on the Sylamore Ranger District of the Ozark National Forest

Vegetation

class

Summer Winter

Range of Range of

Average values Average values

------------____ poundslacre________________

Grass and

grasslike plants

Forbs

Ferns

Preferred browse

Nonpreferred browse

Total

10 5-20 1 o-1

12 4-20 0 0

6 3-10 2 o-4

41 29-73 8 4-13

38 26-41 5 3-6

109 78-139 16 8-18

Available winter forage averaged only 14 lb/acre

with 290 percent consisting of twigs from deciduous

woody plants. Green forage was practically nonexistent.

Mast yields fluctuated widely by year, ranging from

a low of 3 lb/acre in 1968 to 482 lb/acre in 1960 (table

3). On the average, white oak produced more acorns

than black oak, but black oak was more consistent,

producing some acorns each year. There were no regular cycles of high or low acorn production. White oak

acorn yields exceeded 134 lb/acre in 5 out of 19 years,

but only once in 2 consecutive years. Yields were <5

lb/acre for 7 years, and white oak produced no acorns

in 3 years.

Yields of white and black oaks appeared to be operating independently of each other. In some years the

yields were high for both species, but frequently the

yields of both species were low. At other times the yield

was high for one and low for the other.

Acorn yields were highest in the upland hardwoods

(184 lb/acre) where stocking of ma&bearing oaks was

greatest. The upland pine-hardwood type was second

with 96 lb/acre. This habitat occurred on the southand we&facing slopes where pines comprised much of

the stand. Production was lowest in the cedar glades

(17 lb/acre) due to low hardwood densities. Acorn

yields in streambottom hardwoods were quite variable, varying from zero to 55 lb/acre. No acorns were

produced in 13 out of 19 years in this forest type.

Mast yields from species other than oaks averaged

10 lb/acre, but varied from zero in 1968 to 35 lb/acre in

1960 (table 3). Those with the highest yields were dogwood, grape (Vitis spp.), and blackgum. Lesser production was noted for sassafras (Sassafras albidum), black

cherry, smooth sumac (Rhus gkzbra), shining sumac

(R. copallina), and chinkapin (Castanea ozarkensis).

Average production of non-oak mast was greatest in

streambottom hardwoods, at 34 lb/acre, with grape

being the main producer in this forest type. Upland

hardwoods produced an average of 14 lb/acre, mainly

from dogwood and grape. Upland pine-hardwoods produced 5 lb/acre and cedar glade produced < 1 lb/acre.

4

Deer Food and Feeding Habits

Dogwood and lowbush blueberry were the most

important browse species eaten by deer in late winter

(table 4). Species of lesser importance as browse were

eastern redcedar (Juniperus virginiana), New Jersey

tea (Ceonothus americanus), hophornbeam (Ostrya

virginiana), common deerberry, and red maple (Acer

rubrum). There was no deer utilization of black and

white oak, blackgum, farkleberry (Vaccinium

arboreum), greenbriar (Smilax spp.), sassafras, or

winged elm (Ulmus ahtu).

Grasses and sedges constituted 39 percent of the

stomach contents of five deer killed in March 1967

(table 5). Only a moderate amount of acorns was

recorded because they were fairly sparse at this time

of year, and those eaten were either partially decomposed or damaged by insect larvae. Consumption of

normally unpalatable items such as dead oak leaves,

dry twigs, and eastern redcedar suggested a scarcity

of palatable food for the deer. In general, it appeared

that items eaten by deer during the winter were more

closely related to availability than to preference.

Forbs and red maple had the highest importance

rating of plants eaten by deer during the summer

(table 6), followed by blackgum, sassafras, dogwood,

lowbush blueberry, poison-ivy (Z’oxicodendron

radicans), and hackberry (Celtis occident&s). White

oak, hickory, greenbriar, grape, and white ash (Fraxinus americana) were not utilized by deer. The relatively low importance values indicated that most

browse plants were not being utilized by deer and that

ample food was available during the summer.

Pellet count surveys were used to test for seasonal

habitat preferences. During the spring when herbage

and browse were most succulent, deer used all types of

habitat to some extent. Use was fairly evenly divided

among habitat types during the summer and was not

related to mast yields. In the fall when acorns were

available, deer concentrated in the upland hardwood

and streambottom hardwood forest types where mast

was most abundant. These forest types received heavi-
Table 3.-Mast production in the Caney enclosure, located on the Sylamore Ranger District of the Ozark National Forest

Year White

Oak Non-oak

Black Other’ Total Flowering Blackgum Grape Misc.+ Total

dogwood

____

1959

1960

1961

1962

1963

1964

1965

1966

1967

1968

1969

1970

1971

1972

1973

1974

1975

1976

1977

135 2 t

415 31 1

* 69 *

8 22 *

11 4 *

4 5 *

230 24 10

* 14 1

395 10 1 1

* 3 *

27 5 1

5 5 P

60 107 2

1 48 1

2 14 1 1

300 12 5

2 1 *

114 125 4

37 160 9

92 35 3

137

447

69

30

1 5

9

264

1 5

406

3

33

10

169

50

27

317

3

243

206

129

_ poun&/mm____________

1 3

11

18

4

3

3

2

*

24

2

3

1

1

6

*

2

2

5

25

1

6

*

1

P

1

*

3

1

t

P

1

1

t

4

%

2

1

1 2

1

2

3

---___

14

35

20

7

4

1

9

1

23

t

6

1

5

1

4

6

18

26

6

10

l Northern red oak (Quercus rubm), post oak (Q. stc&r.!o~, chinkapin oak, (Q. muehlenbergii).

’ Black cherry, smooth sumac, shining sumac, chinkapin, sassafras.

* Less than l/z lb/acre.

