United States

Department of

Agriculture

Forest Service

Southeastern Forest

Experiment Station

Research Paper

SE-287

Seasonal Mast Availability

for Wildlife in the Piedmont

Region of Georgia

John W. Edwards

David C. GUYM, Jr.

Susan C. Loeb

WINTER

SUMMER
Cover art by:

Suzy Mefferd, Clemson, SC.

November 1993

Southeastern Forest Experiment Station

P.O. Box 2680

Asheville, North Carolina 28802
Seasonal Mast Availability

for Wildlife in the Piedmont

Region of Georgia

John W. Edwards, Research Assistant

Department of Forest Resources

Clemson University

Clemson, South Carolina

David C. Guynn, Jr., Professor

Department of Forest Resources

Clemson University

Clemson, South Carolina

Susan C. Loeb, Research Ecologist

Southeastern Forest Experiment Station

Clemson, South Carolina

ABSTRACT (1990), however, argued that the combination of all other

mast-producing species is of equal or greater importance

than oak to both game and nongame wildlife. This

conclusion was reached because oak and hickory mast is

usually available only in fall and winter, and annual yields

vary widely.

We measured mast production by traditional and buffer species for

2 years on the Piedmont National Wildlife Refuge in Georgia. Our

objectives were to determine how these two types varied on a seasonal,

annual, and habitat basis. Mast from buffer species was more frequent

and diverse than that from traditional mast producers. Our findings

suggest that although traditional mast species are important, buffer

species may play an equally important seasonal role in maintaining

annual diversity.

Keywords: Buffer species, oak, Southeast, traditional.

Introduction

Availability of mast in specific seasons is important to

many wildlife species. Janzen (1971) defined mast

fruiting as the periodic synchronous production of large

seed crops by trees. Huntley (1990) described mast

species as woody species that produce fruits that are not

wind disseminated. Hard mast is fruit with a dry or hard

exterior; soft mast is fruit with a soft or fleshy exterior.

Traditionally, however, discussion of mast has included

only heavy-seeded hard mast, such as acorns (Quercus

spp. L.)’ and hickory nuts (Caryu spp. Nuttall), and

relatively large soft fruits, such as persimmon (Diospyros

virginiana L.), black cherry (Prunus serotina Ehrhart),

and dogwood (Cormsflorida L.). Seeds and fruits from

other woody species like pines (Pinus spp. L.), sweetgum

(Liquidambar styraciflua L.), poison ivy (Rhus

radicans L.), and Virginia creeper (Parthenocissus

quinquefolia L.) have often been ignored. The term

“buffer” has been used in connection with these species

(Huntley 1990) because their mast is believed to decrease

the adverse effects of poor crops of traditional mast

species like oaks and hickories.

Mast is important food for many wildlife species. Many

studies have shown the importance of oaks and hickories

to game animals, such as deer (Odocoileus spp.

Rafinesque),’ squirrels (Sciurus spp. L.), wild turkeys

(Meleugris gallopavo Vieillot),3 and bobwhite quail

(Colinus virginianus L.) (Goodrum and others 197 1,

Kirkpatrick 1990, Kurzejeski 1990, Martin and others

1961, Nixon and Hansen 1987, Nixon and others 1975,

Pelton 1990, Wentworth and others 1990). Huntley

’ Botanical nomenclature follows Radford and others (1968).

The mast-producing species in a habitat depend on the

characteristics of the habitat and past land-use patterns.

Mast production and availability are influenced by the

ages and sixes of plants, the season, inherent fruiting

ability, canopy coverage, age of stand, competition,

climate, and stem density (Stransky and Halls 1980).

Many studies show that acorn yields vary considerably

from tree to tree, species to species, and year to year

(Cypert and Webster 1948, Downs and McQuilkin 1944,

Goodrum and others 1971, USDA 1980). Nixon and

others (1980) and Sork (1983) reported large tree-to-tree

and annual variations in mast yields of hickories. Little

information has been published on the seasonal availability

of hard mast among habitat types.

Pew studies have been conducted on soft mast production.

Lay (1959) found flowering dogwood and blueberry

hawthorn (Crutuegus bruchyucuntha Engelm.) to be

consistent producers during a 4-year study in Texas. Of

the 50 mast species present in slash pine (Pinus elliottii

Engelm.) plantations 1 to 30 years old in southeastern

Georgia, 8 species produced abundant fruit, 11 minor

amounts, and 31 none (Johnson and Landers 1978).

Gallberry (Zlex glabru Aiton) and blackberry (Rubus spp.

L.) were the most abundant mast producers, representing

71 and 15 percent of the total mast yield, respectively. I n

contrast, blackberry (R. argutus Link) accounted for 91

percent of the total mast production in l- to g-year-old

loblolly pine (P. tuedu L.) stands in Mississippi (Camp0

and Hurst 1980). Little is known about seasonal soft mast

production and periodicity in mature forest types.

