USDA United States

m

Department of

 Agriculture

Forest Service

Northeastern

Area

Morgantown, WV

NA-TP-04-99

Changes in Composition

of the

Mixed Mesophytic Forest:

Effects of Succession and Disturbance

Jim Steinman

USDA Forest Service

Northeastern Research Station

Radnor, Pennsylvania

January 1999
Changes in Composition

of the

Mixed Mesophytic Forest:

Effects of Succession and Disturbance

Abstract

This study investigated the hypothesis that air pollution is causing mortality of the larger

overstory trees, which results in a shift in species composition. To determine if the theorized

shifts in species composition have occurred, this study compared historical changes in forest

composition as described by Braun (1940) with more recent changes as quantified by the

Forest Inventory Analysis (FIA) program of the USDA Forest Service. FIA estimated recent

composition changes using records of live, dead, and cut trees from 5,404 randomly sampled

plots.

Analyses suggest that the forest overstory consisted mostly of late-successional species in the

1940s and early- and mid-successional species in the 1980s. Thus, differences were most

likely due to disturbances (insects and diseases, fire, weather, pollution) that killed trees,

which allowed pioneer species to occupy openings. Forest succession may account for the 6

percent of dead trees in the 1980s since the percentages of dead trees were significantly

greater among early-successional species. Percentages and spatial gradients of dead trees of

species tolerant to air pollutants were similar to dead trees of intolerant species. Most of the

4 percent of all trees cut in the 1980s were not late-successional species, which may have

favored successional trends.

Keywords: mixed mesophytic forest, tree mortality, succession, disturbances, air pollutants
Table of Contents

Introduction . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . I

Role of Air Pollution . . . . . . . . . ..~..............~................~.....~........~.~...,..,.,,..,,,.,,,.,.,,,~,.,~, I

Study Design ..~.~...~.~.....~.~~..,~~....~..~~~......~..~..........~.........................~.....,,....~,.,...~. 2

Methods . ..~~.....~.,.,,~,.,,,..,.~,,,....,.,......,.,.......,.~.........,~...~..~.....,~..~.,~,.~,,,,..,,,,.,.~~.,., 2

Study Area .......................................................................................................... 2

Braun Historical Data .......................................................................................... 4

FIA Data ............................................................................................................. 4

Analytical Procedure ........................................................................................... 5

Results and Discussion ...................................................................................... 7

1. Differences in Species Composition in the Last 50 Years ............................. 7

Historical Changes in Southeastern Kentucky .................................................. 7

FIA Data: Composition of Mixed Mesophytic Forest ......................................... 8

2. Rate of Tree Mortality .................................................................................... 9

This section of the study ascertained whether trees were dying

at an accelerated rate ................................................................................. 9

Mortality Rate of Different Species .................................................................. IO

Rate of Mortality in Large Trees...................................................................... 11

Mortality by Geographic Locations .................................................................. 11

3. Causes of Recent Tree Mortality ................................................................. 18

Forest Succession........................................................................................... 18

Logging and Land C/earing ............................................................................. 18

Insects, Diseases, Weather, and Fire ............................................................. 19

Air Pollution ..................................................................................................... 20

4. The Future Forest.. ...................................................................................... 20

Conclusions ...................................................................................................... 20

References ...................................................................................................... .2 I
List of Tables

Table 1 - Characteristics of Ecological Subregions that Contain the Mixed Mesophytic Forest

(McNab and Avers 1994) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3

Table 2 -Distribution of FIA Sample Plots with Trees at least 10 inches DBH (by Forest Type

and Ecological Subregion) .,.....................................................,...................................................... 5

Table 3 -Distribution of all trees (live, dead, and cut) at least 10 inches DBH from 5404 FIA

sample plots located in oak-hickory, northern hardwood, and oak-pine forest types . . . . . . . . . . . . . . . . . . . . . 6

List of Figures

Figure 1. States containing the mixed mesophytic forest and counties sampled by Braun (1950). . . . 1

Figure 2. Delineations of the mixed mesophtyic forest (Braun 1950) and ecological subregions

(McNab and Avers 1994) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2

Figure 3. Species composition in southeastern Kentucky during the 1940s and 1988 as estimated

by Braun (1950) and from FIA sample plots, respectively. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 7

Figure 4. Species composition of oak-hickory forest types by tolerance to competition (FIA data). . . . 8

Figure 5. Species composition of northern hardwood forest types by tolerance to competition

(FIA data) , ..,....,,.....,,,....,......................................*.......................................................................... 9

Figure 6. Percentages of dead trees and removed trees in oak-hickory forest types by tolerance

to competition (FIA data) . ,....~..~~...~~.....~~.,.......~.....~~....I...~~..................~.................~,................~....~~ 1 0

Figure 7. Percentages of dead trees and removed trees in northern hardwood forest types by

tolerance t o competition (FIA data). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 1 1

Figure 8. Percentages of dead trees and removed trees by dbh class and tolerance to competition:

Intolerant (INT), moderately tolerant (MOD), and tolerant (TOL) (FIA data). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 11

Figure 9. Percentages of dead trees and removed trees in oak-hickory forest types by tolerance

to competition and ecological subregion (FIA data). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 12

Figure 10. Percentages of dead trees and removed trees in northern hardwood forest types by

tolerance to competition and ecological subregion (FIA data). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 12

Figures 11-15. FIA plots containing at least three trees greater than 10 inches dbh. . . . . . . . . . . . . . . ...13-17

Figure 16. Percentages of FIA plots containing dead trees of selected species for plots with and

without evidence of any removed trees . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 1 9
Introduction

The mixed mesophytic forest was described by E.

Lucy Braun (1950) as the portion of the Appalachian

region of the United States (Figure 1) that contains a

diverse mixture of mesophytes. Mesophytes are

plants that grow best on moist sites. Field studies

conducted by Braun in the 1940s included descriptions of the many tree species observed in the

overstory (see Appendix Table A-l), including

counts of overstory trees along sampled transects.

Summaries of Braun’s published counts (see Appendix Table B- 1) show species in the following order

of abundance:

1. American beech 6. White oak

2. Sugar maple 7. Basswood

3. Yellow-poplar 8. Chestnut oak

4. Eastern hemlock 9. Hickory

5. American chestnut 10. Yellow buckeye

Various stress factors have contributed to modify the

distribution and species composition (Martin 1992)

of the forest. As with all forest ecosystems, trees

compete for light, water, and nutrients; species

tolerant of competition tend to succeed while intolerant species die (Spurr and Barnes 1992). This

relatively slow process of competition and survival

is often abruptly interrupted by disturbances that kill

trees, create forest openings, alter species composition, and modify forest succession trends (Abrams

and Downs 1990; Abrams and Nowacki 1992).

Disturbances in the mixed mesophytic forest include

urban development, agriculture, logging, fire,

drought, wind storms, forest insects, and diseases

(Hicks and Mudrick 1993). The most influential tree

disease in recent years has been chestnut blight

[Cryphonectria parasitica (Mm-r.) Barr]. Chestnut

blight has virtually eliminated the American chestnut

tree from the forest overstory.

Role of Air Pollution

Air pollutants consisting of nitrogen, sulfur, and

ozone have been present in the United States forests

throughout the latter half of the 20th century. However, most studies to determine the effects of ambient air pollutants on mature forest trees have shown

inconclusive results due to the following factors:

l Difficulty and expense involved in conducting

controlled “cause and effect” field experiments.

l Lack of reference or benchmark data with which

r’

GA

Figure 1. States containing the mixed mesophytic forest and

counties sampled by Braun (1950).

to compare increases in air pollution

data.

l Reliance on evaluations that associate tree mortality and ambient air

pollution along spatial gradients

(Shriner and others 1990).