Table 4.-Frequency of occurrence and utilization and the importance

rating of browse forage in the Caney enclosure, located on

the Sylamore Ranger District of the Ozark National Forest,

March 1965

Species

Frequency Frequency

of of Importance

occurrence utilization rating

__________________percent__________________

Dogwood 37 33 12.21

Lowbush blueberry 41 23 9.43

Eastern redcedar 2 50 1.00

New Jersey tea 4 14 0.56

Hophornbeam 7 8 0.56

Common deerberry 9 6 0.54

Red maple 4 13 0.52

Blackgum 2 0 0

Black oak 14 0 0

Farkleberry 4 0 0

Greenbriar 2 0 0

Sassafras 6 0 0

Winged elm 2 0 0

White oak 27 0 0

Table 5.-Percent composition of stomach contents of five o&r from

the Caney enclosure, located on the Sylnmore Ranger District of the Ozark National Forest, March 1967

Stomach contents Percent

Grasses and sedges 39

Acorns 1 9

Unidentified 1 2

Unidentified browse 7

Low blueberry 6

Forbs 6

Dogwood 4

Dead oak leaves 4

Eastern redcedar 3

!lbtal 100
‘Ihble 6.-Occurrence and utilization frequencies and the importance

rating of forage in the Caney enclosure, located on the

Sylamore Ranger District of the Ozark Nahmal Forest,

August 1965

Species

Frequency Frequency

of of Importance

occurrence utilization rating

mast yields. Honeysuckle seemed especially important

to deer during periods of ice and snow storms when

many acorns were covered.

Nutritional Analysis

Forbs 63 9 5.67

Red maple 29 1 1 3.19

Blackgum 1 1 15 1.65

Sassafras 38 4 1.52

Dogwood 48 2 0.96

Lowbush blueberry 46 2 0.92

Poison-ivy 7 1 2 0.84

Hackberry 4 20 0.60

White oak 51 0 0

Hickory 35 0 0

Greenbriar 7 0 0

Grape 11 0 0

White ash 6 0 0

------____________Percent__________________

est use in the fall, even when mast yields were low. As

the acorn availability decreased during the winter,

deer concentrated in the cedar glade where evergreen

browse and green herbage was more prevalent. Winter

use in the glade was high regardless of mast yields but

higher yet in low mast years. During the winter of

1966-67, when mast yields were low, pellet groups

were only found in cedar glade.

Improved Forage

Yields of rye and clover from 1968-77 ranged from

1,062 lb/acre to 3,860 lb/acre (table 7). Utilization varied directly with the availability of mast. During winters of high mast production, utilization of rye and clover ranged from 11 to 18 percent. During winters of

low mast production, their utilization ranged from 39

to 98 percent of the total production. Deer utilized

mast and native forage in early winter and increased

their use of rye and clover later in the winter as native

foods decreased in availability. The relative contribution of rye and clover to the deer’s diet in a low mast

year was further illustrated when these two species

comprised 96 percent of the stomach contents from five

deer killed in March 1971.

Production of honeysuckle was light from 1969-72

due to the immaturity of plants (table 71, but yields

averaged 2,922 lb/acre during the last 5 years

(1973-77). In 1970-71, leaves of immature honeysuckle plants averaged 143 lb/acre. During the same

period, native green forages preferred by deer produced 2 lb/acre. Thus, 1 acre of honeysuckle produced

as much green winter forage as 70 acres of the undisturbed forest (Segelquist and others 1972).

Utilization of honeysuckle was limited mainly to the

winter period and was inversely correlated with fall

Crude Protein.-Crude protein content of honeysuckle leaves ranged from 11 to 16 percent (table 8) and

was consistently above the 6- to ‘I-percent levels suggested for mature deer maintenance (French and

others 1955). On the other hand, twigs of honeysuckle

were frequently below maintenance protein levels and

often had the lowest protein levels of all forages tested.

Crude protein content of rye and clover ranged from 9-

to 39-percent. Except for the spring of 1970 when samples were collected after rye had matured, these values

were above the 13- to Is-percent levels suggested as

optimum for deer growth (French and others 1955)

and were significantly higher than all other forages.

Except for dogwood twigs, the native species were

generally at or above the deer maintenance levels, but

native species seldom contained adequate protein for

optimum growth of young deer. Protein content of dogwood leaves ranged from 7 to 13 percent.

Protein content was highest when plants were

actively growing; thus, honeysuckle, dogwood, low

panic grasses (Panicurn spp.), and pussytoes (Antennuriu pluntuginifiliu) were highest in this nutrient in

the spring, and rye and clover were highest in the fall

and winter.

Calcium.-Dogwood was usually higher than any

other species in calcium content, ranging from 1.3 to

4.5 percent (table 91, considerably above the 0.45 percent calcium requirement needed to support growth

and skeletal development of weaned fawns (Ulrey and

others 1973). This high calcium content is characteristic of dogwood foliage (Fowells 1965). All forages contained adequate levels of calcium for maintenance,

and, except for honeysuckle twigs and panic grasses,

were sufficient for good antler development (Magruder

and others 1957). In general, the calcium levels were

highest during fall and winter months; this may possibly be related to rainfall patterns.

Phosphorus.-The phosphorus content of honeysuckle leaves and rye and clover foliage was consistently

higher than that for native forage (table 10). As a rule,

phosphorus content was below deer maintenance

levels (0.25 percent) for all forages except the rye and

clover, and this class of forage contained adequate

phosphorus for best antler development only during

the fall after antler growth is essentially complete.

Calcium-to-phosphorus ratios seldom fell within the

recommended range of 1:l to 2:l for livestock. The

ratio was generally highest for dogwood leaves and

twigs and eastern redcedar (1O:l to 4O:l) and lowest in

rye and clover and twigs (<7:1) for all seasons.

6
Table ‘I.-Yield and utilization of improved forage plants in the Caney enclosure,

located on tke Sylumore Ranger District of the Ozark National Forest

Year

Elbon rye and ladino clover Japanese honeysuckle

Yield Utilization Yield Utilization

Poundslacre Percent Pounds/acIV Percent

1968-69 1,980

1969-70 1,678

1970-71 1,303

1971-72 1,824

1972-73 1,062

1973-74 1,625

1974-75 2,226

1975-76 2,063

1976-77 3,860

98

56

57

17

12

11

18

39*

+

67 95

239 61

750 3

2,181 5

2,833 22

2,583 0

3,532 13

3,482 5

* Late collection after elbon rye had gone to seed.

’ No data.