We measured the production of traditional and buffer mast

for 2 years on the Piedmont National Wildlife Refuge in

Georgia. Our objectives were to determine how these two

types varied on a seasonal, annual, and habitat basis.

* Mammalian nomenclature follows Honacki and others (1982).

’ Ornithological nomenclature follows Sptunt and Chamberlain (1970).
Study Description

The Piedmont National Wildlife Refuge (PNWR) is

approximately 40 km north of Macon in Jasper County,

Georgia. The rolling hills of the PNWR are typical of the

Piedmont Plateau. The PNWR was established in 1939 to

demonstrate that depleted land could be restored and

support viable populations of native wildlife. At that

time, the land was severely eroded and the soils were

depleted by over 100 years of cotton farming. Today,

much of the 14,000-ha PNWR is covered by pine and

mixed pine/hardwood forests. Loblolly pine seeded

naturally onto the area. Hardwoods occupied small

sheltered upland valleys and creek bottoms. The forested

stands of the PNWR are actively managed to maintain

populations of deer, wild turkey, red-cockaded

woodpeckers (Picoides borealis Vieillot), and other game

and nongame species.

Upland soil series include Gwimrett, Hiwassee, and

Wilkes with textures ranging from sandy loam to sandy

clay loam. Bottomland soils are Chewacla and Wehadkee

with textures from tine sandy loam.4

Twenty-two permanent transects, 50 m apart and 400 to

900 m long, were established. Reference points were

placed at 50-m intervals along each transect. Six mast

traps were randomly placed on each transect line. Each

mast trap was located 10 m from its assigned reference

point in a randomly selected cardinal direction

(N. ,E.,S. ,W.). Hard and soft mast were collected in

0.00004-ha circular basket traps (modified design of

Thompson and McGinnes 1963). The bottom of the trap

was mesh window screen attached to a wire (9 gauge)

hoop (228.6 cm circumference) to form a basket-shaped

structure. This “basket” was secured on three wooden

stakes, 1 m above the ground, and covered with 2.54-cmmesh poultry wire. Trap placement was intended to

reduce seed predation, but it also excluded mast from

low-growing species, such as blueberry (Vaccinium spp.

L.). Mast traps were checked monthly, and the presence

of fruits was recorded by species. No attempt was made

to quantify mast found in traps.

Stand boundaries were delineated using aerial photographs

and ground reconnaissance. Gn a lo- by 40-m plot in

each stand, d.b.h and species of codominant and dominant

stems were recorded. On a 5- by 20-m plot, d.b.h and

species of midstory stems > 2.5 cm in d.b.h and > 1.4

m in height were recorded. Habitat types were assigned

based on overstory basal area (USDA Forest Service

1988). Individual stands were assigned to one of four

habitat types:

1. Pine

2. Pine/hardwood

3. Hardwood/pine

4. Hardwood

Basal area

(Percent)

> 69 pine

51-69 pine

5 l-69 hardwood

> 69 hardwood

Stands were further classified, based on slope position.

Stands in top and upper-mid positions were classified as

“upland, * and stands in bottomland and lower-mid slopes

were classified as “bottom.” Because the numbers of

pine/hardwood and hardwood/pine stands were low, these

two classes were combined into a mixed pine/hardwood

habitat type. Accordingly, each of the 132 mast traps was

assigned to one of five habitat types: (1) upland pine,

(2) upland mixed pine/hardwood, (3) bottom pine,

(4) bottom mixed pine/hardwood, or (5) bottom

hardwood.

In these habitat types the dominant overstory species

were: loblolly pine in upland pine; loblolly pine,

sweetgum, and white oak (Q. alba L.) in upland mixed

pine/hardwood; loblolly pine and sweetgum in bottom

pine; and swamp chestnut oak (Q. michauxii Nuttall),

willow oak (Q. phellos L.), sweetgum, scarlet oak (Q.

coccineu Muenchh.), and shagbark hickory (C. ovutu

(Miller) K. Koch) in bottom mixed pine/hardwood.

Habitat characteristics, overstory stems/ha, midstory

stems/ha, and basal area are presented in table 1.

Mast items were categorized as traditional or buffer, hard

or soft, winged or nonwinged seeds, and vine fruit.

Comparisons of mast availability among seasons, years,

and habitat types were based on the relative frequency of

mast items. Relative frequency of mast item Mi =

’ Personal communication. 1992. Harold McMichael, soil conservation

technician, USDA, Soil Conservation Service, 206 S. Mulberry St.,

Jackson, GA 30233.