Studies of tree mortality and ambient

air pollution in the northern Appalachian region have only been able to

demonstrate a relationship between

mortality of red spruce and concentrations of air pollutants at high elevations. Even so, two hypotheses exist

for the mixed mesophytic forest.

These hypotheses state that: (1) larger

trees of some species are dying at an

accelerated rate, and (2) mortality is

due to the deposition of airborne

nitrogen and exposure to ozone, which

predispose trees to root pathogens

(Little 1995).

-l-
Study Design Methods

The current analysis focused on testing the hypothesis that trees of some species are dying at accelerated rates. Supportive analyses used quantitative

data from the Forest Inventory and Analysis (FIA)

program of the USDA Forest Service (Hansen and

others 1992). FIA provides the best available source

of high-quality and unbiased information obtained

from an extensive system of randomly selected

locations. Initial analyses were conducted to compare the overstory composition observed by Braun in

the 1940s with the composition observed in the

1980s as gleaned from the FIA data. Subsequent

analyses identified tree species and geographic

locations with percentages of dead trees or cut trees

Study Area

The location of the mixed mesophytic forest as

described by Braun (1950) closely corresponds to

delineations of ecological subregions (Table 1 and

Figure 2) defined by McNab and Avers (1994). Use

of subregion boundaries was desirable to facilitate

comparisons among natural physiographic and

climatic zones instead of political entities (e.g.,

states and counties). The southern extension of the

forest in ecological subregion 23 1 C of Alabama was

retained for analyses, although it has a warmer

climate and more loblolly pine than the other subregions. Likewise, the northern extension in 212G of

Pennsylvania was retained although it represents a

cooler climate than the rest of the forest and has a

higher proportion of black cherry.

that deviate from forest-wide averages. These

results were then used to infer historical differences

in species composition between the 1940s and

198Os, and the recent changes in species composition and forest succession trends.

While this study did not directly

address the hypothesis that air

pollutants cause trees to die,

FIA data were utilized to

identify likely causes of mortality based on the differential

proportions of dead trees among

species and locations. More

rigorous hypothesis testing

would require collection of data

to frequently monitor the

variability in tree health over

time across the study region,

and account for changes in

health and mortality due to

coincident effects of tree

competition and disturbances

such as forest insects and

diseases. Such data are currently

not available for the mixed

mesophytic forest although they

have been collected in other

regions by the ongoing Forest

Health Monitoring and North

American Maple Projects

(Twardus and Mielke 1995).

The Mixed Mesophytic Forest

Ecological Subregions

(k&Nab and Avers 1994)

Figure 2. Delineations of the mixed mesophtyic forest (Braun 1950)

and ecological subregions (M&lab and Avers 1994).

-2-
Table 1

Characteristics of Ecological Subregions that

Contain the Mixed Mesophytic Forest

(McNab and Avers 1994)

Paleozoic Potential Growing Disturbance

Ecological Geo- Parent Forest Elevation Precip (in) Season and

Subregion morphology Materials Soil Taxa Vegetation (ft) Temp (F) (days) Land Use

212 Laurentian Mixed Forest Province

212G dissected

plateau

sandstone, Ultisols

siltstone, lnceptisols

shale

221 Eastern Broadleaf Forest Province

221E dissected

plateau

sandstone, Ultisols

siltstone, lnceptisols

shale, coal

221 H dissected sandstone, Ultisols

plateau shale, coal Inceptisols

2211 Faulted/

folded

monoclinal

mountains

sandstone, Ultisols Appal. oak,

shale, Inceptisols Mixed

Ilimestone Mesoph.

231 Southeastern Mixed Forest Province

231C Faulted/

folded

monoclinal

mountains

stratified

marine

deposits

Ultisols

Inceptisols

Hemlock- N.

hardwoods

Appal. oakpine

Appalachian

hardwoods

Mixed;

mesophytic,

Appal. oak

Oak-hickorypine,

Southeastern

mixed

1000-2400 40-50

46-48

120-l 50 Fire; forestry,

oil,

agriculture

35-45

39-55

120-I 70 Fire, clearing:

agriculture,

urban

650-980 46-55 175 Fire: forestry

800-l 000 46-55 175 Fire; forestry

50-55 200-210 Fire, drought;

60-62 forestry

M221 Central Appalachian Broadleaf Forest - Coniferous Forest - Meadow Province

M221A Faulted/

folded

parallel

ridges

limestone, lnceptisols Appal. oak, 300-4800 35-50 120-l 70 Fire; agric.,

sandstone, Ultisols oak-hickory, 46-60 forestry,

shale pine urban

M221 B Severely sandstone, Inceptisols Mixed 1000-4600 40-60

dissected shale Ultisols hardwoods, 39-54

plateau spruce-fir

M221C Faulted/

folded

monoclinal

mountains

sandstone, Inceptisols Mixed 2000-2600 40-47 140-l 60 Fire;

shale Ultisols mesophytic, 45-50 agriculture,

Appal. oak forestry

11 O-1 60 Fire; forestry

-3-
Braun Historical Data

Reports by Braun (1940; 1950) include a valuable

historical record of the characteristics of the mixed

mesophytic forest during the 1940s. Field studies

conducted by Braun included sampling along

transects averaging 1 mile in length to obtain counts

of living overstory trees. Braun sampled a total of

10,140 trees from 19 counties with most trees

(7,027) located in the Cumberland Mountains of

southeastern Kentucky (see Figure 1). Published

data from the various sampled areas provide a means

to estimate the species composition of the forest

during the 1940s.

Diameters and spacing in tree stands were not

provided in the Braun reports, which precluded

estimation of how trees of different sizes were

distributed in stands. However, descriptions and

within-stand photographs indicate an uneven aged

forest with trees of various sizes. Some trees were

reported to be at least 40 inches dbh (diameter at 4.5

feet above the ground), but photographs often depict

one or two of these large diameter trees surrounded

by trees of smaller sizes.

Counts of dead trees were also not collected in the

Braun data, which prevented comparison with

estimates derived from recent FIA data used in this

study. Although Braun mentioned that American

chestnut was disappearing due to chestnut blight,

Braun did not mention the occurrence of mortality of

other tree species.

FIA Data

The Northeastern Research Station FIA Eastwide

Database (Hansen and others 1992) was used to

estimate recent species composition and percentages

of dead and removed trees, which represent cumulative amounts of mortality and cutting during a

probable period of 10 to 15 years. Data were

collected from an extensive network of randomly

sampled plots measured over a span of four years in

the following states: Alabama (1990), Kentucky

(1988), Ohio (1991), Pennsylvania (1989), Tennessee (1989), and West Virginia (1989). A few plots

in the FIA data included sample plots from Maryland (1986) and Virginia (1992).

The area of each sample plot was commonly l/5

acre, but varied among locations (l/6 to l/4 acre) as

a function of different sampling designs. Variables

used from plot records included approximate locations (latitude and longitude), forest type, stand size

(trees dominated by saplings 1 to 5 inches dbh),

poletimber (5 to 10 inches dbh), or sawtimber (>lO

inches dbh), and stand basal area (total cross-sectional area of trees at 4.5 feet above the ground).

Plot sizes were used to estimate the equivalent

number of trees per acre of each tallied tree. Variables used from each tree record included species,

diameter at breast height (dbh), crown position

(dominant, codominant, intermediate, or suppressed),

and status (live, standing or fallen dead, or cut).

Records of dead trees and cut trees from previous

inventories were used to determine diameter at breast

height and crown position values 10 to 15 years

earlier.