Table 8.-Crude protein content of native and improved forages

1969 to 1970

Forage item Summer Fall Winter Spring Summer

1970 to 1971

Fall Winter Spring

___________________________ Percent of ovendry weight

- Japanese honeysuckle

Leaves

Twigs

Elbon rye and

ladino clover

Flowering dogwood

Leaves

Twigs

Eastern redcedar

Panic grasses

Pussytoes

11 13 14 16 13 15 12 16

5 5 5 7 5 6 5 8

16 39 20 9 24 31 25 18

10 13 10 8 9 12

8 5 4 6 6 6

8 8 7 8 9 6

10 11 9 8 10 15

7 8 8 8 9 10

Table 9.-Calcium content of native and improved forages

Forage item

1969 1970 to 1971

Summer Fall Summer Fall Winter Spring

___________~~~~~~~~~_ Pementofovendryweight ~~~~~~~~_____________

Japanese honeysuckle

Leaves

Twigs

Elbon rye and

ladino clover

Flowering dogwood

Leaves

Twigs

Eastern redcedar

Panic grasses

Pussytoes

1.6 1.5

0.5 0.5

*

1.1

0.4 0.6

1.5 1.4

0.7 0.6

1.3 1.1 1.1 0.9 0.7 0.9

3.2 2.9 2.5 3.2 4.5 1.3

2.1 1.4 2.1 3.2 2.6 2.0

1.5 2.3 1.6 1.7 2.9 2.0

0.5 0.7 0.5 0.8 0.6 0.6

1.6 1.1 1.1 1.5 2.1 1.2

* No data

7
Table IO.-Phosphorus content of native and improved fomges

1969 to 1970 1970 to 1971

Forage item Summer Fall Winter Spring Summer Fall Winter Spring

___________________________ Percentofovendryweight___________________________

Japanese honeysuckle

Leaves 0.21 0.28 0.23 0.28 0.23 0.21 0.18 0.16

Twigs 0.20 0.09 0.09 0.17 0.18 0.14 0.10 0.13

Elbon rye and

ladino clover 0.20 0.74 0.45 0.32 0.42 0.59 0.43 0.16

Flowering dogwood

Leaves 0.13 0.14 0.12 0.26 0.14 0.12 0.17 0.08

Twigs 0.11 0.10 0.14 0.14 0.11 0.10 0.11 0.10

Eastern redcedar 0.13 0.15 0.14 0.14 0.16 0.15 0.14 0.05

Panic grasses 0.12 0.11 0.16 L

0.13 0.08 0.14 0.10

Pussytoes 0.15 0.15 0.11 0.19 0.19 0.14 0.13 0.10

* No data.

‘Iable 1 L-Dry mntter digestibility of native and improved forges

1969 to 1970 1970 to 1971

Forage item Summer Fall Winter Spring Summer Fall Winter Spring

.____________________________________pe~ent_______________________________------

Japanese honeysuckle

Leaves

Twigs

Elbon rye and

ladino clover

Flowering dogwood

Leaves

Twigs

Eastern redcedar

Panic grasses

Puesytoes

* No data.

90 88 92 90 89 91 91 92

41 29 30 46 40 38 36 56

80 94 89 60 82 95 93 80

87 87 91 88 87 88 87 88

57 56 69 68 I 69 69 75

71 69 78 78 75 76 78 76

61 50 62 68 56 56 60 75

88 79 89 88 85 86 87 89

Forage Digestibility.-As determined by the in vitro,

nylon bag, dry matter digestion technique (Tilley and

Terry 1963), honeysuckle leaves were the most highly

digestible (88 to 92 percent) of all forages tested,

except during both fall seasons and one winter season

when rye and clover forages were highest (table 11).

Dogwood leaves were generally the most digestible of

the native forages, and honeysuckle twigs were the

least digestible of improved forage items. Twigs of

woody plants were consistently much less digestible

than the leaves. The digestibility values for dogwood

and honeysuckle leaves were generally slightly higher

than those reported in other studies (Short and others

1975).

Deer Responses

Population Changes 1963-67.-Twenty-nine adult

deer and seven fawns were removed from the enclosure

between March and November 1962 by live trapping

and hunting. The enclosure was restocked with eight

does and two bucks between April and June 1963

(table 12), but one buck died shortly after restocking.

The recruitment (annual production minus mortality)

of fawns in the spring of 1963 was seven.

The Caney enclosure was not censused for deer in

the fall of 1963. The census in March 1964 recorded 16

deer. A fall census was not made in 1964. Four deer

were shot during the winter of 1964-65 to keep deer

numbers at a prescribed level. Five deer were counted

in the census of March 1965. A census error was suspected so a recount was made in April. Six deer were

counted. A subsequent search revealed three deer carcasses, and it was assumed that a major die-off had

occurred during the previous winter when mast yields

were low. There were no surviving fawns in the enclosure during the summer of 1965. Nine deer were

restocked over the summer of 1965 to bring populations up to the desired level. The 1965 fall census of 16

indicated restocked deer had survived and that 1 fawn

had been added by recruitment.

All deer survived the winter 1965-66. There were

no recorded losses over summer. The fall census of 23

deer indicated recruitment of 7 fawns. In the spring of

1967, five deer were removed and necropsied, and one

8
Table 12.-Deer population changes in the Caney enclosure, located

on the Sylamore Ranger District of the Ozark National

Forest, 1963-67

Year Data source Number of deer

1963 Stocked

Recruitment

Unexplained losses

Removed

10

7

1

0

1964 Spring census 16

Recruitment Unknown

Unexplained losses 6

Removed 4

Stocked 0

Fall census No census

1965

1966

Spring census 6

Recruitment 1

Unexplained losses 0

Removed 0

Stocked 9

Fall census 16

Spring census 16

Recruitment 7

Unexplained losses 0

Removed 0

Stocked 0

Fall census 23

1967 Spring census 9

Recruitment 0

Unexplained losses 8

Removed 6

Stocked 0

Fall census 14’

* Includes deer removed for necropsy March l-3,1967.

deer was crippled and presumed dead. Immediately

following the necropsy work, a census drive recorded

only nine deer, and the finding of five carcasses indicated that another die-off had occurred during the winter of 1966-67, another winter of low mast availability. All nine deer survived the summer of 1967, but

there was no recruitment.