2
(frequency of M,/total frequency of all mast items M,

through M,,) x 100. Winter, spring, summer, and fall

seasons included the calendar months January-March,

April-June, July-September, and October-December,

respectively. In each habitat type, occurrences of mast

items from traditional and buffer species were compared

by season and year. An index of mast availability was

calculated by summing the number of traps containing

individual mast items, M, through M,, by mast type,

traditional or buffer, habitat type, season, and year. For

example, if a habitat type contained 10 mast traps and

mast item M, was found in 8 traps, mast item M, in 2

traps, and mast item M, in 1 trap, the mast availability

index would equal 8 + 2 + 1 = 11. Numbers of mast

items recorded in different habitats were adjusted for

unequal numbers of mast traps.

Results

Scientific names, fruiting seasons, and frequencies of

substantial yields for the 25 mast species sampled are

presented in table 2.

Availability of mast, based on relative frequency, varied

by season and habitat. Buffer species pine, yellow350 -

300-

poplar, and elm were among the 3 most common mast

items in 35, 27, and 10, respectively, of the 40

season/year/habitat type categories (tables 3-6).

Traditional species dogwood, oak, and muscadine were

among the 3 most frequently recorded mast items in 11,

6, and 2, respectively, of the 40 season/year/habitat type

categories (tables 3-6). Buffer species were the most

common mast item in 36 of 40 season/year/habitat type

categories, compared with traditional mast items which

were most frequent in only 4 categories; pine was the

most frequent mast in 26 of 40 categories, while oak was

most common in only 2 categories (tables 3-6).

Annual availabilities of elm and sweetgum differed

markedly. Elm mast, which was virtually absent during

1989, was one of the three most common mast items

available in all habitats during the winter of 1990 (table

3). Sweetgum mast, which was common in all habitats

during the fall of 1989, was not common in the fall of

1990 (table 6). In the habitats where their mast was

found, dogwood and oak were consistent producers during

1989 and 1990 (tables 3,5,6). Although seasonal

availability of pine and yellow-poplar (Liriodemfron

tulip&m L.) mast were similar, annual trends differed

(fig. 1). Pine mast availability was moderate in the

WINTER SPRING SUMMER FALL WINTER SPRING SUMMER FALL

1989 1990

Figure l-Seasonal availability of loblolly pine and yellow-poplar on the Piedmont National Wildlife Refuge, Georgia.
winter of 1989, poor in fall/winter of 1989/1990, and

high during the fall of 1990. Yellow-poplar showed the

reverse trend with high production during the winter of

1989 and fall/winter of 198911990, compared with low

availability in the fall of 1990 (table 7).

Across habitat types, species of available mast varied by

season. The most frequent winter mast items included

pine, yellow-poplar, elm, and dogwood (table 3); the most

common spring mast items were pine, yellow-poplar, elm,

maple, and sycamore (table 4); the most common summer

mast items were pine, dogwood, oak, muscadine, and

cherry (table 5); and the most common fall mast items

were pine, yellow-poplar, oak, dogwood, and sweetgum

(table 6). Traditional mast species occurred primarily

during summer and fall, while buffer species were

available throughout the year.

Mast availability varied among habitat types (tables 3-6).

Most frequent mast items in upland habitats included pine,

yellow-poplar, elm, dogwood, and sweetgum. In bottom

pine and bottom mixed pine/hardwood habitats, pine,

yellow-poplar, elm, dogwood, sweetgum, oak, and cherry

were the most common. Most frequent mast items in the

bottom hardwood type were yellow-poplar, oak,

muscadine, pine, and maple. Traditional mast species

occurred more frequently in bottom habitats than in

uplands. Traditional species accounted for 1 or more of

the 3 most common mast items in 46 percent (11 of 24)

of the season/year/habitat type categories in bottom

habitats compared with 38 percent (6 of 16) in the upland

habitats.

The number of individual mast species present, or mast

richness, was generally lower in spring and summer than

in fall and winter (table 8). Mast richness in spring and

summer was quite similar between years. In contrast,

mast richness in winter and fall varied considerably (table

8). Within seasons, mast richness was generally higher in

bottom types than in upland habitats, and bottom

hardwood habitats contained the highest mast richness

(table 8). Buffer species richness within season was

higher in 32 of 40 season/year/habitat categories than

traditional mast richness (table 8). Four of the five

categories where traditional mast richness was higher

occurred in upland mixed pine/hardwood and bottom pine

types during the summer (table 8).