All tree species within the study area were of interest

for evaluation. This comprehensive approach

avoided bias towards any given species and facilitated more robust comparisons among a variety of

species groups. However, greater emphasis was

placed on species that Braun described as predominant in the overstory than on other species. Thus,

analyses were confined to FIA plots within oakhickory, northern hardwood, and oak-pine forest

types that mostly contain representative species of

the mixed mesophytic forest.

Of the selected FIA plots, 68 percent was from oakhickory types and 19 percent was from northern

hardwood forest types. Oak-hickory forest types

were predominant in all ecological subregions except

212G, which had a majority of plots in northern

hardwood forest types. Oak-pine forest types were

represented by less than 10 percent of the plots in

most subregions, and therefore some analyses were

not conducted for these types.

Analyses were also confined to FIA plots within

poletimber- and sawtimber-sized stands and to

dominant and codominant trees at least 10 inches

dbh. Trees of this size were chosen because they

represent most of the relative stocking of mature

stands and most of the removals from logging in the

region (Birch and others 1992). With this criterion, a

total of 5,404 FIA plots and 86,654 overstory trees at

-4-
least 10 inches dbh was chosen from the database

(Table 2 and Table 3). Of this total, 32 groups of

species were each represented by at least 150 trees,

with an additional 12 species placed in a miscellaneous category. Each species was commonly

represented by only one or two trees on a sample

plot, and rarely by more than five trees.

among species that vary in tolerance to competition

and different successional stages of the forest

(USDA Forest Service 1995).

Analytical Procedure

Analytical procedures to evaluate species composition, dead trees, and removed trees used percentages

of numerical counts of trees in the overstory that

were at least 10 inches dbh. Counts of trees were

used because this measure can be interpreted and

collected for later comparison with other data, and

species composition from tree counts can be compared to historical estimates by Braun (1950).

Although Braun did not include measurements of

tree diameters in reported data, it was assumed in

this study that overstory trees were greater than 10

inches in diameter and located in mature stands.

Species composition of the forest was evaluated by

combining tree data from all FIA plots within a

given forest type or ecological subregion. Percentages of trees in each species were tabulated for each

stratum. Tabulations were used to rank species by

their abundance and to determine if any differences

exist among forest types and ecological subregions.

Species composition was also evaluated at six

counties in southeastern Kentucky (counties of Bell,

Clay, Harlan, Letcher, Perry, and Whitley) that

correspond to a primary area sampled by Braun. A

combined total of 3,618 trees from 206 FIA plots in

these counties was used to determine percentages of

each species and compared to those reported by

Braun. Data from FIA plots were then used to

estimate the composition of the forest in the same

counties existing in 1988 and to interpret differences

between surveys as temporal changes.

Species were analyzed individually and in groups Proportions of dead trees and removed trees were

based on whether they were intolerant, moderately analyzed separately using FIA data. Percentages of

tolerant, or tolerant of competition (Burns and all live, dead, and removed trees were used to

Honkala 1990a; 1990b). The tolerance ratings of express the amount of dead and removed trees

species generally correspond to the successional among species in a given stratum. Expressing the

stage in which they predominate, where tolerant amount of dead and removed trees for all species in

species characterize a late successional forest. A each stratum allowed comparison of averages

previous study showed differences in stocking (higher or lower) among species.

Table 2

Distribution of FIA Sample Plots with Trees at least IO inches DBH

(by Forest Type and Ecological Subregion)

Forest Type All

Ecological Subregion

212G 221 E 221H 2211 M221A M221 B M221 C 231C

__________________--____________________------- Percent of p/o& ________________________________________------------.

Oak-Hickory 68.1 30.3 72.4 75.6 82.7 72.5 63.4 89.8 49.3

Northern

Hardwood 18.5 62.6 16.5 2.2 5.5 12.8 31.9 8.1 0.0

Oak-Pine 5.1 1.1 2.7 11.2 7.2 8.3 0.6 0.8 23.5

Mixed 91.7 93.0 91.6 89.0 95.4 93.6 95.9 98.7 72.8

Mesophytic

Types n= 5404 527 1974 a44 226 204 832 521 276

Softwoocl 8.3 6.1 8.4 11.0 4.6 6.4 4.1 1.3 27.2

and

Other Types n = 490 34 181 104 11 1 4 36 7 103

All Types n= 5894 561 2155 948 237 218 868 528 379

-5
Table 3

Distribution of all trees (live, dead, and cut) at least 10 inches DBH

from 5404 FIA sample plots located in

oak-hickory, northern hardwood, and oak-pine forest types

Species Trees

n=

Number of trees per sample plot Plots with at least:

0 1 1-2 1 3-5 1 6-10 1 1 1 + 1 tree 3 trees

Percent of 5404 plots n= n =

Ash sp. 2157 80.7 15.2 2.9 1.0 0.2 1044 221

Aspen sp. 594 95.4 3.0 1.2 0.2 0.1 246 8 2

Basswood sp. 1457 89.5 6.9 2.6 0.8 0.2 568 193

Beech 4446 77.2 12.2 5.8 3.7 1.1 1233 575

Birch sp. 1791 86.7 9.4 2.7 1.0 0.2 717 211

Black cherry 5240 77.4 12.0 5.2 3.2 2.2 1221 571

Black locust 1350 88.9 8.3 2.0 0.7 0.1 601 153

Black walnut 617 93.0 6.0 0.9 0.1 0.0 380 5 5

Blackgum 791 89.2 9.9 0.8 0.1 0.0 581 4 5

Buckeye sp. 280 97.2 2.3 0.4 0.1 0.0 149 2 5

Cucumbertree 786 91.8 6.8 1.1 0.2 0.0 441 7 3

Elm sp. 869 91.9 6.3 1.5 0.3 0.0 440 100

Hemlock sp. 923 93.5 4.0 1.8 0.6 0.1 350 133

Hickory sp. 5208 60.7 26.5 9.7 2.7 0.4 2124 692

Magnolia sp. 158 98.3 1.5 0.2 0.1 0.0 9 4 1 5

Maple, red 8047 58.2 24.7 10.1 4.1 2.9 2259 925

Maple, sugar 4624 73.1 18.7 5.9 2.9 1.3 1452 547

Oak, black 5236 65.9 21.0 8.9 3.6 0.7 1845 711

Oak, chestnut 8114 65.2 15.1 10.4 6.6 2.7 1883 1066

Oak, northern red 7626 56.2 25.9 11.4 4.6 1.9 2367 969

Oak, other red 555 95.6 3.3 0.9 0.2 0.1 239 6 0

Oak, scarlet 3277 77.6 14.5 5.4 2.1 0.4 1210 429

Oak, other white 429 95.7 3.4 0.8 0.1 0.0 233 4 7

Oak, white 8069 56.4 23.2 12.6 5.8 1.9 2356 1100

Pine, loblolly 488 97.7 1.0 0.7 0.4 0.1 124 6 8

Pine, pitch 281 97.2 2.2 0.5 0.0 0.0 149 3 2

Pine, shortleaf 598 95.4 3.3 0.9 0.3 0.1 247 7 0

Pine, Virginia 1171 92.3 4.9 1.8 0.8 0.2 417 152

Pine, white 433 97.1 1.9 0.6 0.4 0.1 157 5 5

Sassafras 463 95.4 3.9 0.5 0.1 0.1 249 3 8

Sweetgum 263 98.0 1.3 0.5 0.1 0.0 107 3 5

Yellow-poplar 9280 58.0 20.2 11.8 6.8 3.1 2267 1174

Other species 1075 87.7 2.1 0.3 0.0 0.0 663 1 6

All Species 86654 0.0 1.6 5.7 i. .,k&!“’ 74.8 5404 5319

-6-
Statistical tests of independence were used to

compare percentages among species and ecological

strata to determine if differences were significant

(p < 0.05) (Agresti 1990). In this procedure, the

Chi-square test statistic was used to determine if

there is significant difference in percentage of dead

or removed trees in a given stratum and the percentage of dead or removed trees in all observations.