These study results indicated that the Caney enclosure would support one deer per 45to-100 acres on a

sustained basis. It was apparent that deer populations

would build up in excess of that density during periods

of average or above-average mast production, but deer

would die off during periods of mast scarcity.

Population Changes 1968-77.-Nine deer were

recorded in the Caney enclosure in 1968 when

improved food plots were established (table 13). Five

adult does were added during the summer. The fall

census of 1968 showed a population of 23 deer, 9 of

which were fawns.

During the winter of 1968-69, rye and clover were

available to deer on the improved food plots, but mast

production was the lowest recorded in the study. The

extra heavy utilization of the rye and clover (98 percent, table 7) indicated that deer were short of food.

Eight deer died during the winter, but 15 survived. In

comparison, only six had survived the 1964-65 winter

under similar food conditions but without food plots.

The fall 1969 census revealed a recruitment of 11

fawns. In contrast, no fawns were born or survived following the winter die-offs of 1964-65 and 1966-67

when no supplemental forage was available.

Twenty-two of the 26 deer present in the fall of 1969

survived over winter. Two deer jumped out of the

enclosure during the fall 1969 census drive, and two

deer were killed by bobcats. A moderate crop of mast

(39 lb/acre) and the availability of improved forage was

probably responsible for this high overwinter survival.

The fall 1970 census resulted in conflicting counts

of 20 to 24 deer. Six fawns were positively identified

during the fall, indicating that adult mortality

equaled or possibly exceeded recruitment during the

summer and fall. No carcasses were found. Escapes

from the pen, poaching, or other unknown factors

could have been responsible for the adult mortality.

Twelve live deer were counted in the spring of 1971.

However, it was assumed that 21 deer survived the

winter because 7 deer were removed for the disease

and parasite study, 1 deer was wounded and presumed

dead, and 1 deer was killed during the census when it

ran into a tree. These last two deer were added to the

removal data (table 13). Even though mast production

was very low in 1970-71 (11 lb/acre), the yields of

native and supplemental forage were adequate for

winter survival of all deer.

The fall 1971 count of 14 deer represented a recruitment of 4 deer. Two old bucks were found dead from

apparently natural causes. High mast yields enabled

all deer to survive the winter and resulted in limited

utilization of improved forages and undetectable use of

native forage.

The fall 1972 count of 18 deer indicated a recruitment of 4 deer over the spring count. One deer was lost

to unknown factors over winter even though mast production in the fall was relatively good (51 lb/acre).

Recruitment was good in the summer of 1973 with

eight deer added to the population. Twenty-four deer

survived the winter of 1973-74, and one was lost to

unknown factors. The survival rate was high even

though mast production was only 31 lb/acre, and utilization of cultivated forages was fairly light. High use

of honeysuckle was noted during a period of snow and

ice coverage.

A tornado damaged approximately 40 percent of the

study area in the spring of 1974. Twenty-eight deer

were counted in the fall, an increase of four over the

spring count. Since eight fawns had been observed in

late June 1974, it was obvious that some mature deer

died or escaped after the tornado damaged the fences.

Because of high mast production in the fall of 1974

(323 lb/acre), use of cultivated forages was light even

9
!hble 13.-Deer population changes in the Canny enclosure, located on the Sylumore Ranger Disrict of the Ozark National Forest following food

1969

1970

Year Data Source

1968 Spring census

Known reproduction

Known mortality

Removed

Stocked

Recruitment

Fall census

Spring census

Known reproduction

Known mortality

Removed

Recruitment

Fall census

Spring census

Knciwn reproduction

Known mortality

Removed

Recruitment

Fall census

Spring census

Known reproduction

Known mortality

Removed

Recruitment

_ Fall census

Spring census

Known reproduction

Known mortality

Removed

Recruitment

Fall census

* Includes deer removed for necropsy.

+ Conflicting counts.

1971

1972

plot establishment, 1968-77

Number of deer

9

9

0

0

5

9

23

1 5

11

8

0

11

26

22

6

2

0

22

20-24+

21’

4

2

9

4

14

14

4

0

0

4

18

though the population of deer was the highest attained

during the study. All deer survived the winter.

Seven newborn fawns in the enclosure and two outside in the wildlife management area were captured

by hand and equipped with radio transmitters in the

spring of 1975. All the instrumented fawns in the

enclosure were either killed by predators or died during the summer. One fawn outside the enclosure died

during the summer; the other survived for 5 months.

Two adult bucks were killed during live-trapping operations, and one doe was found dead from apparently

accidental causes. The net herd reduction during the

summer was one deer; thus, a minimum of two 1975

fawns survived to 6 months.

Five deer were lost during the winter of 1975-76.

Three had been killed by poachers, and a radioequipped buck was found dead with his front leg and

shoulder through the radio transmitter collar. The

other loss was unexplained. During this winter, utilization of improved forages was relatively high, 39 per10

Year Data Source Number of deer

1973 Spring census

Known reproduction

Known mortality

Removed

Recruitment

Fall census

1974 Spring census 24

Known reproduction 8

Known mortality 1

Removed 0

Recruitment 4

Fall census 28

1975 Spring census 28

Known reproduction 10

Known mortality 1 3

Removed 0

Recruitment 0

Fall census 27

1976 Spring census 22

Known reproduction 9

Known mortality 1 1

Removed 0

Recruitment 0

Fall census 21

1977 Spring census 18

Known reproduction 1 1

Known mortality 1 3

Removed 0

Recruitment 2

Fall census 20

17

8

0

8

25

cent for rye and clover and 13 percent for honeysuckle.

Improved forage crops eaten by deer totaled almost

9,000 ovendry pounds, the highest utilization during

the study. Food plots compensated for the low mast

yields (22 lb/acre), and no deer losses were attributed

to food shortages.

Twenty-one deer were counted in the fall of 1976,

one less than in the spring. One adult doe was found

dead from apparent gunshot wounds. Eight newborn

fawns were equipped with transmitters; seven of these

died or were killed by predators before reaching 6

weeks of age. The radio signal from one fawn was lost

after 20 days. When all deer were removed from the

enclosure at the end of the study, there were no deer

from the 1976 fawn class, which verified the high fawn

mortality of 1976. This was the second year in a row

that mortality exceeded reproduction.

