United States

Department of 

Agriculture

Forest Service

Northern 

Research Station

Research Paper NRS-9

Seed Bank Response to 

Prescribed Fire in the 

Central Appalachians

Thomas M. Schuler

Melissa Thomas Van-Gundy

Mary B. Adams

W. Mark Ford
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The Authors

THOMAS M. SCHULER, research forester, is currently engaged in research on the ecology 

and silviculture of Appalachian hardwood forests and the restoration of central Appalachian 

spruce forests at the Northern Research Station’s Timber and Watershed Laboratory in 

Parsons, West Virginia.

MELISSA THOMAS-VAN GUNDY, research forester, joined the Northern Research Station’s 

Timber and Watershed Laboratory at Parsons, West Virginia in 2006 and is engaged in 

research on management and ecology of Appalachian forests.

MARY B. ADAMS, project leader and research soil scientist, is currently engaged in 

research addressing problems of air pollution effects on forests, sustainability of mixed 

species forests in the face of many stressors, and basic nutrient cycling in forests. She 

works with the Northern Research Station at the Fernow Experimental Forest, Timber and 

Watershed Laboratory in Parsons, West Virginia.

W. MARK FORD, research wildlife biologist with the U.S. Army Engineer Research and 

Development Center, is currently engaged in research on environmental threat assessment 

to Department of Defense lands.

Manuscript received for publication 12 August 2009

Cover Photo

Conditions in the study area in June 2005 following the second prescribed fire on the 

Fernow Experimental Forest (April 2005). Note the absence of understory trees and 

the relatively sparse forest floor. The large wound in the tree at right is unrelated to the 

prescribed fire. Photo by Thomas M. Schuler, U.S. Forest Service.

Abstract

We compared pre- and post-treatment seed bank characteristics of woody species after 

two prescribed fires in a mesic mixed-oak forest in the central Appalachians. Nineteen 

woody species were identified from soil samples. Mean species richness and diversity 

declined but evenness did not. The seed bank was dominated by black birch (Betula 

lenta), yellow-poplar (Liriodendron tulipifera), blackberry (Rubus spp.), grapevine (Vitis

spp.), and Hercules club (Aralia spinosa) before burning. Following burning, the median 

density of seed bank propagules declined by 45 percent (247.6 to 138.1 m-2). Black birch, 

yellow-poplar, and grapevine declined by 69, 56, and 40 percent, respectively. The results 

illustrate the importance of the seed bank as a robust source of non-oak regeneration in 

this forest type and of the potential effect of fire in altering it. Oak restoration efforts that do 

not account for this significant source of regeneration in mesic mixed-oak forests likely will 

be unsuccessful.
1

Introduction

Successful regeneration of oak (Quercus spp.) is widely 

recognized as a serious problem throughout the 

hardwood regions of the Eastern and Central United 

States (Abrams 2000). Both managed and unmanaged 

forests show declining oak abundance as overstory oaks 

experience natural mortality or are harvested (Schuler 

2004). Other woody species such as red maple (Acer 

rubrum) have become increasingly abundant over the 

past several decades in number of stems and growingstock volume (Alderman and others 2005, Fei and 

Steiner 2007). Nowacki and Abrams (2008) referred to 

the widespread replacement of oaks with more mesic 

species as the “mesophication” of forests in the Eastern 

United States.

To counter the mesophication of oak forests, silvicultural 

guidelines using the shelterwood method have been 

developed to increase the size of existing oak seedlings, 

as large oak seedlings are more competitive with other 

vegetative competitors following canopy reductions 

(Loftis 1990, Brose and others 1999, 2008). Treatments 

are commonly needed to reduce non-oak competition 

and increase oak seedling size before significant canopy 

reductions. Loftis (1990) proposed a preharvest herbicide 

application to remove understory and midcanopy trees. 

More recently, prescribed fire has been recommended 

to reduce non-oak competition and improve the 

competitiveness of oak seedlings (Abrams 1992, 

Brose and Van Lear 1998, Hutchinson and others 

2005). Factors related to fire effectiveness include site 

characteristics, frequency and intensity, insects, and 

native and nonnative invasive species (Brose and others 

2006, Glasgow and Matlack 2007).

