United States 

Department of 

Agriculture 

Forest Service 

Northeastern Forest 

Experiment Station 

Research Paper NE-607 

Abundance and Distribution of 

Vegetation under Four Hardwood 

Stands in North-Central 

West Virginia 

G. W. Wendel 
Abstract 

Forest floor samples were collected from four hardwood forest stands in West Virginia to study species composition, abundance, and distribution of vegetation that 

originated from seeds, rootstocks, rhizomes, and so on. The abundance and distribution of plants on square-foot sections of forest floor that were lifted and 

moved to the greenhouse indicate that under the most ideal conditions upwards 

of 800,000 stems per acre may occur during the first year. Sweet birch was the 

most abundant tree species, blackberry the most abundant shrub, and species of 

violets were the most abundant herbaceous plants. Implications of this vegetation 

on the regeneration of the forest are discussed. 

The Author 

G. W. WENDEL, research forester, joined the Northeastern Forest Experiment Station's Timber and Watershed Laboratory, Parsons, West Virginia, in 1962. He 

received a B.S. degree in forestry from Iowa State University in 1951, and a master's degree in forestry from Yale University in 1957. 

Manuscript received for publk.ation 22 June 1987. 

Northeastern Forest Experiment Station 

370 Reed Rd., Broomall, PA 19008 

October 1987 
Introduction 

Control of species composition is a major regeneration 

problem in central Appalachian hardwood forests. After 

harvest cutting, dense stands of tree seedlings, shrubs, 

vines, and herbaceous plants become established. The species composition of the future forest is often determined at 

this time. The situation is further confounded by variation in 

the silvical requirements and relative desirability of tree 

species, changes in site quality over short distances, and 

the destruction of tree seeds by birds, mammals, insects, 

and diseases. Slower growing desirable species such as 

newly germinated oak seedlings are often overtopped by 

less desirable plant species that make it difficult to bring the 

oak through. This paper presents estimates of the quantity 

and distribution of species that were observed in 12-inch x 

12-inch x Cinch samples of forest floor from four hardwood 

stands on two areas with different site indexes. These data 

provide an insight to the competition problem and serve as 

a basis for designing studies to cope with it. 

The Study Area 

All of the forest floor sampling was done on 1/2-acre growth 

plots that were established earlier as part of the Fernow 

Experimental Forest growth study. The Fernow Experimental Forest is a 5,000-acre outdoor laboratory of the Northeastern Forest Experiment Station located in Tucker County 

near Parsons, West Virginia. Two forested areas encompassing three site index classes were sampled. Area I is about 39 

acres of undulating, relatively flat land at an average elevation of about 2,300 feet. Area II approximately 80 acres in 

size has moderately steep topography with slopes mostly 

between 20 and 40 percent. Elevation on Area II ranges 

between 1,800 and 2,500 feet. 

Average oak site index (Trimble and Weitzman 1956) for 

Area I, which varied little in topography and soil, was 70. 

Area II varied widely in site index and was divided into three 

oak site index strata: 64, 70, and 80. 

The soils on both areas are silt loams derived from acid 

sandstone and shale from the Mauch Chunk and Pocono 

geologic formation. Calvin channery silt loam, a loamyskeletal, mixed mesic Typic Dystrochrepts, is the predominant soil on both areas with scattered areas of Dekalb silt 

loam, a loamy-skeletal mixed mesic Typic Dystrochrepts 

near the ridge tops. Both soils are moderately well-drained 

and 2 to 4 feet deep to bedrock. 

Precipitation averages 58 inches annually and is well distributed throughout the year. The average length of the 

frost-free season is about 145 days. 

The existing forest is composed of second growth hardwood timber that came in after the cutting during the early 

1900's, old residual trees of species that were in low 

demand during the early cutting, and trees that became 

established as the American chestnut died in the 1930's. 

The three stands on Area II were stratified by oak site index. 