Mast availability index was least in summer, intermediate

in spring, and greatest in fall and winter. Annual

variation (percent) in the mast availability index during

summer ranged from 0 in bottom pine to 106 in upland

pine, from 18 in upland mixed pine/hardwood to 91 in

bottom hardwood during spring, from 11 in upland pine

to 43 in bottom hardwood during fall, and from 7 in

upland mixed pine/hardwood to 52 in bottom hardwood

during winter (table 9). While the mast availability index

during fall and winter remained relatively constant in

upland habitats, bottom types showed higher

fluctuations-bottom hardwood had a 43-percent decrease

in fall and a 52-percent increase in winter. Bottom types

had higher mast availability indices within seasons than

did upland habitats; bottom hardwood had the highest,

except in the spring of 1989 (table 9). The buffer species

mast availability index was greater in 36 of 40

season/year/habitat type categories than the traditional

mast availability index (table 9). The traditional mast

availability index was higher in upland mixed

pine/hardwood, bottom pine, and bottom hardwood during

the summer.
Discussion

Mast richness and abundance are functions of site

conditions, weather, and past land use. Such

environmental constraints limit potential mast-producing

trees, shrubs, and vines. Once mast-producing species

are established, their inherent yield and periodicity will

determine mast abundance in a habitat. The fruiting

patterns of the species that are present also influence

seasonal mast abundance. We found mast richness to be

generally lower in spring and summer and higher during

fall and winter. This would be expected because only 3

of the 24 mast species present are known to fruit

predominantly in spring and only 5 during the summer; in

contrast, 10 are known to fruit in winter and 21 during

the fall. Mast richness was higher in bottom habitats than

upland types. Because loblolly pine is the single most

dominant pine species in all habitats, the presence of

hardwood species has a significant effect on mast

richness. Although the numbers of overstory hardwood

stems/ha decreased from bottom to upland habitats, a

strong midstory hardwood presence remained in all

habitats. However, the midstory hardwoods in the upland

types were small and their likely contribution to mast

richness was minimal.

Fire strongly influences species composition on the

PNWR. One purpose of prescribed fire is understory

hardwood control. Most stands are burned in winter at 3-

to 4-year intervals. Particularly in the uplands, burning

limits the presence of large vines, shrubs, and midstory

hardwoods. Midstory species maple, black cherry, and

dogwood do not begin to bear fruit until 4, 5, and 6 years

of age, respectively (USDA Forest Service 1974). Of the

mast items sampled on the PNWR, 15 of 24 were

produced by midstory hardwoods or vines. Although

many of these species do not produce large mast crops,

most are believed to be fairly consistent producers.

Either through constructed barriers, topography, or

moisture conditions, bottom habitats on the PNWR are

more protected from the effects of fire. As a result,

mast-producing hardwoods, shrubs, and vines are more

prevalent in bottom midstories.

In an area on the Piedmont of North Carolina where fire

had been excluded, the differences in species composition

between uplands and bottoms were less distinct than those

we observed at the PNWR (Oosting 1942). In the

absence of fire, there was greater overlap in species

composition among upland pine, bottom pine, and bottom

hardwood habitats. Johnson and Landers (1978) found

that mast yields from huckleberry (Gaylussacia spp.

HBK) and blueberry responded positively in frequently

(< 5 yr) burned areas, while other species were less fire

resistant and failed to attain mast-producing size. In

addition to mast species richness, improved site quality in

bottom types, including better soils, less drought, and

limited fire, may increase the yields of some species in

these habitats (USDA 1980).

Mast from buffer species was found to be more common

and diverse than that from traditional mast producers.

The species producing the most common buffer mast

items were pine, yellow-poplar, elm, ash, and maple; the

most common traditional mast items were from dogwood,

black cherry, oak, and blackgum. Traditional mast was

produced primarily in late summer and fall, while buffer

mast was more common throughout the year. During

summer-a season of low mast production-black cherry,

dogwood, and oak contributed significantly to the

availability of mast. Among oak, hickory, black walnut

(Juglans nigra L.), and beech (Fugus grandifolia

Ehrhart)-the traditional hard mast species occurring on

the study area-only oak mast was recorded in our mast

traps. Availability of oak mast on the PNWR was

relatively low in all habitats except the bottom hardwood

type. Our estimate may be less than was actually

available due to arboreal seed predators, which may

consume 11 to 13 percent of the acorns before they fall

(Cypert and Webster 1948, Kearby and others 1986).

Seed predation of varying degrees also occurs in other

mast-producing species, but its occurrence is not well

documented. In contrast to oak mast, traditional soft mast

was available in several seasons and habitat types.

The term “buffer” has been used to describe species

whose crops may compensate for low yields from

traditional mast species like oaks and hickories. Huntley

(1990) suggested that the term “buffer” is inappropriate.

Although he agreed that they serve a compensatory role,

he recognized that non-oak mast production is of primary

importance to many wildlife species even where acorns

5
are available. The importance of oak mast to wildlife has

been well documented (McGee 1990). Acorns are

consumed by 30 southeastern wildlife species (Martin and

others 1961). In comparison, buffer species such as pine,

yellow-poplar, and sweetgum are reportedly used by 22,

8, and 10 species, respectively. However, the

relationship between mast richness and wildlife use is not

exact. Each wildlife species may use several mast items

during a particular season, depending on availability and

individual preferences. Therefore, overlap is likely. For

example, of the 30 southeastern wildlife species known to

consume oak mast, 15 have been found to also consume

pine mast. Additional information quantifying mast

production and species specific use is needed.