Enhancing oak competitiveness through prescribed 

burning is more likely on sites with xeric characteristics 

(Thomas-Van Gundy and others 2007). In part, mesic 

sites dry more slowly and develop less flammable 

forest litter (non-oak species), which makes it difficult 

to conduct a successful prescribed fire (Nowacki and 

Abrams 2008). There might be significant additional 

resistance to oak restoration efforts on mesic sites due to 

the extant seed bank. Seed banks are an important aspect 

of site characteristics that can play a critical role in the 

regeneration process, but the effect of prescribed fire on 

seed bank dynamics has only recently been explored.

In the relatively xeric Ozark Mountains of Arkansas, 

Schuler and Liechty (2008) found a low abundance of 

seed bank propagules in a mixed-oak forest, both woody 

and herbaceous, before and after prescribed burning. 

However, in more mesic conditions, seed banks can store 

substantial quantities of viable seed of both shrub and 

arboreal species (Wendel 1987, Adams and others 2004), 

illustrating the capacity of the seed bank to store non-oak 

propagules and interfere with oak restoration. Little is 

known about the extent to which prescribed fire alters 

seed bank characteristics in the central Appalachians and 

thus its effectiveness in promoting oak regeneration.

In this study we compared seed bank characteristics 

before and after prescribed burning in a mesic mixedoak forest in West Virginia. Unlike studies that have 

assessed the seed bank indirectly by reporting the number 

of new seedlings following burning (Hutchinson and 

others 2005), we sampled the forest floor before and after 

burning to gain a better understanding of the effects of 

fire on the seed bank itself. We report the number of 

propagules, species, and diversity characteristics before 

and after two prescribed fires. This research is part 

of a larger, ongoing study designed to determine the 

feasibility of using prescribed fire in conjunction with 

shelterwood harvesting in the central Appalachians to 

facilitate the sustainability of mixed-oak forests.

Methods

Site Description

The study was conducted in the Canoe Run watershed 

of the Fernow Experimental Forest (39.03o N, 79.67o 

W) in east-central West Virginia. The Fernow is in 

the Allegheny Mountains of the Central Appalachian 

Broadleaf Forest (McNab and Avers 1994). The 

mean growing season is 145 days (May to October); 

mean annual precipitation is about 1,430 mm and is 

distributed evenly throughout the year (Pan and others 

1997). Temperatures during the growing season usually 

are moderate and moisture deficits during the growing 

season are uncommon (Leathers and others 2000). 

Species composition in the forest overstory is complex 
2

throughout much of the Fernow and best described as 

mixed-mesophytic (Braun 1950). However, within this 

smaller study area, northern red oak (Quercus rubra), 

chestnut oak (Q. prinus), and white oak (Q. alba) are the 

most common overstory species in descending order of 

importance as measured by basal area. Other common 

associates in the study area are red maple, sugar maple (A. 

saccharum), black birch (Betula lenta), and yellow-poplar 

(Liriodendron tulipifera). Treatment areas encompass 

approximately 31 ha and site elevations range from about 

600 to 770 m asl. Site index is about 70 for northern red 

oak (Schnur 1937) throughout the study area.

Experimental Design and Data Analysis 

As part of a larger and ongoing study, we monitored the 

response of the seed bank on twenty 0.202-ha rectangular 

permanent growth plots treated with prescribed fire.

Within each growth plot we sampled the forest floor 

and the upper horizon of the soil at randomly located 

distances along a transect running across each growth 

plot during March of 2000 (before prescribed fire) and 

in March 2007 (after two prescribed fires). We followed 

the methodology of Wendel (1987). We collected four 

samples per transect plot in 2000 and three samples per 

plot in 2007. Samples of the forest floor were cut with a 

shovel to a depth of 10 cm around the inside of a wooden 

template (30 by 30 cm). The samples were placed in 

wooden trays with a mesh hardware cloth bottom and 

lined with paper. We labeled samples as to location 

and plot and then placed flats on greenhouse benches 

randomly (Fig. 1); the flats were watered as needed. Seeds 

that germinated and or root stocks that sprouted were 

allowed to grow until summer to allow identification and 

counts of tree and shrub species in each tray. Nothing 

else was done to stimulate germination nor did we 

attempt to census herbaceous vegetation that was present.