Eighteen deer survived the winter of 1976-77.

Three adult deer carcasses accounted for all missing

deer. These losses were not attributed to lack of nutri-
tious food because mast production was 269 lb/acre,

and utilization of improved forage crops was extremely

low. Eight fawns that were captured and radio

equipped during the spring either died or were killed

by predators.

All 20 deer in the enclosure were removed by shooting from January through March of 1978. Age structure of these deer indicated a fawn survival of two

from the 1977 fawn class, none from the 1976 fawn

class, and two from the 1975 fawn class. Thus, only

four fawns survived to adulthood during the period

from 1975 to 1978. Reproductive potential as determined from necropsy analysis averaged 1.54 fetuses

per breeding-age doe with a range of 1.34 to 1.86.

Thus, fawn survival was only a small percentage of

the potential production.

These results indicated that the deer carrying

capacity of the enclosure with food plots was one deer

per 21 to 33 acres, compared to one deer per 45 to 100

acres without supplemental forage. The average fall

population without supplemental food plots was 13

deer (1 deer per 46 acres), whereas the average was 23

deer (1 deer per 26 acres) when supplemental food

plots were available. Deer removed for necropsy were

counted as surviving deer. Based on this evaluation,

the number of deer available to the hunter in the fall

was increased by 77 percent when supplemental food

plots were available to augment fluctuating mast supplies.

Mortality factors.-A concerted effort to isolate mortality factors began with the capture of fawns in 1975.

Twenty-five fawns were captured, 18 in the enclosure

and 7 in the adjacent wildlife management area.

Motion-sensitive transmitters were attached to fawns

captured in 1975, and a heatrsensing function was

added to transmitter units in 1976 and 1977. The combination of motion- and temperature-sensing functions

was effective in determining when mortality occurred.

The majority of captured fawns was found in light

understory in open hardwood and pine-hardwood timber stands. Most fawns were found bedded in the open

at the base of trees, occasionally in dead or blowndown tree tops, and usually in slight depressions in

the ground. A few were observed in heavy cover on

food plots.

Fawns were never located in the same place on 2

consecutive days. One set of twin fawns was captured

less than 1 hour after birth. Each fawn was found in a

separate location the night after being fitted with

transmitters. The fawns were not located together

again, although they were never more than 300 yards

apart for the 15 days they were monitored. The dam

was never more than 300 yards from one of the twins,

but she was seldom in direct contact with the fawns

except during nursing and grooming periods. She

would groom and nurse the twins and change their

locations daily, usually in early morning and late

afternoon. Typically, fawns were relatively inactive

during the first 2 to 3 weeks of life. Activity, size of

range, and the amount of time spent with the dam

increased with age.

None of the 18 radio-equipped fawns survived inside

the enclosure, and only 2 of 7 were known to have survived outside the enclosure. Predators killed 18, 2

starved, 2 died from accidents, 1 died from disease,

and 2 were unaccounted for.

Of the predator kills, 13 were due to coyotes and 5 to

bobcats. Predator-killed fawns averaged 14 days of age

with a range of 8 to 61 days. The two deaths from starvation occurred at the ages of 14 and 17 days and were

the result of abandonment by does. The abandoned

fawns were inactive in one location for 2 to 5 days and

then moved at random for 0.4 to 1 mile before dying. It

appeared that one predator-related death occurred to a

fawn near death from starvation.

An effort in 1977 to control predators by trapping

was of little consequence because only one coyote was

caught. Observation data indicated that the enclosure

may have served as a portion of the home range for

approximately three coyotes and two bobcats.

Necropsy of 14 fawns indicated 1 case of disease.

Acute interstitial pneumonia with hemorrhagic disease was suspected.

Causes of adult deer mortality were investigated in

1975-77 by attaching motion-sensitive radio transmitters to 16 adults. Three died or were killed by

poaching, accident, or predation, and one died when a

leg became entangled in the radio collar. Nine of the

transmitters failed due to antenna breakage, and one

failed because of internal malfunctions. Seven additional deaths were recorded for adult deer not fitted

with transmitters: three by poachers, two by deer

traps, and two by undetermined causes.

Herd Behavior.-Generally, adult bucks were less

active than other age or sex classes of deer in all seasons except fall. Adult males traveled, bedded, and fed

in groups of two to six individuals. Usually these

groups were made up of the same deer. These male

groups began to disperse in October, and individual

activity increased to a peak during the main breeding

period of October 29 to November 6. At this time, adult

bucks moved almost constantly throughout the enclosure. Adult males, generally active at night, were

highly active during daylight hours only during the

breeding period and regrouped after the breeding season. Yearling bucks were not as structured in their

activities and frequently fed and traveled with adult

doe family groups.

Adult does generally associated with their female

offspring. One group consisted of an old doe with her

young of the year, her 3-year-old daughter with her

young of the year, and a yearling female. Adult does

spent up to 75 percent of their time in selected highuse areas. There was some intermix between female

11
family groups and bucks in late winter and spring.

Adult females were somewhat more active during the

breeding period, but much less than bucks, and were

seldom found outside their core areas. Doe groups

broke up during late spring with the pregnant does

usually traveling and bedding alone until their newborn fawns were old enough to travel. Doe-fawn groups

began to re-form in midsummer and remained basically intact the remainder of the year.

Internal Parasites.--Five of the 19 deer removed at

the termination of the study (January-February

1978) were necropsied by personnel from the Southeastern Cooperative Wildlife Disease unit of the University of Georgia. Adult meningeal worms

(Parelaphostronglyus tenuis) and their larvae were

found in all five deer, but they were not associated

with significant tissue damage. Abomasal helminth

counts ranged from zero to 43 per deer for Ostertugiu

mossi and 60 to 571 for Skrjabinugiu spp. These low

levels indicated that food supplies were adequate for

the period preceding collection.

Abdominal worms (Seteriu yehi), gullet worms (Gongylonemu pulchrum), and intestinal worms (Capilluriu

bovis) were of little importance at the low levels

encountered. Blood and muscle protozoans Vheileriu

cerui and Sarcocystis sp.) were present, but neither

parasite was a significant mortality factory. Arthropod parasites were at a level commonly encountered on

deer in the Southeast.