Our findings suggest that although traditional mast species

such as oak, dogwood, and black cherry are important,

buffer species such as pine, yellow-poplar, elm, and

maple may play equally important seasonal roles in

maintaining the diversity of fauna found on the PNWR.

Although “buffer” may be an accurate term for certain

seasons or during periods of high mast yield by traditional

species, our results demonstrate the importance of buffer

species in ammal mast production. The diets of wildlife

species include numerous mast items. Both traditional

and buffer mast are common foods. Harlow and Van

Lear (1990) suggested that increases in mast diversity

should be accompanied by increased wildlife diversity.

Empirical data to support this hypothesis are unavailable,

however.

To maximize the benefit to wildlife and to minimize

variations in mast availability, we suggest promoting both

traditional and buffer mast species in conjunction with

maintaining species diversity. We would promote only

species endemic to habitats. This type of management

would allow buffer species to provide mast during seasons

when traditional mast species are not fruiting. During

traditional mast seasons, buffer species would compensate

when traditional mast crops fail. In addition, they would

add variety to wildlife diets.

Acknowledgments

Support for our study was provided by the USDA Forest

Service and the Department of Forest Resources, Clemson

University. R.L. Downing, R.F. Harlow, J.L. Landers,

and D. Van Lear provided helpful comments on earlier

drafts. We thank the staff of the Piedmont National

Wildlife Refuge for cooperation in this study.

6
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7
TABLES

Table 14tand characteristics of habitat types #ampled on the Piedmont National Wildlife Rofugo, Georgia

Over-story (stems/ha) Midstory (stems/ha) Basal area' (m'/ha)

Habitat

t-YE=

Pine Hardwood Pine Hardwood overetorv Midetorv

N Mean S D Mean SD Mean S D Mean S D Mean SD Mean SD

Upland

pine 2 2 115.9 56.5 10.2 16.7 331.8 891.4 763.6 829.5 12.1 4.9 7.8 8.2

Upland mixed

pine/hardwood 6 95.8 40.5 79.2 43.1 1550.0 2379.7 2166.7 1153.5 10.5 6.5 14.8 6.4

Bottom

pine

Bottom mixed

pine/hardwood

Bottom

hardwood

5 260.0 185.9 25.0 25.0 200.0 158.1 1260.0 602.5 22.0 4.5 9.0 3.4

3 58.3 14.4 58.3 14.4 33.3 57.7 1700.0 1652.3 16.7 8.4 19.2 2.6

7 7.1 18.9 175.0 79.1 0.0 0.0 1414.0 715.1 21.6 10.9 11.6 4.3

N = number of stande in sample; SD = standard deviation.

a Pine and hardwood combined.
Table P--scientific names, mast type, fruiting seasons, and periodicity of mast

species sampled on the Piedmont National Wildlife Refuge, Georgia (USDA Forest

Service 1974)

Fruiting

Common name Scientific name Mast types season Periodicitya

Ash

Black cherry

Blackgum

Dogwood

Elm

Hackberry

Hawthorn

Japanese

honeysuckle

Hornbeam

Ironwood

Maple

Muscadine

Oak

Pine

Poison ivy

Possumhaw

holly

Smilax

Supplejack

Sweetgum

Sycamore

Trumphet vine

Virginia

creeper

Winged sumac

Yellow-poplar

Fraxinus spp. L.

Prunus serotina Ehrhart

Nyssa sylvatica L.

Cornus florida L.

Ulmus spp. L.

Celtis occidentalis L.

Crataegus spp. L.

Buffer-WS Fall/winter 3-5 yr

Trad-soft Summer/fall Annual

Trad-soft Fall N/A

Trad-soft Fall/winter l-2 yr

Buffer-WS Winter/spring 2-3 yr

Trad-soft Fall

Buffer-NW Fall

Lonicera japonica Thunberg Buffer-vine Fall/winter

Ostrya virginiana (Miller) K. Koch Buffer-NW Fall

Carpinus caroliniana Walter Buffer-NW Fall

Acer spp. L. Buffer-WS Spring

Vitis rotundifolia Michaux Trad-soft Summer/fall

Quercus spp. L. Trad-hard Fall/winter

Pinus spp. L. Buffer-WS Fall

Rhus radicans L. Buffer-vine Fall/winter

Ilex decidua Walter

Smilax spp. L.

Berchemia scandens (Hill) K. Koch

Liquidambar styraciflua L.

Platanus occidentalis L.

Campsis radicans L.

Buffer-NW Fall/winter

Buffer-vine Fall/winter

Buffer-vine Summer/fall

Buffer-WS Fall

Buffer-WS Spring

Buffer-vine Summer/fall

Parthenocissus quinquefolia L.

Rhus copallina L.

Liriodendron tulipifera L.