We calculated Shannon-Weiner’s diversity index (H’) 

(Whittaker 1972), Pielou’s (1969) evenness index 

(J’), and species richness based on species presence 

and density for each experimental unit through 

time (Magurran 1988); often seed bank data are not 

distributed normally (Hyatt 1999) and most species in 

our sample did not exhibit normality. As a result, we used 

the Wilcoxon signed-rank test to test for differences in 

species abundance before and after prescribed burning. 

The null hypothesis is that the median difference between 

the pair of observations is zero. We used a paired t-test 

Figure 1.—Samples of forest floor on greenhouse benches allowed identification and counts of 

tree and shrub species.

Mary Beth Adams, U.S. Forest Service
3

to examine changes in diversity measures related to fire 

treatments through time after determining that patterns 

of normality and homogeneity of variance had been met. 

The level of statistical significance for all tests was alpha = 

0.05. It should be noted that the samples before and after 

prescribed fire do not account for changes that might 

have occurred in the absence of fire.

Prescribed Fire: Treatments and 

Characteristics

The study area was treated with two prescribed fires just 

before the growing seasons in 2002 and 2003 and again 

in 2005. Because the first prescribed fire in April 2002 

was interrupted due to poor weather, we were able to 

burn only one-half of the site. The remaining portion 

was burned the following spring (2003). We conducted 

a second prescribed fire in April 2005 so that the entire 

study site has been burned twice. We used the strip-head 

fire technique; ignition was by hand-held drip torches. 

We wanted to generate flame lengths of about 1 m with a 

rate of spread (ROS) of about 1 to 3 m per minute based 

on BEHAVE (v. 3.0.2) predictions using Fuel Model 9. 

Meteorological data at the time of the fires were obtained 

from a U.S. Environmental Protection Agency weather 

station (PAR107) located about 4.5 km north of the 

study area.

We monitored fire behavior at 10 locations for each 

prescribed burn. We used Hobo Type K data loggers 

connected to stainless steel probes 25 cm long with 

an isolated thermocouple at the tip (Onset Computer 

Corp.).1

 This methodology is similar to that used by 

Iverson and others (2004). Temperatures were recorded 

every 2 seconds by the probe and data logger and were 

recorded at five points at each instrumented growth plot 

during the fires. We set temperature probes at each corner 

on a square grid (9.1 by 9.1 m) and one in the center 

of the grid. Because the data loggers had synchronized 

clocks, we were able to calculate the fire’s rate of spread 

at each plot location using the elapsed time between the 

rapid initial fire-induced elevated temperatures (> 32  ̊C) 

logged by adjacent data loggers. We have not attempted 

to correlate fire effects with fire characteristics at the 

plot level because not all plots were instrumented with 

temperature probes, and probe grids were not sized 

consistently with our seed bank measurements. However, 

we do report general characteristics of fire behavior 

for each fire for comparison with similar studies as 

recommended by Brose and others (2006).

Differences in factors such as air temperature, relative 

humidity, and wind speed (Table 1) contributed to 

variability in fire behavior. We observed mostly moderate 

or less intense fire behavior during all of our burns 

(i.e., flame lengths less than 1 m stemming from the 

combustion of leaf litter and 1-hour surface fuels). Actual 

fire spread rates were greater than those predicted by 

modeling but were similar to those reported in Ohio 

(Iverson and others 2004).

1

The use of trade, firm, or corporation names in this 

publication is for the information and convenience of the 

reader. Such use does not constitute an official endorsement or 

approval by the U.S. Department of Agriculture or the Forest 

Service of any product or service to the exclusion of others that 

may be suitable.

Table 1.—Weather and fire behavior characteristics for prescribed burns on the Fernow 

Experimental Forest, West Virginia. Air temperature (Air) and relative humidity (RH) given for start 

and finish times of fires; rate of spread (ROS), maximum probe temperature (PT), and wind speed 

(WS) are means with standard deviations in parentheses

Date Air RH ROS PT WS No. days 

since rain

o

C Percent m/min oC km/hr

4/12/2002 23 - 22 38 - 39 9 (4) 302 (153) 4 (3) 3

4/4/2003 21 - 17 41 - 64 15 (15) 251 (88) 9 (2) 5

4/11/2005 21 - 22 13 - 12 44 (83) 327 (122) 8 (2) 3

4/15/2005 14 - 13 21 - 23 12 (9) 324 (66) 14 (2) 7
4

Results

The seed bank was dominated by black birch, yellowpoplar, blackberry (Rubus spp.), grapevine (Vitis spp.), 

and Hercules club (Aralia spinosa). These five species 

comprised 70 and 76 percent of seed bank propagules 

before and after prescribed burning, respectively. 