To compare percentages of dead trees in sampled

FIA plots at different locations, only plots with at

least three live, dead and/or removed trees of a

subject species were used. Percentages of plots with

dead trees present or absent among forest types and

ecological subregions were

compared.

In the 194Os, Braun (1950) reported that 84 percent

of the overstory trees in the Cumberland Mountains

of southeastern Kentucky consisted of 10 species

(see Figure 3 and Appendix Table B-l). Data

collected from FIA plots in the same counties during

To determine if dead trees

and removed trees on the

same plot were associated,

each plot was designated as

having removals present or

absent depending on evidence

of at least one removed tree

of any species. To facilitate

statistical tests of association,

analyses were confined to

species represented on at least

50 sample plots. The percentages of plots with dead trees

of individual species were

determined for the groups of

plots with and without

removals, and tests of independence were used to

determine if percentages were

significantly different (p c

0.05) for groups that were

compared.

Intolerant Species

yellow-poplar

hickory sp.

scarlet oak

black locust

pine sp.

ash sp.

sweet birch

black walnut

black cherry

Total

n 1988: 38%

•l 1940s: 18%

I 0 5 10 15 20

percent of trees

Moderately Tolerant Species

chestnut oak

white oak

black oak

north. red oak

cucumbertree Total

chestnut n 1988: 37%

elm sp.

0 5 10 1 5 20

percent of trees

Tolerant Species

Maps of plot locations and

their corresponding attributes

were also used to show where

dead trees were present or

absent for individual tree

species. Spatial distributions

of plots containing dead trees

were visually interpreted to

determine if plots occurred in

groups or if they appeared to

be randomly scattered.

red maple

beech

sugar maple

basswood sp.

blackaum

hemlock sp.

yellow buckeye

m

23%

53%

5 10 15 20

percent of trees

Results and Discussion

1. Differences in Species

Composition in the Last 50 Years

Historical Changes in Southeastern

Kentucky
1988 showed that a different set of 10 species

accounted for 83 percent of the overstory. Oak,

hickory, red maple, and yellow-poplar were more

abundant; beech, sugar maple, hemlock, basswood,

and buckeye were less frequent. American chestnut

was notably absent.

Braun estimated that the American chestnut comprised 10 percent of the overstory and observed that

trees were dying from chestnut blight. The eventual

loss of chestnut from the overstory is a well-known

event. However, corresponding decreases in American

beech, sugar maple, eastern

hemlock, and yellow buckeye

did not coincide with any

disease or insect pest outbreak

(e.g., beech bark disease

[Nectria coccinea var.

faginata (Pers.:Fr.)] or hemlock woolly adelgid [Adelges

tsugae Annand]). Because

these species are tolerant of

limited growing conditions

and therefore predominate

late successional forests, it is

not likely that they were outcompeted by other species. It

is more likely that disturbances such as logging and

land clearing created forest

openings that were rapidly

colonized by early successional species well adapted to

increased light, moisture, and

nutrient availability.

FIA data show that early

successional species, including yellow-poplar, scarlet oak,

hickory, black locust, Virginia

pine, and red maple, were

more abundant in 1988. The

increase of red maple also

corresponds to the capability

of this species to easily

regenerate and out-compete

other species on a range of

hydric to xeric sites (Burns

and Honkala 1990a; 1990b).

FIA Data: Composition of Mixed

Mesophytic Forest

Two-thirds of all FIA plots measured throughout the

mixed mesophytic forest between 1986 and 1992

were within oak-hickory forest types, while only 19

percent of the plots represented northern hardwood

types (see Table 1). Figure 4 (derived from Appendix Table C-l) shows that the oak-hickory types

throughout the forest had a composition similar to

southeastern Kentucky in 1988 (see Figure 3).

Figure 4. Species comp&ition of oak-hickory forest types by tolerance to

competition (FIA data).

Oak-hickory Forest Types

Intolerant Species

yellow-poplar

hickory sp.

scarlet oak

black locust

pine sp.

ash sp.

sweet birch

black walnut Total: 38%

black cherry

0 5 10 1 5 20

percent of trees> 10” dbh

Moderately Tolerant Species

chestnut oak

white oak

black oak

north. red oak

cucumbertree

chestnut (Absent in FIA Survey) Total: 44%

elm sp. f

0 5 10 15 2 0

percent of trees > 10” dbh

Tolerant Species

red maple

beech

sugar maple

basswood sp.

blackgum

hemlock sp. Total: 16%

yellow buckeye I

0 5

percent of 2:s > 10” dbh

1 5 2 0

-8-
Conversely, Figure 5 shows that the composition of

northern hardwood forest types more closely resembled the composition of southeastern Kentucky

in the 1940s. These results suggest that the mixed

mesophytic forest region in the late 1980s were at an

earlier stage of succession than that observed by

Braun in the 1940s.

By definition, the oak-hickory forest types contain

fewer tree species tolerant of competition than the

northern hardwood types. FIA data show that only

16 percent of the trees in the oak-hickory type were

of tolerant species as compared to 49 percent of the

trees in the northern hardwood type. Most of this

difference was due to greater percentages of red

maple, sugar maple, American beech, basswood, and

hemlock in the northern hardwood type. Conversely, high percentages of oak species in the oakhickory type explain why 44 percent of the trees are

moderately tolerant of competition, while only 9

percent of the trees in the northern hardwood type

were moderately tolerant. The two groups of forest

types are similar in that at least 40 percent of their

trees were intolerant of competition. However,

intolerant species in the oak-hickory type were

mostly yellow-poplar, scarlet oak, and hickory

Northern Hardwood Forest Types

Intolerant Species

yellow-poplar

hickory sp.

scarlet oak

black locust

pine sp.

ash sp.

sweet birch

black walnut Total: 41%

black cherry I

0 5 10

percent of trees> 10” dbh

15 20

Moderately Tolerant Species

chestnut oak

white oak

black oak

north. red oak

cucumbertree

chestnut (Absent in FIA Survey) Total: 9%

elm sp. 1

0 5

percent of trze > 10” dbh

15 20

Tolerant Species

red maple

beech

sugar maple

basswood sp.

blackgum

hemlock sp. Total: 49%

yellow buckeye -I

0 5 10 15 20

percent of trees > 10” dbh

Figure 5. Species composition of northern hardwood forest types by

tolerance to competition (FIA data).

species, and those in the

northern hardwood type are

mostly black cherry, sweet

birch, and ash species.

Various pine and other oak

species were not common and

mostly found in the minor

component of oak-pine forest

types (see Appendix Table Cl). Pine species comprised

about 40 percent of the oakpine forest and consisted of

Virginia pine, shortleaf pine,

loblolly pine, eastern white

pine, and pitch pine. Only 7

percent of the trees in this type

were of species tolerant of

competition.

2. Rate of Tree

Mortality

This section of the study

ascertained whether

trees were dying at an

accelerated rate.

Data show that percentages of

dead trees in the oak-hickory

and northern hardwood types

were similar at 6 and 7 percent,

respectively. About 6 percent

of the trees in the northern

hardwood types were removed

as compared to 3 percent in the

oak-hickory types.

-9-
Records of dead trees and removed trees in the FIA

database represent cumulative amounts of mortality

and cutting that occurred during the lo- to 15year

period between measurements of sampled plots.