Buffer-vine Summer/fall

Buffer-NW Fall/winter

Buffer-WS Fall/winter

Annual

N/A

N/A

N/A

3-5 yr

Annual b

N/A

Variable

3-13 yr

N/A

N/A

N/A

N/A

2-3 yr

l-2 yr

N/A

Annual

Annual

Annual

ws = winged seed;

N/A = no information available;

a Period between good mast crops.

b

For red maple (A. rubrum).

Trad = traditional;

NW = nonwinged seed.

9
Tablo 3dolativo frequency of uintcrr mast items by habitat typo ou the Phdmont National Wildlife Rofugo,

Qmorgia

Upland mixed Bottom mixed

Unland Dine Dine/hardwood Bottom Dine Dine/hardwood Bottom hardwood

Raet item 1989 1990 1989 1990 1989 1990 1989 1990 1989 1990

Blackgum

Dogwood

Elm

Hackbarry

Honeysuckle

Hornbeam

Ironwood

Maple

Oak

Pine

Poison ivy

Possumhaw

Smilax

Supplejack

Sweetgum

Sycamore

Trumphet vine

Winged sumac

Yellow-poplar

0.5

3.1

UP

0.5

N P

N P

N P

N P

N P

80.6

N P

N P

1.0

N P

0.5

N P

N P

N P

13.6

N P

2.2

17.7

N P

0.5

N P

N P

5.4

N P

47.8

1.6

1.6

3.2

N P

5.9

1.1

N P

N P

12.4

N P

N P

N P

N P

N P

N P

N P

N P

N P

80.0

3.3

N P

3.3

N P

3.3

N P

N P

N P

10.0

N P

3.1

25.0

N P

3.1

N P

UP

3.1

N P

37.5

UP

N P

3.1

N P

9.4

N P

N P

UP

15.6

UP

7.2

N P

1.4

N P

N P

N P

N P

N P

63.8

1.4

1.4

N P

N P

1.4

N P

N P

N P

23.2

N P

1.3

23.4

N P

3.9

N P

1.3

5.2

N P

23.4

1.3

1.3

3.9

N P

13.0

N P

N P

N P

22.1

UP

10.8

N P

N P

2.7

2.7

N P

N P

N P

54.1

2.7

N P

2.7

N P

5.4

N P

N P

N P

18.9

N P

5.0

27.5

N P

N P

5.0

N P

7.5

N P

17.5

5.0

5.0

2.5

N P

17.5

N P

N P

N P

1.5

N P UP

3.3 1.1

1.6 17.6

5.0 N P

3.3 2.2

N P N P

N P 4.4

3.3 15.4

N P 6.6

25.0 6.6

18.3 6.6

N P 2.2

N P N P

3.3 1.1

3.3 8.8

11.7 7.7

1.6 N P

1.6 3.3

18.3 16.5

UP = not present in sample.

Table O-Relative frequency of spring mast items found by habitat type on the Piedmont National Wildlife

Rofugo, Georgia

Upland mixed Bottom mixed

IJDland Dine Dine/hardwood Bottom Dine Dine/hardwood Bottom hardwood

Wast item 1989 1990 1989 1990 1989 1990 1989 1990 1989 1990

Ash

Blackgum

Black cherry

Dogwood

Elm

Naple

Pine

Possumhaw

Smilax

Sweetgum

Sycamore

Winged sumac

Yellow-poplar

N P N P

N P 1.5

N P N P

1.0 N P

1.0 26.2

N P 3.1

69.1 47.7

N P N P

N P N P

N P 1.5

5.2 N P

N P N P

23.7 20.0

N P

N P

N P

N P

N P

N P

63.6

UP

N P

N P

18.2

N P

18.2

N P

N P

N P

N P

23.1

15.4

30.8

N P

N P

N P

15.4

N P

15.4

N P

N P

3.8

N P

N P

N P

46.2

N P

N P

N P

1.1

N P

42.3

5.0

N P

10.0

N P

20.0

15.0

15.0

N P

N P

N P

5.0

N P

30.0

6.3

N P

N P

N P

N P

12.5

50.0

N P

6.3

N P

N P

N P

25.0

6.7

N P

N P

N P

33.3

33.3

6.7

N P

N P

6.7

N P

N P

13.3

N P

N P

NP

N P

N P

4.3

17.4

4.3

N P

N P

30.4

13.0

30.4

4.5

N P

N P

N P

25.0

20.5

4.5

N P

N P

NP

15.9

4.5

25.0

NP = not present in sample.