The site index 80 area is typical of Society of American 

Foresters (SAF) cover type 59-yellow-poplar-white oaknorthern red oak (Eyre 1980) (Table 1). This type includes a 

large component of yellow-poplar, white oak, and northern 

red oak. At lower elevations black locust, white ash, and 

other mesic site species may be present. At higher elevations, American beech, red and sugar maple, and other species may become more abundant. The site index 64 and 70 

areas are more typical of SAF cover type 55-northern red 

oak. On the more moist sites a rather large component of 

yellow-poplar, black cherry, sugar maple, beech, white ash, 

and white oak may be present. On drier sites, other oaks, 

hickories, black gum, and red maple may be more 

abundant. 

Methods 

On Area I, ten 1/2-acre square growth plots were sampled. 

Forest floor samples were 12-inches square and were lifted 

from 16 locations established on a 25- by 25-foot grid in 

each growth plot, 160 samples in all. On Area II, two 1/2- 

acre circular growth plots were sampled in each site index 

stratum. Twenty forest floor samples were lifted from five 

locations at 15-foot intervals on each cardinal direction from 

plot center in each plot, 40 samples from each site index 

strata. Because of greenhouse space limitations, Area II 

samples were collected in November 1982, and Area I samples were lifted in November 1983. 

Forest floor samples were carefully cut with a machete to a 

depth of 4 inches around the inside of a wooden template. 

The sample trays were constructed of wood, 12-inches x 

12-inches x Cinches, with a 1M-inch mesh hardware cloth 

bottom. The cut block of forest floor was carefully lifted 

with a spade and placed in a tray. Where stoniness prevented lifting the sample intact, the sample was carefully 

gathered with a trowel and placed in a tray. Usually a single 

layer of leaves was placed on the hardware cloth bottom in 

the tray to prevent soil sifting during transport of the 

samples. 

The samples were labeled as to location and plot and randomly placed on greenhouse benches. No heat was maintained in the greenhouse during the winter, but flats were 

watered occasionally when the soil looked dry. About midMarch the automatic overhead watering system was started. 

Water was applied through a misting system for 2 hours 

each morning, and the greenhouse was heated to keep the 

flats from freezing during the early spring. Seeds that ger- 
Table 1.-Average number and average basal area, ftYacre, of stems larger than 1.0 inch d.b.h. in the four stands sampled 

Area I Area II 

Site Index 70 Site Index 64 Site Index 70 Site Index 80 

Species Number Basal area Number Basal area Number Basal area Number Basal area 

Eastern hemlock 

Sweet birch 

White oak 

Chestnut oak 

Red oak 

Cucumbertree 

Fraser magnolia 

Yellow-poplar 

Black locust 

Sugar maple 

Red maple 

Sourwood 

Hickories 

American beech 

Downy serviceberry 

Black cherry 

American basswood 

White ash 

American chestnut 

Scarlet oak 

Sassafras 

Black gum 

Dogwood 

Striped maple 

Eastern hophornbeam 

Slippery elm 

Total 172 133 534 143 380 174 41 9 147 

minated and/or root stocks that sprouted were allowed to 

grow until midSeptember after which time species in each 

tray were identified, counted, and the number recorded by 

tray. 

Forty-four plant species were identified in the trays at the 

end of the study. A few sparsely occurring grasses and 

sedges were observed in the trays but were not identified. 

Tree Species 

Sweet birch was by far the most abundant and frequent of 

the nine tree species observed in all of the trays (Table 2). 

Numbers ranged from 126,000 per acre to more than 

450,000 per acre with a frequency-percentage of flats containing at least one sweet birch-ranging from 68 to 98 percent. On the three Area il sites, sweet birch was most 

abundant in the trays that were collected from the site index 

64 site. 

Yellow-poplar was the second most abundant species. Projected field abundance was estimated at more than 100,000 

seedlings per acre on the oak site index 80 area and about 

23,000 per acre on the site index 64 area. Yellow-poplar frequency ranged from 73 percent on site index 80 to 40 percent on the site index 64 area. 
Table 2.-Projected average number of tree seedlings (tho~sandlacre)~ and frequencyb of occurrence in forest floor 

samples collected at four locations in West Virginia 

Area I Area I I 

Site Index 70 Site Index 64 Site Index 70 Site Index 80 

Species Number Frequency Number Frequency Number Frequency Number Frequency 

Sweet birch 

Yellow-poplar 

Sourwood 

Sassafras 

Fire cherry 

Red oak 

Black locust 

Fraser magnolia 

Red maple 

Sugar maple 

Black cherry 

Total 

a Rounded to nearest 1,000. 