Nonetheless, following prescribed burning, the median 

density of seeds and other vegetative propagules in 

the seed bank was reduced significantly by 44 percent 

(Table 2), according to the Wilcoxon signed-rank test 

results. Birch, yellow-poplar, and grapevine abundance 

declined significantly by 69, 56, and 40 percent, 

respectively (Table 2). Conversely, we found no evidence 

that blackberry abundance was affected by prescribed 

burning as afterward it became the most abundant 

species due to the decreased abundance of other species. 

Only black locust (Robinia pseudoacacia) exhibited 

significantly greater abundance after burning than 

before; however, it remained a relatively minor species, 

accounting for slightly more than 2 percent of the overall 

sample in 2007.

We identified 19 woody species from the soil samples. 

Mean species richness declined significantly from 8.35 

to 6.60 after burning (P < 0.001) based on results of the 

paired t-test. H’ also declined after burning, from 1.676 

to 1.421 (P = 0.005), though J’ did not (P = 0.241). 

American beech (Fagus grandifolia), chestnut oak, 

sourwood (Oxydendrum arboreum), Fraser magnolia 

(Magnolia fraseri), and sumac (Rhus sp.) were present, 

though not abundant, before prescribed burning, but 

absent afterward and contributed to the decline in 

mean species richness. Other species of relatively low 

abundance in the sample included white and red oak, 

cucumbertree (Magnolia accuminata), sassafras (Sassafras 

albidum), black cherry (Prunus serotina), striped maple 

(Acer pensylvanicum), red maple, and greenbrier (Smilax

sp.). We found no evidence that the abundance of 

these species in the seed bank was altered by prescribed 

burning.

Discussion

Following prescribed fire, seeds in the forest floor may 

germinate, remain viable but not germinate, or be killed 

by the fire (Fig. 2). At the inception of this study, we 

knew that the seed bank was a robust source of woody 

regeneration but there was little information on how 

repeated prescribed fires might affect it. We hypothesized 

that repeated fires reduces the number of viable seeds in 

the seed bank and fundamentally alters post-fire stand 

dynamics. Here we focus on the seed bank itself because 

little is known about the direct effects of fire on the seed 

bank in mesic forests. The seed bank may contribute to 

new seedlings following perturbations to the forest, for 

example during different stages of a shelterwood harvest. 

Before the onset of burning, seeds of black birch and 

yellow-poplar together totaled nearly 100 m-2, reflecting 

values reported for the central Appalachians by Wendel 

Table 2.—Median density of seed bank propagules of most abundant species before and 

after two prescribed fires in a mesic mixed-oak forest in West Virginia (minor species 

not listed individually but included in total)

Number of propagules m-2

Species Pre-burn Post-burn Pa

Betula lenta 47.1 14.4 <0.01

Rubus spp. 45.7 53.8 0.82

Liriodendron tulipifera 44.4 19.7 <0.01

Vitis spp. 24.2 14.4 0.04

Aralia spinosa 12.1 3.6 0.18

Sassafras albidum 5.4 0 <0.01

Acer rubrum 4.0 0 0.08

Robinia pseudoacacia 0 3.6 <0.01

Total 247.6 138.1 < 0.01

a

Wilcoxon signed-rank test
5

(1987) and Adams and others (2004). Seed densities for 

black birch and yellow-poplar were reduced significantly 

following prescribed burning but remained above 30 

m-2. Thus, even after two prescribed fires, the seed bank 

remains an abundant source of potential tree regeneration 

that may overwhelm many competitors, including 

small oak seedlings (Beck and Hooper 1986, Brashears 

and others 2004). In more xeric mixed-oak forests it 

has been reported that the seed bank harbors an order 

of magnitude less of seeds of tree species, regardless of 

recent fire history (Schiffman and Johnson 1992, Schuler 

and Liechty 2008). Seed bank differences between 

xeric and mesic mixed-oak forests create fundamentally 

different challenges to sustaining oak species. In xeric 

forests, we believe the primary challenge is to overcome 

the buildup of advanced regeneration of shade-tolerant 

species. In mesic conditions, a significant additional step 

is to overcome the buildup of non-oak competitors in the 

seed bank.