10
Tsblo Sdelstive frequency of summer maat items found by habitat type on the Piedeont National Wildlife

Rofug., Oeorgia

Upland mixed Bottom mixed

UDlsnd D ine Dine/hardwood Bottom Dine Dine/hardwood Bottom herdwood

PLenf item 1989 1990 1989 1990 1969 1990 1989 1990 1989 19w

Ash

Blackgum

Black cherry

Dogwood

Hornbeam

Huscadine

Oak

Pine

Poison ivy

Possumhaw

Smilax

Winged sumac

Yellow-poplar

N P

6.3

N P

6.3

N P

N P

N P

75.0

N P

N P

N P

N P

12.5

N P

15.2

3.0

6.1

N P

N P

N P

66.7

3.0

N P

N P

NP

6.1

N P

N P

25.0

25.0

N P

N P

N P

50.0

N P

N P

N P

N P

N P

N P

14.3

14.3

28.6

N P

N P

14.3

28.6

N P

N P

N P

H P

N P

N P

N P

18.2

18.2

N P

36.4

N P

27.3

N P

N P

N P

N P

N P

18.2

9.1

18.2

N P

N P

N P

18.2

36.4

N P

N P

N P

N P

N P

20.0

N P

N P

N P

N P

N P

N P

40.0

20.0

.NP

20.0

N P

N P

28.6

14.3

N P

28.6

14.3

N P

N P

14.3

N P

N P

N P

N P

N P

4.8 15.8

N P 10.5

N P N P

4.0 5.3

N P U P

23.8 N P

33.3 42.1

4.8 10.5

4.0 5.3

14.3 N P

N P N P

4.8 N P

4.8 10.5

NP = not present in sample.

Table 6delatire frequency of fall mast items found by habitat type on the Piedmont National Wildlife

Refuge, Qeorgia

Uld'd B tt d

UDlsnd Dine DiEe%ar%~d Bottom Dine DiEe/%r%gEd Bottom hardwood

Mast item 1989 1990 1989 1990 1989 1990 1989 1990 1989 1990

Ash 0.4

Blackgum 0.4

Dogwood 5.4

Hackberry N P

Hawthorn 0.4

Honeysuckle 0.4

Hornbeam 0.8

Ironwood N P

Maple N P

Huscadine N P

Oak N P

Pine 50.6

Poison ivy N P

Possumhaw 0.4

Smilax 2.5

Supplejack N P

Sweetgum 8.7

Sycamore N P

Trumphet vine N P

Virginia creeper N P

Winged sumac N P

Yellow-poplar 29.0

N P

1.8

2.8

N P

N P

N P

N P

N P

N P

N P

0.5

83.9

N P

N P

N P

N P

N P

N P

N P

N P

N P

11.1

N P N P

N P N P

9.1 la.4

N P N P

N P N P

2.3 N P

NP N P

N P N P

N P N P

N P N P

4.5 2.0

25.0 75.5

N P N P

N P N P

2.3 N P

N P NP

27.3 2.0

N P N P

2.3 N P

N P N P

N P N P

27.3 2.0

N P

1.0

10.6

N P

N P

1.9

N P

N P

2.9

1.0

N P

24.0

3.8

N P

3.8

N P

la.3

N P

N P

1.0

N P

31.7

1.4

1.4

9.6

N P

N P

1.4

N P

N P

N P

N P

2.7

63.0

1.4

N P

N P

N P

N P

N P

N P

N P

N P

19.2

8.3

N P

14.6

N P

4.2

2.1

a.3

N P

N P

N P

2.1

18.8

N P

N P

2.1

N P

18.8

N P

N P

N P

NP

20.8

13.9 1.7

N P N P

8.3 6.0

N P 0.a

N P N P

N P 1.7

N P N P

N P 4.3

N P 0.8

N P 2.6

2.8 19.8

63.9 8.6

2.8 6.0

N P 3.4

N P N P

N P N P

2.8 16.4

N P N P

N P 0.8

N P 2.6

N P 2.6

5.6 21.6

6.2

2.5

8.6

N P

N P

N P

N P

N P

N P

N P

27.2

35.8

2.5

N P

U P

1.2

N P

1.2

2.5

1.2

3.7

7.4

NP = not present in sample.

1 1
Table 'I-umber of mast traps containing pine and yellow-poplar seeds by year,