Analyses of data from the mixed mesophytic forest

show that 6 percent of trees with at least 10 inches

dbh were still standing or fallen dead and 4 percent

have been removed (see Appendix Table C- 1).

Mortality Rate of Different Species

Percentages of dead trees varied greatly among

species, which indicate different rates of mortality

and consequent changes in

species composition. In the

oak-hickory forest type, 7

percent of trees intolerant of

competition were dead compared to 4 percent of tolerant

species and 5 percent of

species that were moderately

tolerant (Figure 6). Species in

the northern hardwood type

that were tolerant of competition also had lower proportions

of dead trees than species that

were intolerant or moderately

tolerant (Figure 7 on next

page). These differences

suggest that most of the recent

mortality is related to the

dynamics of forest succession.

Intolerant Species

yellow-poplar

hickory sp.

scarlet oak

black locust

pine sp.

ash sp.

sweet birch Mean

black walnut I Dead: 7 %

black cherry

0 5 1 0 15 2 0 25

percent of trees> 10” dbh

Moderately Tolerant Species

to Dutch elm disease (DED) and elm yellows. DED

is caused by a fungus [Ophiostoma ulmi (Buism.)

Nannf.] and elm yellows is caused by a phytoplasm

(Hicks and Mudrick 1993).

Tree cutting also accounts for recent mortality. In

the oak-hickory type forest, loblolly pine, shortleaf

pine, and Virginia pine showed the highest percentages of removed trees (9 to 25 percent) (see Figure 6

and Appendix Table A-3). Black oak, northern red

oak, black cherry, black walnut, and sweetgum were

also being removed faster than average from the

oak-hickory forest. Within the northern hardwood

Oak-hickory Forest Types

In the oak-hickory forest type,

species with the greatest

proportions of dead trees were

elm, black locust, pitch pine,

Virginia pine, and scarlet oak.

Species with the greatest

percentages of dead trees in

the northern hardwood type

were black locust, birch,

cucumbertree, elm, hemlock,

and chestnut oak. Most of

these species were intolerant

chestnut oak

white oak

black oak

north. red oak

cucumbertree

chestnut

elm sp.

Mean

0 5 1 0 15 2 0 2 5

percent of trees > 10” dbh

Tolerant Species

red maple

beech

sugar maple

basswood sp.

blackgum Mean

hemlock sp.

yellow buckeye

of competition and representative of early stages of forest

succession. Although elm is 0 5 1 0 1 5 2 0 2 5

moderately tolerant of compe- percent of trees > 10” dbh

tition, high percentages of Figure 6. Percentages of dead frees and removed frees in oak-hickory

dead elm were most likely due forest types by tolerance to competition (F/A data).

-lO-
forest, species moderately tolerant of competition and beech. However, for all combined species

were removed more frequently than the tolerant and intolerant of competition (Figure 8), there were more

intolerant species (see Figure 7 and Appendix Table

C-l). For example, black oak, chestnut oak, northern red oak, and white oak accounted for most of the

13 percent of removed trees. Another 10 percent

each of moderately tolerant (cucumbertree and elm)

and intolerant species (black walnut and yellowpoplar) were also removed.

dead trees in the 10 to 15 inch class.

lo T DBH class (inches)

Rate of Mortality in Large Trees

Species in this study were represented by two sizes

of trees: (1) those that were 10 to 15 inches dbh, and

(2) trees larger than 15 inches dbh (see Appendix

Table C-2). Proportions of dead trees were significantly greater among larger trees of hickory, scarlet

oak, shortleaf pine, Virginia pine, elm, magnolia,

INT MOD TOL

Dead

INT MOD TOL

Removed

Northern Hardwood Forest Types

Intolerant Species

I

scarlet oak

black locust

pine sp.

ash sp.

sweet birch

black walnut

black cherry

n Dead: 8%

0 Removed: 6%

I

0 5 10 10Yibh 2 0 2 5

percent Of trees>

Moderately Tolerant Species

chestnut oak I white oak

T

J

black oak J

north. red oak

cucumbertree

chestnut I Dead: 9%

elm sp.

0 5 1 0 1 5 2 0 2 5

percent percent of trees > IO” dbh

rolerant Species

red maple

beech

sugar maple

basswood sp.

blackgum

hemlock sp. 4

J

yellowbuckeye :AI :

Mean

II Dead: 5%

Cl Removed: 5%

I

0 5 1 0 1 5 2 0 2 5

percent of trees > lo” dbh

Figure 7. Percentages of dead frees and removed frees in

northern hardwood forest types by tolerance to competition

(FIA data).

Figure 8. Percentages of dead trees and

removed trees by dbh class and tolerance to

competition: In tolerant (INT), moderately

tolerant (MOD), and tolerant (TOL) (FIA data).

In general, more of the larger trees (~15

inches dbh) were removed than trees that

were 10 to 15 inches dbh for most species.

These results indicate selective logging of

timber species and support previous studies

showing that changes in forest composition

and structure are partly due to a disproportionate removal of large trees of marketable

species (Birch and others 1992).

Mortality by Geographic Locations

Significantly greater percentages of dead

trees belonged to intolerant species within

oak-hickory forests (Figure 9) at the Appalachian plateau (ecological subregions 212G,

221E and 221H) than in mountainous areas

(subregions M221A, M221B, and M221C).

Specifically, at least 10 percent of the aspen,

birch, black locust, sassafras, scarlet oak, and

Virginia pine were dead when they occurred

in the plateau subregions (Appendix Tables

C-3 through C-10). Percentages of dead trees

of moderately tolerant species (mostly oak)

were also greater in the northern hardwood

forests of the Appalachian plateau than in

other subregions (Figure 10).

-ll-
Oak-hickory Forest Types Northern Hardwood Forest Types

Intolerant Species Intolerant Species

0’

212G

‘ii, 221E

? !

9

M221A

m M221B

Z

0

M221 c

$

221 H

8

221 I

w 231C

Mean

n Dead: 7%

0’

212G

*b 221E

? !

‘:

M221 A

fn M221 B

3 M221C

‘g 221H Mean

2 2211 (insuffIcient data) n Dead: 8%

w 231C

0 5

percent of t&k> lo” dbh

II 20

Moderately Tolerant Species Moderate1

percent of trees> lo” dbh

5 2120

P 221 E

9 M221 A

ul M221 B

jj M221C

9) 221H

w”‘0 2211

231C

Mean

m Dead: 5%

01 Removed: 4%4

0

Tolerant Species

5 oft& 15 20

parcent > IO” dbh

s 2120 221E

P M22lA

‘:

tn M221 B

b

M22lC

9) 221 H

2211

231C

Meen

I Dead: 4%

0 6 10 15 20

percent of trees > 10” dbh

Figure 9. Percentages of dead trees and removed

trees in oak-hickory forest types by tolerance to

competition and ecological subregion (FIA data).

Percentages of removed trees were also greater in

the northern hardwood forest types of subregions

212G, 221E, M221B, and M221C (see Figure 10).

Species with the greatest proportions of removed

trees in the mountainous subregions of M221B and

M221C were cucumbertree, red maple, oak, and

yellow-poplar. Black walnut, elm, oak, and yellowpoplar were removed more than other species in

subregion 221E, while hemlock, beech, and oak

were more frequently removed in subregion 212G.

In the oak-hickory forest, subregions 212G and

M221B also had greater percentages of removed

trees than other subregions.

Figures 11-15 on the following pages show the

locations of plots with dead trees and illustrate the

differences in mortality among ecological subregions. Throughout the region, an average of 20

s

2120

-5 221E

? !