Although the seed bank we studied was robust, overall 

abundance decreased substantially after burning due to 

decreases in black birch, yellow-poplar, and grapevine 

even though post-fire sampling occurred 2 years after 

the last prescribed fire. During this time, the seed bank 

could have been partially replenished from surrounding 

overstory trees. As a result, our estimates may be 

conservative regarding total seed reductions. However, 

the significant reduction in black birch is intriguing 

because past research demonstrated that prescriptions 

designed to enhance oak seedling development also 

enhanced birch seedling germination and development, 

and birch can be a particularly problematic species 

in oak regeneration (Schuler and Miller 1995). Both 

black birch and the primary oak species in the central 

Appalachians are intermediate in shade tolerance 

(Trimble 1975) and can respond positively to partial 

reductions in overstory density (Thomas-Van Gundy 

and Schuler 2008). Ecologically significant seed 

bank reductions of black birch and yellow-poplar 

should equate to a higher incidence of understory oak 

persistence and greater probability of oak recruitment 

into the overstory. We cannot yet conclude that the 

significant reductions in birch and yellow-poplar we 

documented are ecologically significant in meeting goals 

for oak restoration.

Figure 2.—Typical forest floor conditions in June 2005 following the second prescribed fire 

(April 2005). 

Thomas M. Schuler, U.S. Forest Service
6

The implication with regard to blackberry resilience to 

prescribed burning in our study is not fully understood. 

Blackberry is a common early successional, shadeintolerant shrub that persists until trees species emerge 

through the shrub layer and form a continuous canopy 

over it. Dense blackberry thickets can compete vigorously 

with hardwood seedlings (Whitney 1986), but also may 

benefit hardwood seedlings by making them less visible 

to browsers. Blackberry is a true seed banker in that some 

seeds may remain viable in the seed bank for more than 

a century. However, seeds are less abundant in stands 

older than 95 years (Graber and Thompson 1978). We 

speculate that higher levels of blackberry abundance 

in the seed bank relative to some tree species do not 

preclude oak seedling persistence and development, but 

additional research is needed to understand how oak and 

blackberry interact following prescribed fire in the mixedoak forest type.

In areas where the seed bank does constitute an 

ecologically important source of non-oak propagules, 

particularly when it includes species of intermediate 

shade tolerance such as black birch, efforts to reduce the 

role of the seed bank in the regeneration process likely 

will be required to achieve management goals for oak. 

Seeds from most tree species are considered transient 

in temperate woodlands (Thompson 1992), but the 

annual seed input into the forest floor from common 

oak competitors in contemporary forests effectively 

creates a persistent seed bank. Black birch and yellowpoplar may persist for as many as 8 years in the seed 

bank (Beck and Hooper 1986), and these species may 

produce abundant seed crops nearly every year (Olson 

1969, Brinkman 1974). Accordingly, efforts to reduce 

seed bank propagules (Shearin and others 1972) that 

do not address seed bank input, likely will have limited 

temporal effect. Repeated landscape-level burning by 

Native Americans and settlers (Van Lear and Waldrop 

1989) may have altered seed bank dynamics by altering 

overstory composition at spatial scales large enough to 

slow renewal. To mimic this process today, it is essential 

to consider the recruitment potential of species from 

both within and outside of the area being managed as 

well as the spatial scale and shape of the area needed to 

achieve desired results. Small management units with 

a high proportion of edge are unlikely to inhibit seed 

bank renewal from external sources. Future research may 

evaluate the size and shape of burn units that are required 

to slow renewal.

In the past decade, prescribed burning of eastern 

mixed-oak forests has become a more widely accepted 

restoration approach as indicated by the number of 

new National Forest management plans that have 

incorporated fire into management objectives (Yaussy 

and others 2008). The continued use of fire to promote 

oak regeneration and restoration will hinge on a better 

understanding of when, where, and how to effectively 

apply prescribed fire in conjunction with other 

treatments. In West Virginia, landscape modeling points 

to the relatively more xeric conditions of the Ridge and 

Valley Province as the most likely area in which mixedoak forests will be sustained through management 

intervention (Thomas-Van Gundy and others 2007). 

However, Nowacki and Abrams (2008) asserted that 

prescribed burning on mesic oak sites should be a high 

priority because species shifts are occurring more rapidly 

and will be more difficult to reverse than on xeric sites. 