season, and habitat type on the Piedmont National Wildlife Refuge, Georgia

Year Upland Upland mixed Bottom Bottom mixed Bottom

and season pine pine/hardwood pine pine/hardwood hardwood

PINE

1989

Winter

Spring

Summer

Fall

1990

Winter

Spring

Summer

Fall

1989

Winter

Spring

Summer

Fall

1990

Winter

Spring

Summer

Fall

154 24 44 20 1 5

6 7 7 1 2 8 4

1 2 2 3 2 0

122 11 2 5 9 1 0

89 1 2 1 8 7 6

31 4 3 1 2

2 2 2 4 1 0

182 3 7 46 2 3 2 9

26 3 1 6

23 2 11

2 0 0

70 1 2 3 3

23 5 1 7 3 1 5

1 3 2 6 2 11

2 0 0 0 0

24 1 1 4 2 6

YELLOW-POPLAR

i

0

10

1 1

7

0

25

12
Tab10 g-Umber of different mast items in each habitat type by seaa and year OII the Piedmont National Wildlife

Rafugo, Neorgia

Year

and season

Upland mixed Bottom mixed

UDland Dine Dine/hardwood Bottom Dine

Traditional Buffer

Dine/h ardwood

Traditional Buffer Traditi OnSl

Bottom hardwoo$

Buffer Traditional Buffer Traditionam

1989

Winter 3 4 0 5 2 5 1 7 2 1 1

Spring 1 4 0 3 1 3 0 5 0 6

Summer 2 2 2 1 3 1 0 4 3 6

Fall 2 9 2 5 3 8 2 8 4 1 2

1990

Winter 1 1 0 1 I 1 1 0 1 9 2 1 2

Spring 1 5 0 5 1 6 0 6 0 7

Sumner 3 3 4 1 3 2 2 3 3 4

Fall 3 2 2 3 3 5 2 5 3 9

Table g-Index

Refuge, Qoorgia

pf seasonal mast availability by habitat type and year on tha Piedmont National Wildlife

Upland mixed Bottom mixed

Year UDland Dine Dine/hardwood Bottom Dine Dine/hardwood

and eeason Traditional Buffer Traditional B Uffer

Bottom hardwu

Traditional Buffer Traditional Buffer Traditional Buff1989

Winter 7.0 184.0 0.0 125.3 22.4 235.4 28.4 234.3 23.7 260.3

Spring 1.0 96.0 0.0 45.9 3.7 93.4 0.0 113.6 0.0 108.9

Sumner 2.0 14.0 8.4 8.4 29.9 11.2 0.0 35.5 61.5 37.9

Fall 14.0 226.0 25.1 158.7 40.6 340.1 56.8 284.0 160.9 388.1

1990

Winter 4.0 181.0 4.2 129.5 3.7 284.0 14.2 269.8 33.1 397.6

Spring 1.0 64.0 0.0 54.3 7.5 67.3 0.0 106.5 0.0 208.3

Sumner 8.0 25.0 20.9 8.4 18.7 22.4 21.3 28.4 52.1 33.1

Fall 11.0 206.0 41.8 162.9 37.4 235.4 28.4 221.2 146.7 236.1

aEetimate calculated by summing the number of mast traps containing individual mast items M, through Mn, by habitat

type. *eason, and year. For example, if a habitat type contains 10 mast traps, and mast item Ml is found in 8 of the

traps, mast item H2 in 2 traps, and mast item M3 in 1 trap, the estimated mast availability would be 8 + 2 + 1 I 11.

Numbers of mast items recorded in different habitats were adjusted for unequal numbers of mast traps.

1 3

Edwards, John W.; Guynn, David C., Jr.; Loeb, Susan C. 1993.

Seasonal mast availability for wildlife in the Piedmont Region of

Georgia. Res. Pap. SE-287. Asheville, NC: U.S. Department of

Agriculture, Forest Service, Southeastern Forest Experiment Station.

13 PP.

We measured mast production by traditional and buffer species for 2 years on

the Piedmont National Wildlife Refuge in Georgia. Our objectives were to

determine how these two types varied on a seasonal, ammal, and habitat

basis. Mast from buffer species was more frequent and diverse than that

from traditional mast producers. Our findings suggest that although

traditional mast species are important, buffer species may play an equally

important seasonal role in maintaining annual diversity.

Keywords: Buffer species, oak, Southeast, traditional.

Edwards, John W.; Guynn, David C., Jr.; Loeb, Susan C. 1993. i

Seasonal mast availability for wildlife in the Piedmont Region of

Georgia. Res. Pap. SE-287. Asheville, NC: U.S. Department of i

Agriculture, Forest Service, Southeastern Forest Experiment Station. I

13 PP. I

We measured mast production by traditional and buffer species for 2 years on ;

the Piedmont National Wildlife Refuge in Georgia. Our objectives were to

determine how these two types varied on a seasonal, annual, and habitat I

basis. Mast from buffer species was more frequent and diverse than that I

from traditional mast producers. Our findings suggest that although I

traditional mast species are important, buffer species may play an equally

important seasonal role in maintaining annual diversity.

/

Keywords: Buffer species, oak, Southeast, traditional. I

I
The Forest Service, U.S. Department of

Agriculture, is dedicated to the principle of

multiple use management of the Nation’s forest resources

for sustained yields of wood, water, forage, wildlife, and

recreation. Through forestry research, cooperation with the

States and private forest owners, and management of the

National Forests and National Grasslands, it strives-as

directed by Congress-to provide increasingly greater

service to a growing Nation.

USDA policy prohibits discrimination because of race,

color, national origin, age, religion, sex, or disability.

Person who believe they have been discriminated against in

any USDA-related activity should immediately contact the

Secretary of Agriculture, Washington, DC 20250.