Percentage of flats with at least one seedling. 

Sassafras also occurred in trays from all sites but was most 

abundant and well distributed on the site index 64 area70,000 per acre with 73 percent frequency. Sassafras was 

also abundant on Area I site index 70, but its occurrence 

was considerably less than that in the trays from the Area II 

site index 64. Sassafras frequency was low in the trays from 

Area II, site index 70 and 80. 

Red oak seedlings ranged from about 1,000 to more than 

6,000 per acre on areas with site index 64 and 70. No red 

oaks were observed in the trays from the site index 80 area. 

Seedling frequency in trays from the site index 64 and 70 

areas ranged from 3 to 5 percent. A heavy population of red 

squirrels, chipmunks, and other rodents was observed on 

both areas, and most of the uneaten acorns were observed 

to be weevil infested. 

Sourwood abundance averaged between 36,000 and 74,000 

seedlings per acre in the trays from site index 64, Area II 

and site index 70, Area I. Frequency ranged from 31 to 68 

percent. 

Other tree species that emerged in the trays were fire 

cherry, black locust, Fraser magnolia, red maple, sugar 

maple, and black cherry. Most of these were poorly distributed occurring in only a small percentage of the trays. 

Semi-woody and shrub species 

Of the nine shrub species identified in the flats, the most 

abundant was blackberry. Blackberry stems ranged from 

about 100,000 per acre to more than 200,000 per acre (Table 

3). Frequency ranged from 73 to 88 percent of the trays. 

Blackberry was the most abundant in trays from the two site 

index 70 areas and least abundant in those from site index 

64. 

Wild grape was the second most abundant species in this 

class. The number of wild grape ranged from 5,000 to 

100,000 seedlings per acre. Trays from the site index 64 area 

contained fewer wild grape seedlings than those from the 

site index 70 or 80 sites. Frequency ranged from 8 percent 

for trays from the site index 64 area to 61 percent for the 

trays from the higher site index areas. 
Table 3.-Projected average number of shrub and semi-woody stems (tho~sandlacre)~ and frequencyb of occurrence in 

forest floor samples collected at four locations in West Virginia 

Area I Area II 

Site Index 70 Site Index 64 Site Index 70 Site Index 80 

Species Number Frequency Number Frequency Number Frequency Number Frequency 

Striped maple 

Devils-walkingstick 

Blackberry 

Wild grape 

Flame azalea 

Teaberry 

Blueberry 

'Flowering dogwood 

Total 283 - 245 - 258 - 185 - 

a Rounded to nearest 1,000. 

Percentage of flats with at least one seedling. 

Devils-walkingstick was abundant in the trays from the site 

index 64 and 70 (Area I) areas (Table 3). Frequency was 41 

and 23 percent, respectively, for the two sites. On Area II, 

site index 70 and 80, there were fewer stems per acre and 

only 10 percent of the trays contained devils-walkingstick. 

Many of the other shrubs observed-teaberry, blueberry, 

azalea and flowering dogwood-were restricted to the site 

index 64 area. The most abundant was teaberry. 

Herbaceous Species 

Twenty-seven species of forbs, grasses, and ferns were 

identified in the flats. Sparse amounts of other grasses and 

sedges were not identified. 

Species of violets were the most abundant and well distributed-in the trays from the better sites (Table 4). Field estimates ranged from 34,000 to about 169,000 stems per acre. 

Violet frequency ranged from 39 to 80 percent in the trays 

from the site index 70 and 80 areas, compared to 20 percent 

in the trays from the area with site index 64. 