Based on these results it was concluded in this study

that deer mortality was not parasite induced. The abomasal parasite count indicated a deer population

below the nutritional carrying capacity of the range

(Eve and Kellogg 1977). Similar conclusions were

drawn from necropsies performed on five deer collected from the surrounding wildlife management

area.

Genetics.-An indication of genetic problems from

inbreeding in the confined deer herd was suggested by

liver and kidney examinations with horizontal starch

gel electrophoresis (Price and others 1979). The average individual heterozygosity, ranging from 2.3 to 4.7

percent, was much lower for deer within the enclosure

than for white-tailed deer in other parts of the range.

Low genetic variability of the deer population may

have been associated with the low herd productivity

and high fawn mortality in the latter stage of the

study.

92 ft2/acre for all forest types except the cedar glade,

where it averaged 68 ft2/acre (table 14). Overstory canopies were heavy and continuous for all types except

the glade. Hardwood saplings were dominant in all

types except in the transitional pine-hardwood type

where pole size timber predominated. Streambottom

hardwoods had the highest basal area of sawtimber.

Pines were a major component of the timber overstory

only in the upland pine-hardwood and transitional

pine-hardwood stands.

Understory browse cover ranged from 14 percent in

upland hardwood to 31 percent in the pine-hardwood

stands. Species contributing most to cover were eastern redcedar, lowbush blueberry, grape, greenbriers,

white oak, and American elm (Ulmus americana).

Summer Forage Yields

Summer forage yields averaged 179 lb/acre and varied from 128 to 516 lb/acre (table 15). Yields tended to

be lowest in growing seasons with below average rainfall and generally increased with improved moisture

availability. The high forage yields in 1975 were a

direct response to the opening up of the canopy by the

1974 tornado. Opening of the stands reflects the

extent to which forage yields may be increased

through timber-cutting practices, which have been

recommended for oak-hickory forests (Torgerson and

Porath 1984).

Grasses, sedges (Carez spp.), forbs, and ferns

accounted for 31 percent of the average yield, and

browse for 69 percent. Composites, legumes, and mints

made up the bulk of the forbs, while panic grasses and

bluestems were the most common grasses. About half

the browse was classified as preferred by deer. Dogwood, lowbush blueberry, and common deerberry were

the most abundant species of preferred browse. Oaks

and hickories were the dominant of nonpreferred

species.

Winter Forage Availability

Winter forage was scarce, ranging from 14 to 103

lb/acre (table 16). Deciduous browse twigs made up

about 90 percent of the total winter vegetation. Preferred green forage, consisting primarily of panic

grasses, sedges, pussytoes, and eastern redcedar averaged only about 3 lb/acre. The relatively high yields in

1976 were a result of the 1974 tornado.

Big Spring Enclosure

Mast Yields

Woody Plants

Timber stands were largely a mixture of many size

and age classes, with only a few acres of even-aged

hardwood stands. Tree basal area ranged from 83 to

Mast yields averaged 73 lb/acre but ranged from 3 to

206 lb (table 16). Acorns made up 86 percent of all

mast. White oak acorns were most abundant, followed

by black, northern red, blackjack (Q. marilandica), and

12
Table Il.-Tree basal area in the Big Spring dser enclosure, located on the Sylumore Ranger District of the Ozark

National Forest

Forest types

Upland Upland Cedar Streambottom Transitional All types

hardwood pine-hardwood glade hardwood pine-hardwood combined

Hardwood

Sapling

Pole

Sawtimber

Total

hardwood

Pine

Sapling

Pole

Sawtimber

Total pine

Total timber

34 30 30 32 16 31

33 12 1 3 25 30 24

23 12 18 30 1 8 21

90 54 61 87 64 76

0 8 1 1 2 2

0 2 2 1 6 1

1 19 4 3 12 7

1 29 7 5 20 10

91 83 68 92 84 86

Table 15.-Summer fomge yields in the Big Spring enclosure, located on the Syhwre Ranger District

of he Ozark National Forest

Date

Grasses and

grasslike

plants

Forbs and Preferred

ferns browse

Nonpreferred

browse

Total

vegetation

1963

1964

1965

1966

1967

1968

1969

1970

1971

1972

1973

1974

1975

Average

* No data.

20 13 50 67 150

14 14 44 56 128

28 33 50 59 170

23 24 53 45 145

22 1 5 40 47 128

22 1 5 45 50 132

23 19 51 47 140

15 18 49 54 136

. * . * *

19 21 54 61 155

26 29 66 62 183 * * * l *

65 138 142 171 516

25 31 58 65 179

chinkapin oaks. Fruit of the dogwood, grape,

blackgum, and sassafras made up 14 percent of the

mast.

Acorn yields were less than 20 lb/acre for 4 years

and greater than 100 lb/acre for 4 years. White oak

was the most erratic major mast producer. No white

oak acorns were produced for 4 years and less than 5

lb/acre another 4 years. Some black oak acorns were

produced every year, but less than 9 lb/acre were produced 6 years.

The upland hardwood forest type produced the most

mast (114 lb/acre), followed by the upland pinehardwood type (79 lb/acre), the streambottom hardwood type, (59 lb/acre), and the dry cedar glade (25

lb/acre).

Deer Food and Feeding Habits

Deer diets varied widely between seasons. Based on

forage utilization estimates and stomach contents,

deer ate forbs, grasses, sedges, mushrooms, and the

succulent portions of browse during spring and summer. During these seasons deer used all forest types as

indicated by pellet group counts.

Deer diets during fall and winter varied largely

with mast availability. When plentiful, mast made up

almost the entire diet, and deer concentrated in the

most heavily wooded forest types. Grapes were heavily

used as soon as they began to drop. When acorn crops

were low, all mast was soon expended, and deer turned

to green winter forages such as eastern redcedar,

13
l’hble 16.-Winter fomge availability and mast yields in Big Spring enclosure, located on the Sylummv

Ranger District of the Ozark National Forest

Winter forage availability Mast yields

Year

Grasses and

grasslike

plants

Forbs and

ferns Browse Total ACOIllS Other Total

1963

1964

1965

1966

1967

1968

1969

1970

1971

1972

1973

1974

1975

1976

1977

Average

l

2

1

2

1

1

2

*

3

2

l

*

1

*

*

2

1

0

1

1

3

l

4

4

l

*

.