They concluded that an alternative stable-state in the 

composition of mesophytic hardwood species will occur 

in the absence of fire. We assert that this stability may 

be linked functionally to seed bank characteristics that 

develop in advance of other factors already identified 

by Nowacki and Abrams (2008) such as less flammable 

forest litter of non-oak species. It is unclear whether the 

alternative stable-state favoring mesophytic hardwoods 

has already occurred when and where the seed bank 

is an abundant source of non-oak propagules, but 

oak restoration efforts that do not account for this 

potentially significant source of regeneration in mesic 

mixed-oak forests likely will be unsuccessful. Although 

the inferences from our study are limited due to the 

spatial scale of our experimental design, the results 

illustrate the potential importance of the seed bank 

in the oak regeneration process. Research is needed to 

determine where the seed bank is a limiting factor to 

oak sustainability and whether prescribed fire and other 

silvicultural treatments can be used to change seed bank 

characteristics in favor of oak and sympatric species.
7

Literature Cited

Abrams, M.D. 1992. Fire and the development of oak 

forests. Bioscience. 42(5): 346-353.

Abrams, M.D. 2000. Fire and the ecological history 

of oaks in the eastern United States. In: Yaussy, D., 

comp. Proceedings: workshop on fire, people, and 

the central hardwoods landscape; 2000 March 13-14; 

Richmond, KY. Gen. Tech. Rep. NE-274. Newtown 

Square, PA: U.S. Department of Agriculture, Forest 

Service, Northeastern Research Station: 46-55.

Adams, M.B.; Burger, J.; Zelazny, L.; Baumgrass, J. 

2004. Description of the Fork Mountain longterm soil productivity study: site characterization. 

Gen. Tech. Rep. NE-323. Newtown Square, PA: 

U.S. Department of Agriculture, Forest Service, 

Northeastern Research Station. 40 p.

Alderman, D.R.; Bumgardner, M.S.; Baumgrass, J.E. 

2005. An assessment of the red maple resource in 

the northeastern United States. Northern Journal of 

Applied Forestry. 22: 181-189.

Brinkman, K.A. 1974. Betula L. Birch. In: Schopmeyer, 

C.S., tech. coord. Seeds of woody plants in the United 

States. Agric. Handb. 450. Washington, DC: U.S. 

Department of Agriculture: 252-257.

Beck, D.E.; Hooper, R.M. 1986. Development of 

a southern Appalachian hardwood stand after 

clearcutting. Southern Journal of Applied Forestry. 

10: 168-172.

Brashears, M.B.; Fajvan, M.A.; Schuler, T.M. 2004. 

An assessment of canopy stratification and tree 

species diversity following clearcutting in Central 

Appalachian hardwoods. Forest Science. 50(1): 54-

64.

Braun, E.L. 1950. Deciduous forests of eastern North 

America. Philadelphia, PA: Blakiston Co. 596 p.

Brose, P.H.; Van Lear, D.H. 1998. Responses of 

hardwood advance regeneration to seasonal 

prescribed fires in oak-dominated shelterwood 

stands. Canadian Journal of Forest Research. 28: 331-

339.

Brose, P.; Van Lear, D.; Cooper, R. 1999. Using 

shelterwood harvests and prescribed fire to 

regenerate oak stands on productive upland sites.

Forest Ecology and Management. 113: 125-141.

Brose, P.H.; Schuler, T.M.; Ward, J.S. 2006. Responses 

of oak and other hardwood regeneration to 

prescribed fire: what we know as of 2005. In: 

Dickinson, M.B., ed. Fire in oak forests: delivering 

science to land managers: proceedings of a conference; 

2005 November 15-17; Columbus, OH. Gen. 

Tech. Rep. NRS-P-1. Newtown Square, PA: U.S. 

Department of Agriculture, Forest Service, Northern 

Research Station: 123-135.

Brose, P.H.; Gottschalk, K.W.; Horsley, S.B.; Knopp, 

P.D.; Kochenderfer, J.N.; McGuinness, B.J.; Miller, 

G.W.; Ristau, T.E.; Stoleson, S.H.; Stout, S.L. 2008. 

Prescribing regeneration treatments for mixed-oak 

forests in the Mid-Atlantic region. Gen. Tech. Rep. 

NRS-33. Newtown Square, PA: U.S. Department 

of Agriculture, Forest Service, Northern Research 

Station. 100 p.