Other common species that occurred on all sites were plantainleaf sedge, mountain spleenwort, pokeweed, and 

common cinquefoil. Knotweed occurred only in the trays 

common cinquefoil. Knotweed occurred only in the trays 

from Area II, site index 64. Among the species occurring in 

trays from the two better sites in Area II were enchantersnightshade, white snakeroot, two-leaved toothwort, Indian 

tobacco, wild stonecrop, cleavers, and wood nettle. There 

were more herbaceous species in Area I trays from site 

index 70 than there were from Area II samples, which also 

had an oak site index of 70. 

Discussion 

When forest stands in the Central Appalachians are disturbed by clearcutting, fire, or mechanical site preparation, 

rapid revegetation of the areas usually occurs (Kochenderfer and Wendel 1983; Wendel and Trimble 1968). Numerous observations on clearcut areas show that by the end of 

the second growing season site occupancy is usually 

attained. The potential for more than 800,000 stems per acre 

to become established in the first year as estimated from the 

observations in this study lends credence to the statement 

that "nature abhors a vacuum." In natural ecosystems that 

are scheduled to be managed, this can lead to problems. 

On the one hand, site stability is rapidly restored and on the 

other, an abundance of undesirable species often creates 

drastic and costly control procedures. 
Table 4.-Projected average number of herbaceous plants (thousandlacre) and frequencya of occurrence in forest floor 

samples collected at four locations in West Virginia 

Area l Area II 

Site Index 70 Site Index 64 Site Index 70 Site Index 80 

Species Number Frequency Number Frequency Number Frequency Number Frequency 

Plantainleaf sedge 

Violet 

Mountain spleenwort 

Pokeweed 

Common cinquefoil 

Upright yellow 

wood sorrel 

Deertongue grass 

Fireweed 

Woodnettle 

Richweed 

Shield fern 

Whorled loose strife 

Wild yam 

Enchanters-nightshade 

White snakeroot 

Two-leaved toothwort 

Virginia avens 

Indian tobacco 

Wild stonecrop 

Buttercup 

Dodder 

Cleavers 

Trillium 

Knotweed 

Wild cranesbill 

Total 

a Percentage of flats with at least one seedling. 
The important aspect is that ground cover is quickly established and that the loss in soil and nutrients is curtailed by 

plant roots and recycling of nutrients through the plants. 

Marks (1974) has emphasized the importance of early successional species such as fire cherry in accomplishing this. 

Similarly, the large number and wide distribution of sweet 

birch, yellow-poplar, blackberry, wild grape, and devilswalkingstick observed in the flats perform the same role in 

the natural environment. Sweet birch seedlings probably 

arose from seeds disseminated the previous winter, whereas 

much of the yellow-poplar, blackberry, wild grape, devilswalkingstick, arose from seed stored in the forest floor. 

Studies have shown that wild grape, yellow-poplar, black 

cherry, and sassafras seeds are known to retain their 

viability in the forest floor for several to many years (Clark 

and Boyce 1964; Wendel 1977,1981 ). 

Numerous grasses and forbs were identified in the flats. 

Some of these may have originated from seed stored for 

relatively long periods but this was not determined. Other 

species probably are biennials or perennials that are components of the spring or summer flora and originated from 

existing rootstocks. 

The impact of herbaceous plants on tree reproduction in 

central Appalachian forests is not well understood. However, it has been shown in other areas that plant antagonisms exist between and among many species. With such a 

large and varied flora encountered in this study, there are 

undoubtedly some allelopathic relationships. Many studies 

in different areas support this contention (Horsley 1977; 

Fisher et al. 1978; Gabriel 1975; Tubbs 1973,1976; Hook and 

Stubbs 1967). 

The immediate effect of early and rapid natural revegetation 

of disturbed forest sites is site stabilization. Over the longer 

term, however, regeneration of forest stands may be 

delayed or seriously curtailed. A knowledge of the potential 

species mix for various sites and parent stand compositions 

can provide the forest manager with insight to this problem 

and highlight specific areas where preharvest treatment and 

research might be needed. 

Literature Cited 

Clark, F. Bryan; Boyce, Stephen G. 1964. Yellow-poplar seed 

remains viable in the forest litter. Journal of Forestry. 