2

M221 A

tn M2210

5

M22lC

j’ 221H

5

di

2211 (Insufficient data)

231C

0 5

percent of tr& > 10” dbh

16 20

Tolerant Species

s

212G

.6 221E

e

‘?

M22lA

tn M221 B

2

M221C

‘s) 221H Mean

2211 (lnsufflcient data) I Dead: 5%

231 C

0 5 t o 16 20

psccent of trees > 10” dbh

Figure 70. Percentages of dead trees and removed

trees in northern hardwood forest types by tolerance

to competition and ecological subregion (FIA data).

percent of the plots contained dead trees. However,

significantly more plots in subregions 212G, 221E,

and 221H contained dead trees of several species

(see Appendix Table D- 1). Subregion 2 12G contained plots with the highest number of dead trees,

where dead birch, hemlock, sugar maple, and

chestnut oak were found on at least 30 percent of the

plots containing these species. Red maple was the

only species with dead trees on less than 20 percent

of the sampled plots. Subregions 221E and 22 1H

also contained many species with dead trees on more

than 20 percent of the plots (see Appendix Table

D-l). In 221E, significantly more than 20 percent of

the plots contained dead black locust, elm, scarlet

oak, Virginia pine, hickory, black cherry, and beech.

Significantly greater percentages of plots in subregion 221H contained dead beech, hickory, black oak,

and scarlet oak.

-12-
Dead Trees greater than 10 inches DBH:

* Present ‘ Absent

Figure 7 1. F/A plots containing at least three trees greater than 70 inches cfbh.

-13-
Black Cherry 21x

7 .

EcoSoqluf Number Pmxntwllh

221E 201 27

221H 0 -

2211 3 -

2316 0 -

Y221A 9 11

M221B 132 11

M22X 3 -

All 571 2 1

V M221C 23 32

Au 73 11

Black Locust

1

Ecokglcal Number Pwcentwttt

subregkm ofPbta OeadTrem

212Q 0 39

221E 78 -

221H 3 51

2211 5 -

231c 0 -

M221A 23 26

M221B 33 27

Elm species

v

221E

221H

2211

231c

M221A

M221B

M22lC

Dead Trees greater than 10 inches DBH:

* Present * Absent

Figure 12. F/A plots containing at /east three trees greater than 70 inches dbh.

-14-
Dead Trees greater than 10 inches DBH:

l Present * Absent

Figure 13. FIA plots containing at least three trees greater than 10 inches dbh.

-15
Chestnut Oak

Dead Trees greater than 10 inches DBH:

* Present * Absent

Figure 14. FIA plots containing at least three trees greater than 10 inches dbh.

-16-
Dead Trees greater than 10 inches DBH:

g Present * Absent

Figure 15. FIA plots containing at least three trees greater than 10 inches dbh.

-17-
The maps also show that plots containing dead trees

were often in proximity to plots with no dead trees.

The interspersion of plots with and without dead

trees and lack of well-defined spatial patterning

suggests that tree mortality is not associated with

any widespread disturbance. Exceptions occur

within subregion 221E, which had more plots with

dead hickory and yellow-poplar; a similar situation

was observed in southeastern Ohio. Additionally,

the northern section of subregion M221B has a

greater frequency of plots with dead red maple,

chestnut oak, northern red oak, and white oak than

the southern section.

3. Causes of Recent Tree Mortality

Forest Succession

Most dead trees in the mixed mesophytic forest were

probably the result of competition among species

during forest succession. FIA data showed that

species with the greatest percentage of dead trees

were mostly trees that occupied forests at early

stages of succession, were intolerant of competition,

and which were not among the list of species

deemed characteristic of the forest 50 years ago

(Braun 1950). These species include aspen, sweet

birch, black locust, scarlet oak, sassafras, and

Virginia pine. Conversely, species with lower than

average percentages of dead trees were moderately

tolerant or tolerant of competition and were characteristic of the mixed mesophytic forest in a mid- to

late-successional stage. The greater proportion of

dead trees of early-successional species suggests that

the composition of the forest tends to change to one

that more closely resembles the historical forest.

Rates of forest succession may vary among geographic locations as a function of inherent differences in climate, physiography, and soils that affect

competitive interactions among tree species. These

influences are somewhat evident from differential

percentages of dead trees of some species among

ecological subregions. The co-occurrence of northern hardwood forest types and oak-hickory types

may also indicate that some forest stands have

progressed to late-successional stages sooner than

others. However, some of these stands will not

progress to a cover of late-successional species

because of restrictive site conditions (Spurr and

-18-

Barnes, 1992). Oak-hickory forest types represent

the climax stage of succession in these cases.

Logging and Land Clearing

Cutting of trees is a disturbance that has occurred

mostly on private non-industrial land where larger

trees of more marketable species are selectively

harvested (Birch and others 1992). FIA data showed

that 4 percent of all overstory trees (consisting

mostly of oak, sugar maple, black cherry, and

yellow-poplar) in the region have been cut. These

data indicate that species with the greatest proportion of removed trees are not the same as those with

the greatest percentages of dead trees. Removal of

trees therefore represents a selective disturbance

with a distinct influence on changes in species

composition. The tendency to cut early- and midsuccessional species may be accelerating forest

succession processes. This hypothesis could not be

tested with data used in this study, but results from

other studies have demonstrated recent increases in

species tolerant and moderately tolerant of competition (Abrams and Downs 1990).

Some of the mortality in the forest may also be a

consequence of poorly planned logging operations

which wound and weaken remaining trees (Nichols

and others 1994). At least one-third of the FIA plots

containing dead black cherry, chestnut oak, or

hickory species also had stumps of removed trees

(Figure 16; see also Appendix Table D-2). In

contrast, less than 20 percent of the plots that

contained dead trees of these species had no evidence of tree cutting. Dead trees of ash, red maple,

sugar maple, northern red oak, scarlet oak, and

yellow-poplar were also more frequently associated

with stumps of removed trees. Additional analyses

indicated that dead trees of several species were

more frequently associated with removed trees in

221E and M221B than in other subregions (see

Appendix Tables D-3 and D-4). Species associated

with evidence of logging included black cherry,

sugar maple, red maple, chestnut oak, white pine,

and yellow-poplar. These results indicate that a

portion of tree mortality in the mixed mesophytic

forest is related to logging injury. Even so, analyses

in this study do not offer definitive evidence because

trees on some of the FIA plots were probably cut

after neighboring trees died.
Intolerant Species

Virginia pine

black locust

scarlet oak

sweet birch

hickory sp.

ash sp.

black cherry

yellow-poplar

0 10 2 0 30 40

percent of plots with dead trees present

Moderately Tolerant Species

elm sp.

black oak

north. red oak

chestnut oak

cucumbertree

white oak

10 2 0 3 0 40

percent of plots with dead trees present

‘olerant Species

beech

red maple

hemlock sp.

sugar maple

basswood sp.

10 20 3 0 40

percent of plots with dead trees present

Evidence of Removals: II Present E3 Absent

Figure 76. Percentages of F/A plots containing dead trees of selected

species for plots with and without evidence of any removed trees.

Disturbances caused by changes in land use to nonforested conditions are also evident. Population

growth and urban encroachment have reduced some

of the forested acreage; however, the effects of land

use have been partially offset by the reversion of

abandoned farms to forest. Although the effects

caused by logging and land clearing are conspicuous,

it was beyond the scope of this study to assess how

much forestland has recently changed use.

Insects, Diseases,

Weather, and Fire

Forest insects, diseases,

weather events, and fire

regularly occur within the

mixed mesophytic forest and

are likely explanations for

some of the recent tree

mortality. Although exact

locations of past disturbances were not available,

the percentages and locations

of dead trees of some species

are indicative of several

documented disturbances

(Hicks and Mudrick 1993;

Twardus and Mielke 1995).