Fei, S.; Steiner, K. 2007. Evidence for increasing red 

maple abundance in the eastern United States. 

Forest Science. 53: 473-477.

Glasgow, L.S.; Matlack, G.R. 2007. The effects of 

prescribed burning and canopy openness on the 

establishment of two non-native plant species in 

a deciduous forest, southeast Ohio, USA. Forest 

Ecology and Management. 238: 319-329.

Graber, R.E.; Thompson, D.F. 1978. Seeds in the 

organic layers and soil of four beech-birchmaple stands. Res. Pap. NE-401. Broomall, PA: 

U.S. Department of Agriculture, Forest Service, 

Northeastern Forest Experiment Station. 8 p.

Hutchinson, T.F.; Sutherland, E.K.; Yaussy, D.A. 

2005. Effects of repeated fires on the structure, 
8

composition, and regeneration of mixed-oak forests 

in Ohio. Forest Ecology and Management. 218: 210-

228.

Hyatt, L.A. 1999. Differences between seed bank 

composition and field recruitment in a temperate 

zone deciduous forest. American Midland Naturalist. 

142(1): 31-38.

Iverson, L.R.; Yaussy, D.A.; Rebbeck, J.; Hutchinson, 

T.F.; Long, R.P.; Prasad, A.M. 2004. A comparison of 

thermocouples and temperature paints to monitor 

spatial and temporal characteristics of landscapescale prescribed fires. International Journal of 

Wildland Fire. 13: 311-322.

Leathers, D.J.; Grundstein, A.J.; Andrew, W.E. 2000. 

Growing season deficits across the northeastern 

United States. Climate Research. 14: 43-55.

Loftis, D.L. 1990. A shelterwood method for 

regenerating red oak in the southern Appalachians.

Forest Science. 36: 917-929.

Magurran, A.E. 1988. Ecological diversity and its 

measure. Princeton, NJ: Princeton University Press. 

912 p.

McNab, W.H.; Avers, P.E.. 1994. Ecological subregions 

of the United States: section descriptions. Admin. 

Publ. WO-WSA5. Washington, DC: U.S. Department 

of Agriculture, Forest Service. 267 p.

Nowacki, G.J.; Abrams, M.D. 2008. The demise of 

fire and “mesophication” of forests in the eastern 

United States. Bioscience. 58: 123-138.

Olson, D.F., Jr. 1969. Silvical characteristics of yellowpoplar (Liriodendron tulipifera L.). Res. Pap. SE-48. 

Asheville, NC: U.S. Department of Agriculture, Forest 

Service, Southeastern Forest Experiment Station. 16 p.

Pan, C.; Tajchman, S.J.; Kochenderfer, J.N. 1997. 

Dendroclimatological analysis of major forest 

species of the central Appalachians. Forest Ecology 

Management. 98: 77-87.

Pielou, E.C. 1969. An introduction to mathematical 

ecology. New York: John Wiley and Sons. 286 p.

Schiffman, P.M; Johnson, W.C. 1992. Sparse buried 

seed in a southern Appalachian oak forest: 

implications for succession. American Midland 

Naturalist. 127: 258-267.

Schnur, G.L. 1937. Yield, stand, and volume tables 

for even-aged upland oak forests. Tech. Bull. 560. 

Washington, DC: U.S. Department of Agriculture. 

88 p.

Schuler, J.L.; Liechty, H.O.. 2008. Seed bank 

emergence following prescribed burning in the 

Ozark Highlands. In: Jacobs, Douglas F; Michler, 

Charles H., eds. Proceedings, 16th central hardwood 

forest conference; 2008 April 8-9; West Lafayette, 

IN. Gen. Tech. Rep. NRS-P-24. Newtown Square, 

PA: U.S. Department of Agriculture, Forest Service, 

Northern Research Station: 516-524.

Schuler, T.M. 2004. Fifty years of partial harvesting 

in a mixed mesophytic forest: composition and 

productivity. Canadian Journal of Forest Research. 

34: 985-997.

Schuler, T. M.; Miller, G.W. 1995. Shelterwood 

treatments fail to establish oak reproduction on 

mesic forest sites in West Virginia. In: Gottschalk, 

K.W.; Fosbroke, S.L.C., eds. Proceedings, 10th 

central hardwood conference. Gen. Tech. Rep. 