621564567. 

Eyre, F. H., ed. 1980. Forest cover types of the United States 

and Canada. Washington, DC: Society of American Foresters. 148 p. 

Fisher, R. F.; Woods, R. A.; Glavicic, M. R. 1978. Allelopathic 

effects of goldenrod and aster on young sugar maple. 

Canadian Journal of Forest Research. 8(1):1-9. 

Gabriel, William J. 1975. Allelopathic effects of black walnut 

on white birches. Journal of Forestry. 73(4):234-237. 

Hook, D. D.; Stubbs, J. 1967. An observation of understory 

growth retardation under three species of oaks. Res. 

Note SE-70. Asheville, NC: U.S. Department of Agriculture, Forest Service, Southeastern Forest Experiment 

Station. 2 p. 

Horsley, Stephen B. 1977. Allelopathic inhibition of black 

cherry by fern, grass, goldenrod, and aster. Canadian 

Journal of Forest Research. 7(2):205-216. 

Kochenderfer, J. N.; Wendel, G. W. 1983. Plant succession 

and hydrologic recovery on a deforested and herbicided 

watershed. Forest Science. 29(3):545-558. 

Marks, P. L. 1974. The role of pin cherry (Pnmus pensylvanica L.) in the maintenance of stability in northern hardwood ecosystems. Ecological Monographs. 4473-88. 

Trimble, G. R., Jr.; Weitzman, Sidney. 1956. Site index studies of upland oaks in the northern Appalachians. Forest 

Science. 2:152-173. 

Tubbs, Carl H. 1973. Allelopathic relationship between 

yellow birch and sugar maple seedlings. Forest Science. 

19(2):139-145. 

Tubbs, Carl H. 1976. Effect of sugar maple root exudate on 

seedlings of northern conifer species. Res. Note NC-213. 

St. Paul, MN: US. Department of Agriculture, Forest Service, North Central Forest Experiment Station. 2 p. 

Wendel, G. W. 1977. Longevity of black cherry, wild grape, 

and sassafras seed in the forest floor. Res. Pap. NE-375. 

Upper Darby, PA: U.S. Department of Agriculture, Forest 

Service, Northeastern Forest Experiment Station. 6 p. 

Wendel, G. W. 1981. Longevity of summer grape seed stored 

in the forest floor. Wildlife Society Bulletin. 9(2):157-159. 

Wendel, G. W.; Trimble, G. R., Jr. 1968. Early reproduction 

after seed-tree harvest cuttings in Appalachian hardwoods. Res. Pap. NE-99. Upper Darby, PA: US. Department of Agriculture, Forest Service, Northeastern Forest 

Experiment Station. 16 p. 
List of Species Mentioned in this Paper 

Tree Species Shrubs and Herbaceous Vegetation 

Common Name Scientific Name 

American basswood Tilia americana L. 

American beech Fagus grandifolia Ehrh. 

American chestnut Castanea dentata (Marsh.) Borkh. 

Black cherry Prunus serotina Ehrh. 

Black gum Nyssa sylvatica Marsh. 

Black locust Robinia pseudoacacia L. 

Chestnut oak Quercus prinus L. 

Cucumbertree Magnolia acuminata L. 

Downy serviceberry Amelanchier arborea (Michx.f.) Fern. 

Eastern hemlock Tsuga canadensis (L.) Carr. 

Eastern hophornbeam Ostrya virginiana (Mill.) K.Koch 

Fire cherry Prunus pensylvanica L. f. 

Flowering dogwood Cornus florida L. 

Fraser magnolia Magnolia fraseri Walt. 

Hickories Carya sp. Nutt. 

Red maple Acer rubrum L. 

Sugar maple Acer saccharum Marsh. 

Northern red oak Quercus rubra L. 

Sassafras Sassafras albidum (Nutt.) Nees 

Scarlet oak Quercus coccinea Muenchh. 

Slippery elm Ulmus rubra Muh I. 

Sourwood Oxydendrum arboreum (L.) DC. 

Striped maple Acer pensylvanicum L. 

Sweet birch Betula lenta L. 