In addition to preventing

American chestnut from

regaining status in the

overstory, chestnut blight

also infects scarlet oak

(Torsello and others 1994).

This disease may explain

why more than 10 percent of

the scarlet oak analyzed in

this study were dead (see

Appendix Table C-l). The

particularly high percentage

of dead scarlet oak in

subregion 2211 implies that

this species will become a

smaller component of the

overstory here than in some

other areas.

In the last two decades,

gypsy moth [Lymantria

dispar L.] defoliation and

beech bark disease have caused the deaths of the oak

and beech components of the forests in western

Pennsylvania, Maryland, and northern counties of

West Virginia. High percentages of dead elm in the

forest may be attributed to Dutch elm disease and

elm yellows. Other insects, diseases, weather

events, and fires have also caused tree mortality of

other species and have played a role in shaping the

composition of the forest (Hicks and Mudrick 1993).

-19-
Air Pollution

Tree species of the mixed mesophytic forest that are

sensitive to sulfur, nitrogen and/or ozone air pollutants are: ash, white oak, yellow-poplar, black cherry,

loblolly pine, Virginia pine, eastern white pine,

birch, and aspen (Shriner and others 1990). While

this study did not attempt to quantify the relationships between tree mortality and air pollution, FIA

data show that several species sensitive to pollutants

had percentages of dead trees equal to or less than

species insensitive to pollutants. For example,

yellow-poplar, a species sensitive to air pollutants,

has significantly less dead trees than black locust,

which is said to be tolerant of pollutants. In addition, dead trees of pollutant-sensitive species did not

occur along well-defined spatial gradients. Fewer

dead trees were found in mountainous ecological

subregions, which is naturally subjected to greater

amounts of pollutants.

4. The Future Forest

Forest succession is a dynamic natural process that

will continue to cause changes in species composition of the mixed mesophytic forest as it matures.

Different disturbances to the forest are likely to recur

in the future, and just like any weather phenomenon,

their locations, timing, and magnitude are not

predictable. The effects of air pollutants on forest

trees are also still uncertain and can only be determined by continual monitoring of the forest.

Continual monitoring of the forests comes from two

sources: FIA and Forest Health Monitoring (FHM)

programs at each of the eight research stations of the

USDA Forest Service. Each Forest Service research

station is required by law to conduct FIA programs.

Northeastern Area, State and Private Forestry,

conducts the FHM program in cooperation with the

National Association of State Foresters (NASF).

FIA measures sampled plots to estimate a variety of

resource attributes. All states containing mixed

mesophytic forests are remeasured every 1 to 5

years. Analyses in this study pertaining to the

overstory of the mixed mesophytic forest could

easily be applied to new FIA data as these become

available. Historical comparisons of species composition and percentages of dead trees and removed

trees as estimated in this study can be made to

quantify changes in forest composition.

The FHM program includes measurements from

sampled plots that are similar in design to those

implemented by the FIA program (USDA Forest

Service 1997). In the mixed mesophytic forest,

FHM plots are currently measured at randomly

selected locations in Maryland, Ohio, Pennsylvania,

Virginia, and West Virginia at the time this publication was written (1998). The FHM plots are less

numerous than FIA plots, but include more measurements of forest health including conditions of tree

crowns and evidence of tree damage. The abundance and diversity of pollutant-sensitive lichens and

ozone-sensitive plants are also quantified on or near

the FHM plots. Analyses of data from these surveys

will provide more detail about the conditional status

of the mixed mesophytic forest at the turn of the

century.

Conclusions

This study showed the interplay between various

factors involved in forest succession and the disturbances associated with temporal and spatial changes

in the mixed mesophytic forest. Analyses of data

published by Braun (1950) and FIA data collected by

the Northeastern Research Station showed the

following:

cl

cl

cl

Two-thirds of all analyzed FIA plots were in the

oak-hickory type. This suggests that most of the

forest in the 1980s was in an earlier stage of

succession than that observed by Braun in the

1940s.

Six percent of all trees at least 10 inches in

diameter on FIA plots were observed as standing

or have fallen (dead), and 4 percent had been

removed.

Differential proportions of dead trees among

species correspond to expected mortality from

competition during forest succession; species

intolerant of competition had the greatest

proportions of dead trees.

-2o-
q For most species intolerant of competition,

percentages of dead trees that were 10 to 15

inches in diameter were significantly greater

than of dead trees larger than 15 inches.

0 A greater proportion of dead trees belonged to

early successional species, which suggests that

forest composition is changing to one that more

closely resembles the historical forest as described by Braun.

0 Differences in dead trees among locations were

most evident in ecological subregion 212G

covering north central Pennsylvania, where dead

sweet birch, hemlock, sugar maple, and chestnut

oak were found on at least 30 percent of the

sampled plots containing these species.

0 Dead trees of species sensitive to pollutants did

not occur along well-defined spatial gradients;

fewer dead trees were found in mountainous

ecological subregions where trees were exposed

to higher amounts of pollution.

*****

References

Abrams, M.D.; Downs, J.A. 1990. Successional

replacement of old-growth white oak by mixed

mesophytic hardwoods in southwestern

Pennsylvania. Can. J. For. Res. 20: 1864- 1870.

Abrams, M.D.; Nowacki, M.D. 1992. Historical

variation in fire, oak recruitment, and postlogging accelerated succession in central

Pennsylvania. Bull. Torr. Bot. Club. 119: 19-

28.

Agresti, A. 1990. Categorical data analysis. John

Wiley and Sons, Inc. New York, NY. 558 p.

Birch, T.W.; D.A. Gansner; S.L. Arner; R.H.

Widmann. 1992. Cutting activity on West

Virginia timberlands. North. J. Appl. For. 9:

146-148.

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

America. MacMillan Publishing Co., Inc. New

York, NY. 596 p.

Braun, E.L. 1940. An ecological transect of Black

Mountain, Kentucky. Ecological Monographs.

10: 193-241.

Burns, R.M.; Honkala, B.H., tech. coords. 1990a.

Silvics of North America: 1. Conifers.

Agriculture Handbook 654. USDA Forest

Service. Washington, D.C. Vol. 1,675 p.

Bums, R.M.; Honkala, B.H., tech. coords. 1990b.

Silvics of North America: 1. Hardwoods.

Agriculture Handbook 654. USDA Forest

Service. Washington, D.C. Vol. 2, 877 p.

Hansen, M.H., Frieswyck, T., Glover, J.F., Kelly, J.F.

1992. The Eastwide forest inventory data base:

users manual. Gen. Tech. Rep. NC-151. St.

Paul, MN: USDA Forest Service, North

Central Forest Experiment Station. 48 p.

Hicks, R.R.; Mudrick, D.A. 1993. 1993 forest health:

a status report for West Virginia. West

Virginia Department of Agriculture.

Charleston, West Virginia. 68 p.

Little, C.E. 1995. The dying of the trees: the

pandemic in America’s forests. Penguin Books

USA Inc. New York, NY. 275 p.

Martin, W.H. 1992. Characteristics of old-growth

mixed mesophytic forests. Natural Areas

Journal. 12: 127-135.

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

subregions of the United States: Section

descriptions. Administrative Publication WOWSA-5. Washington, DC: USDA Forest

Service. 267 p.

Nichols, M.T.; Lemin, R.C., Jr.; Ostrofsky, W.D.

1994. The impact of two harvesting systems on

residual stems in a partially cut stand of

northern hardwoods. Can. J. For. Res. 24:

350-357.