2

. l

* *

13 14

17 21

14 16

16 17

18 19

19 21

20 25

l *

14 21

23 29 * *

103 103

. *

26 29

5 6 11

23 5 28

149 9 158

44 1 45

190 16 206

3 0 3

45 3 48

9 0 9

* l *

51 3 54

23 13 36

. * *

4 26 30

113 32 145

160 21 181

63 10 73

* No data.

panic grasses, pussytoes, and sedges. At these times,

deer concentrated on the glade, where these forages

were most abundant. However, the scarcity of green

forage forced deer to eat considerable amounts of deciduous browse twigs and dead deciduous browse leaves

during periods of extreme mast shortage. Chemical

analyses of the most commonly eaten winter forages

indicated that the most sought after forages had the

highest nutritional content (Segelquist and others

19731.

Deer Responses 1963 Through 1971

In 1963, the enclosure was stocked with 10 does and

1 buck (table 17). From 1963 through 1971, 32 deer

were known to have been born in the enclosure (19

females and 13 males). The total number of deer

stocked and born was 29 does and 14 bucks. Seventeen

does and 10 bucks were shot or trapped and removed,

and 2 does and 1 buck were found dead. The remaining

10 does and 3 bucks apparently died or escaped from

the enclosure because their remains were never found.

Productivity of the deer herd was extremely low

through 1971. Based on the number of does r2 years

old present each spring and a fawning rate of 1.57

fawns per doe as determined from fetal counts, at least

67 fawns should have been born. Records were

obtained on only 32 deer born in the enclosure. These

figures illustrate the disparity between expected and

observed herd productivity.

Deer conditions ranged from poor to good depending

upon the time of year and quantity and quality of

available food. Deer captured in traps during the

spring and summer were all in fair-to-good condition.

Some large bucks taken from the enclosure during

periods of mast scarcity in late winter were in poor

condition. Deer were never found starving, but they

were in poorer shape after mast shortages and were

presumably more susceptible to other environmental

stresses.

Deer were hosts to a number of internal and external parasites, but only those of the protostrongylid

group were present in significant numbers. Adult

meningeal worms were present in most deer, and their

larvae were present in the lungs of all deer examined.

Unidentified protostrongylid larvae were found in the

lungs of deer. Parasitologic and physiologic data indicated that protostrongylid larvae, interacting with

other factors, may have affected deer herd health at

some times.

Poaching, predation, and escapes are unaccountable

factors that may have affected deer numbers. One deer

was shot and killed by a poacher, but its ear tag was

ultimately recovered. A bobcat killed one yearling

buck.

Based on the information collected from 1963 to

1971, the 675-acre enclosure was capable of supporting 10 to 13 deer or 1 deer per 50 to 70 acres. It

appeared that the carrying capacity was determined

by the scarcity of winter foods, which at times could

have been compounded by parasitism.

14
Table 17.-Deer population changes in the Big Spring enclosure,

located on the Syhnore Ranger District of the Ozark

National Forest, 1963-71

Date Data Number of deer

1963

1964

1965

1966

1967

1968

1969

1970

1971

Stocked 11

Recruitment 5

Fall population 16

Spring population 16

Recruitment 7

Shot 5

Losses 2

Fall population 23

Spring population 16

Recruitment 2

Trapped and removed 2

Fall population 16

Spring population 16

Recruitment 5

Shot 5

Fall population 21

Spring population 16

Recruitment 5

Killed in drive 1

Losses 7

Fall population 21

Spring population

Recruitment

Losses

Fall population

13

1

4

14

Spring population 10

Recruitment 6

Losses 4

Fall population 16

Spring population

Recruitment

Losses

Shot

Fall population

1 2

0

0

7

1 1

Spring population

Recruitment

Shot

4

1

5

DISCUSSION

Study results indicate that the deer carrying capacity of Ozark upland forests is generally low and

dependent on several factors that fluctuate within and

among years. Food availability is the most obvious

limiting factor and can be manipulated to some extent

by management practices. As shown in this study, the

90 to 145 lb/acre of available forage during the growing season seems adequate to support a moderate population of deer. But 85 percent or more of this forage is

unavailable to deer during the critical winter period,

and most is deciduous, low-quality browse. Thus, on

the basis of native forage yields, the heavily wooded

Ozark forests are limited in their potential to support

an overwinter population that may accrue during the

summer.

Native forage availability is also governed by habitat types and logging activity. For example, the Big

Spring enclosure, with its 311 acres of cedar glade and

streambottom hardwoods, consistently produced more

native forage than the Caney enclosure, with only 42

acres of these high forage producing types. Green winter forage production in the Big Spring enclosure was

three times greater than in the Caney enclosure due

to cedar glade acreage (Segelquist and others 1969).

Also, tree basal area in the Big Spring enclosure was

only 86 ft2/acre compared to 101 ft2/acre in the Caney

enclosure.

Forage yields are unlikely to change much from

year to year as long as timber stand conditions remain

fairly stable. The extent to which timber alterations

may influence forage yields was shown by the threefold increase of forage yields in the Big Spring enclosure during the growing season after the tornado blew

down many of the larger trees. Presumably the deercarrying capacity would be increased with a greater

availability of forage, particularly during the winter,

but the extent is not predictable from the results of

this study.

The deer carrying capacity of Ozark forests such as

the ones in this study is about one deer per 45 to 100

acres when the deer subsist solely on native forage.

The more glade forest types, the greater the capacity.

The problem of forage sparsity is further accentuated by the low nutritional quality. Although the protein content of most native species tested was adequate for deer maintenance most of the year, it seldom

reached a level needed for the optimum growth of

young deer. The phosphorus level was inadequate in

all cases, even for body maintenance, and the calciumto-phosphorus ratio was much higher than recommended.