White ash Fraxinus americana L. 

White oak Quercus alba L. 

Yellow-poplar Liriodendron tulipifera L. 

Common Name 

Blackberry 

Blueberr.4 

Buttercup 

Cleavers 

Common cinquefoil 

Common dodder 

Deertongue grass 

Devils-walkingstick 

Enchantersnightshade 

Fireweed 

Flame azalea 

Indian tobacco 

Knotweed 

Mountain spleenwort 

Plantainleaf sedge 

Pokeweed 

Richweed 

Shield fern 

Teaberry 

Trillium 

Two-leaved toothwort 

Upright yellow 

wood sorrel 

Violets 

Virginia avens 

White snakeroot 

Whorled loosestrife 

Wild cranesbill 

Wild grape 

Wild stonecrop 

Wild yam 

Woodnettles 

Scientific Name 

Rubus sp. 

Vaccinium spp. L. 

Ranunculus L. spp. 

Galium Aparine L. 

Potentilla simplex Michx. 

Cuscuta gronovii Willd. 

Panicum clandestinum L. 

Aralia spinosa L. 

Circaea quadrisulcata (Maxim.) 

Franch. and Sav. 

Erechtites hieracifolia L. Raf. 

Rhododendron calendulaceum 

(Michx.) Torr. 

Lobelia inflata L. 

Polygonum spp. L. 

Asplenium montanum Willd. 

Carex plantaginea Lam. 

Phytolacca americana L. 

Pilea pumila (L.) Gray 

Dyropteris spinulosa (0. F. Muell.) 

Watt 

Gaultheria procumbens L. 

Trillium L. spp. 

Dentaria diphylla Michx. 

Oxalis structa L. 

Viola spp. L. 

Geum virginianum L. 

Eupatorium rugosum Houtt. 

Lysimachia quadrifolia L. 

Geranium maculatum L. 

Vitis L. spp. 

Sedum ternatum Michx. 

Dioscorea villosa L. 

La~ortea canadensis (L.) Wedd. 

7 

*U. S. GOVERNMENT PRINTING OFFICE: 1987/748-076/60002 
Wendel, G. W. 1987. Abundance and distribution of vegetation under four 

hardwood stands in north-central West Virginia. Res. Pap. NE-607. 

Broomall, PA: U.S. Department of Agriculture, Forest Service, Northeastern Forest Experiment Station. 6 p. 

Forest floor samples were collected from four hardwood forest stands in 

West Virginia to study species composition, abundance, and distribution of 

vegetation that originated from seeds, rootstocks, rhizomes, and so on. The 

abundance and distribution of plants on square-foot sections of forest floor 

that were lifted and moved to the greenhouse indicate that under the most 

ideal conditions upwards of 800,000 stems per acre may occur during the 

first year. Sweet birch was the most abundant tree species, blackberry the 

most abundant shrub, and species of violets were the most abundant herbaceous plants. Implications of this vegetation of the regeneration of the 

forest are discussed. 
Headquarters of the Northeastern Forest Experiment Station are in Broomall, Pa. 

Field laboratories are maintained at: 

0 Amherst, Massachusetts, in cooperation with the University of Massachusetts. 

0 Berea, Kentucky, in cooperation with Berea College. 

0 Burlington, Vermont, in cooperation with the University of Vermont. 

0 Delaware, Ohio. 

0 Durham, New Hampshire, in cooperation with the University of New Hampshire. 

Hamden, Connecticut, in cooperation with Yale University. 

0 Morgantown, West Virginia, in cooperation with West Virginia University, 

Morgantown. 

0 Orono, Maine, in cooperation with the University of Maine, Orono. 

0 Parsons, West Virginia. 

0 Princeton, West Virginia. 

0 Syracuse, New York, in cooperation with. the State University of New York 

College of Environmental Sciences and Forestry at Syracuse University, 

Syracuse. 

0 University Park, Pennsylvania, in cooperation with the Pennsylvania State 

University. 

Warren, Pennsylvania. 

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should be reported to the Secretary of Agriculture, Washington, DC 20250.