NE-197. Newtown Square, PA: U.S. Department 

of Agriculture, Forest Service, Northeastern Forest 

Experiment Station: 375-388.

Shearin, A. T.; Bruner, M.H.; Goebel, N.B. 1972. 

Prescribed burning stimulates natural regeneration 

of yellow-poplar. Journal of Forestry. 70: 482-484.

Thomas-Van Gundy, M.; Nowacki, G.J.; Schuler, 

T.M. 2007. Rule-based mapping of fire adapted 

vegetation and fire regimes for the Monongahela 

National Forest. Gen. Tech. Rep. NRS-12. Newtown 

Square, PA: U.S. Department of Agriculture, Forest 

Service, Northern Research Station. 24 p.
9

Thomas-Van Gundy, M.; Schuler, T.M. 2008. Deferred 

rotation harvests in central Appalachia: 20- and 25-

year results. In: Jacobs, Douglas F.; Michler, Charles 

H., eds. Proceedings, 16th central hardwood forest 

conference; 2008 April 8-9; West Lafayette, IN. Gen. 

Tech. Rep. NRS-P-24. Newtown Square, PA: U.S. 

Department of Agriculture, Forest Service, Northern 

Research Station: 423-435.

Thompson, K. 1992. The functional ecology of seed 

banks. In: Fenner, M., ed. Seeds: the ecology of 

regeneration in plant communities. Wallingford, UK: 

CAB International: 231-258.

Trimble, G.R. 1975. Summaries of some silvical 

characteristics of several Appalachian hardwood 

trees. Gen. Tech. Rep. NE-16. Upper Darby, PA: 

U.S. Department of Agriculture, Forest Service, 

Northeastern Forest Experiment Station. 5 p.

Van Lear, D.H.; Waldrop, T.A. 1989. History, uses, 

and effects of fire in the Appalachians. Gen. Tech. 

Rep. SE-54. Asheville, NC: U.S. Department of 

Agriculture, Forest Service, Southeastern Forest 

Experiment Station. 20 p.

Wendel, G.W. 1987. Abundance and distribution of 

vegetation under four hardwood stands in northcentral West Virginia. Res. Pap. NE-607. Broomall, 

PA: U.S. Department of Agriculture, Forest Service, 

Northeastern Forest Experiment Station. 7 p.

Whitney, G.G. 1986. A demographic analysis of Rubus 

idaeus and Rubus pubescens. Canadian Journal of 

Botany. 64: 2916-2921.

Whittaker, R.H. 1972. Evolution and measurement of 

species diversity. Taxon. 21: 213-251.

Yaussy, D.A.; Nowacki, G.J.; Schuler, T.M.; Dey, 

D.C.; DeGayner, E.J. 2008. Developing a 

unified monitoring and reporting system: a key 

to successful restoration of mixed-oak forests 

throughout the central hardwood region. In: 

Deal, R.L., ed. Proceedings of the 2007 national 

silviculture workshop: integrated restoration of 

forested ecosystems to achieve multi-resource benefits. 

Gen. Tech. Rep. PNW-733. Portland, OR: U.S. 

Department of Agriculture, Forest Service, Pacific 

Northwest Research Station: 281-286.
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Schuler, Thomas M.; Thomas Van-Gundy, Melissa; Adams, Mary B.; Ford, W. Mark. 

2010. Seed bank response to prescribed fire in the central Appalachians.

Res. Pap. NRS-9. Newtown Square, PA: U.S. Department of Agriculture, Forest 

Service, Northern Research Station. 9 p.

Pre- and post-treatment seed-bank characteristics of woody species were 

compared after two prescribed fires in a mesic mixed-oak forest in the central 

Appalachians. Nineteen woody species were identified from soil samples. Mean 

species richness declined but evenness did not after prescribed burning. The 

seed bank was dominated by black birch, yellow-poplar, blackberry, grapevine 

and Hercules club before burning. Following burning, the median density of 

seed bank propagules declined by 45 percent. Black birch, yellow-poplar, and 

grapevine declined by 69, 56, and 40 percent, respectively. The results illustrate the 

importance of the seed bank as a robust source of non-oak regeneration in mixedoak forests and of the potential effect of fire in altering it.

KEY WORDS: mixed-mesophytic, West Virginia, oak regeneration, forest restoration
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