The availability of mast, which consists mainly of

acorns, adds a potentially large, although highly

erratic, dimension to the deer carrying capacity. Deer

populations in both enclosures fluctuated drastically

according to mast availability. Mast yields during

1964-65 were 11 and 28 lb/acre in the Caney and Big

Spring enclosures, respectively. During the 1964-65

winter, deer numbers in Caney declined from one per

33 acres to one per 100 acres and in Big Spring from

one deer per 38 acres to one per 42 acres. During the

winter of 1966-67 with mast yields of 16 pounds per

acre, deer in the Caney enclosure declined from one

deer per 28 acres to one per 40 acres. In the Big Spring

enclosure with mast yields at 45 lb/acre, the overwinter population remained at one deer per 31 acres.

The negative repercussion of meager acorn crops on

mature deer was further accentuated by low fawn sur15
viva1 the following spring.. For example, after the winters of 1964-65 and 1966-67, no fawns survived in

the Caney enclosure, and there was only a 13-percent

increase of deer numbers in the Big Spring enclosure.

Usually the deer populations fluctuated less in the Big

Spring than in the Caney enclosure-probably

because of greater winter forage availability and

because mast yields were more consistent in the Big

Spring enclosure.

The fact that the overwinter carrying capacity and

the reproductive rate of deer fluctuate highly in relation to mast availability poses difficult problems in

the harvest strategy of deer. Data from this study suggest that special attention be given to an assessment

of expected mast yields prior to the fall hunting season. If mast yields, particularly acorns, are low, extra

hunting effort should be exerted to reduce the overwintering deer population. Under low mast conditions,

winter survival would be low, and those surviving

would experience a poor reproductive success the following spring. On the other hand, if herd numbers

were reduced at or below carrying capacity in the fall,

the surviving adult does would be more likely to go

through the winter in reasonably good shape and produce healthy fawns the following spring and summer.

Other studies have also shown the importance of adult

doe nutrition on fawn survival (Verme 1977, Log&

1973).

When acorn yields are moderate to high, there is

less need to impose a heavy harvest strategy. But here

again, a deer herd maintained at or below the carrying capacity will likely experience high reproductive

rates (McCullough 1984).

As visualized in the planning stages of this study, a

practical possibility for increasing the deer carrying

capacity of Ozark forests and ameliorating or cushioning the effects of highly fluctuating mast yields would

be to establish food plots that consistently produce

high-quality food during winter, the most critical part

of the year. Efforts to do so in the Caney closure were

successful.

The fir&year forage yields on food plots established

in 1968 were low and not sufficient to fully compensate for the low yields of native forage and mast. As a

result of this general food scarcity, utilization of native

and improved forage was high. Eight adult deer died,

and deer numbers declined from one deer per 26 acres

to one deer per 40 acres. However, this was a significantly higher carryover than the one deer per 100

acres that survived the winter of 1964-65 when no

food plots were available.

Deer that survived the 1968-69 winter produced 11

fawns the following summer, and the fall population in

1969 was 1 deer per 23 acres. This was the only significant reproduction recorded following any winter dieoff during the study. After the acreage of food plots

was expanded in 1969 and became fully productive,

the overwinter populations became fairly stable at one

deer per 21 acres. This was almost double that of one

deer per 46 acres prior to food plot establishment.

Where food plots are needed to compensate for a

deficiency in winter feed, it should be recognized that

food plots will be of less importance when acorn yields

are high. During such times, deer eat little else other

than acorns until the acorn supply is exhausted. Thus,

an assessment of acorn production should be made

each fall before time and money are expended to plant

and fertilize annual crops that will be utilized lightly

by deer.

Even though adult deer were not suffering from a

lack of nutritious food after the food plots were established, mortality still occurred. The cause was not

always obvious. Old age, poaching, predation, accident,

and unknown natural causes were often involved.

When food plots were available, it appeared that low

fawn survival was the major factor limiting the population. Fawns were usually born at or very near the

expected rate (as determined from prenatal fetal

counts), but they seldom survived 6 months. Of the 18

fawns outfitted with radio transmitters in 1975 and

1978, 77 percent were killed by predators, 6 percent

died from starvation, 11 percent had equipment loss or

failure, and 6 percent died from disease. Although

most of this mortality was associated with predators,

it is believed that predators were mainly the removal

mechanism and that genetic or other unidentified factors were involved with this mortality.

The study results emphasize several important facts

pertinent to deer and habitat management in the

Ozarks. Sparsity of high-quality native foods during

the winter and highly variable mast yields combine

with disease, parasites, and predators to restrict overwinter deer populations to a low level. Scarcity of quality forage can be alleviated to a large extent by timber-cutting practices (Torgerson and Porath 1984) and

by establishing high-quality food plots. Even so, the

populations may be curtailed by low fawn survival

resulting from predation, disease, parasites, and other

possibly interacting factors, such as genetics.

LITERATURE CITED

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832 p.

Donaldson, David; Hunter, Carl; Holder, T.H. 1951.

Arkansas deer herd. Little Rock, AR Arkansas

Game and Fish Commission. 72 p.

16
Eve, J. Hammond; Kellogg, Forest E. 1977. Management implications of abomasal parasites in southeastern white-tailed deer. Journal of Wildlife Management. 41(2): 169-177.

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for the in vitro digestion of forage crops. Journal of

British Grassland Society. 18: 104-111.

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Schoepke, B.L.; McGee, W.T.; Keahey, K.K. 1973.

Calcium requirements of weaned white-tailed deer

fawns. Journal of Wildlife Management. 37(2):

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17
Persons of any race, color, national origin, sex, age, religion, or with any handicapping condition are welcome to use and enjoy all facilities, programs, and services of the USDA. Discrimination in any form is strictly against agency policy,

and should be reported to the Secretary of Agriculture, Washington, DC 20250.
Rogers, M.J.; Halls, L.K.; Dickson, J.G. 1990. Deer Habitat in the Ozark

forests of Arkansas. Res. Pap. SO-259, New Orleans, LA: U.S. Department of Agriculture, Forest Service, Southern Forest Experiment Station. 17 p.

Two enclosures of 590 and 675 acres were constructed and stocked

with white-tailed deer (0docoiZeu.s uirginianus) to determine the deer

carrying capacity of an Ozark mountain forest and to evaluate the

impact of winter food plots on deer survival and productivity.

Keywords: Carrying capacity, disease, forage, mast, parasites, predators, white-tailed deer.