-21-
Shriner, D.S.; Hecks, W.W.; McLaughlin, S.B.;

Johnson, D.W.; Irving, P.W.; Joslin, J.D.;

Peterson, C.E. 1990. Response of vegetation to

atmospheric deposition and air pollution.

NAPAP SOS/T Report 18, In: Acidic

deposition: state of science and technology,

Volume 3. National Acid Precipitation

Program. Washington, DC. 206 p.

Spurr, S.H.; B.V. Barnes. 1992. Forest succession.

In: Forest ecology. Krieger Publishing

Company. Malabar, Florida: 399-420.

Torsello, M.L.; Davis, D.D.; Nash, B.L. 1994.

Incidence of Cryophonectria parasitica

cankers on scarlet oak (Quercus coccinea) in

Pennsylvania. Plant Disease. 78: 3 13-3 15.

Twardus, D.B.; Mielke, M.E., coord. eds. 1995.

Forest health highlights: northeastern states.

USDA Forest Service. Radnor, PA. 134 p.

USDA Forest Service. 1997. Forest health

monitoring 1997 Field Methods Guide. USDA

Forest Service, National Forest Health

Monitoring Program, Research Triangle Park,

NC 27709.325 p.

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forest health problems. NA-TP-07-95. Radnor,

PA: USDA Forest Service, Northeastern Forest

Experiment Station. 9 p.

-22-
Appendices

/

-23-
List of Appendices

Table A-l

Common and Scientific Names of Trees in the Overstory of the Mixed Mesophytic Forest . . . . . . . . . . . . . . . . . . . . . A-l

Table B-l

Species Composition of Different Geographic Areas as Summarized from Data by Braun (1950) . . . . . . . . . . B-l

Table C-l

Percentages of species composition (C), dead trees (D), and removed trees (R)

by forest type and shade tolerance for all combined ecological subregions. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . C-l

Table C-2

Percentages of dead trees and removed trees on FIA sample plots by diameter class. . . . . . . . . . . . . . . . . . . . . . . . . . . C-2

Table C-3

Percentages of species composition (C), dead trees (D), and removed trees (R)

by forest type and shade tolerance for ecological subregion 212G. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . C-3

Table C-4

Percentages of species composition (C), dead trees (D), and removed trees (R)

by forest type and shade tolerance for ecological subregion 221 E. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . c-4

Table C-5

Percentages of species composition (C), dead trees (D), and removed trees (R)

by forest type and shade tolerance for ecological subregion M221 A. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . C-5

Table C-6

Percentages of species composition (C), dead trees (D), and removed trees (R)

by forest type and shade tolerance for ecological subregion M221 B . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . C-6

Table C-7

Percentages of species composition (C), dead trees (D), and removed trees (R)

by forest type and shade tolerance for ecological subregion M221 C. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . c-7

Table C-8

Percentages of species composition (C), dead trees (D), and removed trees (R)

by forest type and shade tolerance for ecological subregion 221 H. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . C-8

Table C-9

Percentages of species composition (C), dead trees (D), and removed trees (R)

by forest type and shade tolerance for ecological subregion 221 L. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . c-9

Table C-l 0

Percentages of species composition (C), dead trees (D), and removed trees (R)

by forest type and shade tolerance for ecological subregion 231 C. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . c-10

Table D-l

Percentages of FIA plots with dead tree species by ecological subregion. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . D-l

Table D-2

Percentages of FIA plots with dead tree species on plots with and without tree removals. . . . . . . . . . . . . . . . . . . . . . D-2

Table D-3

Percentages of FIA plots with dead tree species on plots with and without tree removals. . . . . . . . . . . . . . . . . . . . . . D-3

Table D-4

Percentages of FIA plots with dead tree species on plots with and without tree removals. . . . . . . . . . . . . . . . . . . . . . D-4
Table A-l

Common and Scientific Names of Trees in the Overstory

of the Mixed Mesophytic Forest

Common Name Scientific Name

Ash, Green

Ash, White

Aspen, Blgtooth

Aspen, Quaking

Basswood, American

Basswood, American

Beech, American

Birch, Sweet

Birch, Yellow

Blackgum

Buckeye, Ohio

Buckeye, Yellow

Cherry, Black

Chestnut, American

Cucumbertree

Elm, American

Elm, Slippery

Hemlock, Carolina

Hemlock, Southern

Hickory, Bitternut

Hickory, Mockernut

Hickory, Pignut

Hickory, Shagbark

Hickory, Shellbark

Locust, Black

Magnolia, Fraser

Maple, Red

Maple, Sugar

Oak, Black

Oak, Chestnut

Oak, Northern Red

Oak, Scarlet

Oak, Shumard

Oak, Southern Red

Oak, White

Persimmon

Pine, Eastern White

Pine, Loblolly

Pine, Pitch

Pine, Shortleaf

Pine, Virginia

Sassafras

Sweetgum

Walnut, Black

Yellow-Poolar

Fraxinus pennsyivanica Marsh.

Fraxinus americana L.

Popuius gandidetata Michx.

Popuius tremuioides Michx.

Tiiia americana L

T/i/a heterophyiia Vent.

Fagus grand/flora Ehrh

Betuia ienta L.

Betuia aiiehaniensis Britton

Nyssa syivatica Marsh.

Aescuius giabra Wild

Aescuius octandra Marsh.

Prunus serotina Ehrh.

Castanea dentata (Marsh.) Borkh.

Magnolia acuminata L.

Uimus americana L.

Ulmus rubra Muhl.

Tsuga caroiiniana Englem.

Tsuga canadensis (L.) Carr.

Carya cordiformis (Wangenh.) K. Koch

Carya tomentosa (Poir.) Nutt.

Carya giabra (Mill.) Sweet

Carya ovata (Mill.) K. Koch.

Carya iaciniosa (Michx. f.) Loud.

Robinia psuedoacacia L.

Magnolia frasefi Walt.

Acer rubrum L.

Acer saccharum Marsh.

Quefcus veiutina Lam.

Quercus prinus L.

Quercus rubra L.

Quercus coccinea Muenchh.

Quercus shumardii Buckl.

Quercus faicata Michx.

Quercus aiba L.

Diospyros virginiana L.

Pinus strobus L.

Pinus taeda L.

Pinus rigida Mill.

Pinus echinata Mill.

Pinus virginiana Mill.

Sassafras aibidum (Nutt.) Nees

Liquidambar styracifiua L.

Jugians nigra L.

Liriodendron tuiioifera L.

A-l
Table B-l

Species Composition of Different Geographic Areas

as Summarized from Data by Braun (1950)

Species

Cumberland Appalachian Allegheny All Areas

Mountains Plateau Mountains (n=10140)

(n=7027) (rk2250) (n=663)

__-_--_-_--__________ percent of all trees _____----_--__________

Ash sp. 1.3 2.6 5.6 1.9

Birch sp. 1.4 1.0 11.5 2.2

Black cherry 0.1 0.0 2.1 0.2

Black locust 0.2 0.0 0.1 0.2

Hickory sp. 3.4 6.7 0.9 3.9

Oak scarlet 0.7 0.0 0.5

Pine sp. 1.4 0.1 1.0

Sassafras 0.1 0.0 0.0

Yellow-poplar 9.6 12.5 9.4

Walnut black 0.5 0.9 0.6

Ail lntoierants 16.5 23.7 20.3 19.6

Chestnut 9.5 1.5 13.4 6.1

Elm sp. 0.1 1.0 0.3

Magnolia sp. 1.4 2.1 5.5 1.9

Oak, black 0.4 1.3 0.5

Oak, chestnut 5.9 2.7 0.9 4.6

Oak, n. red 3.0 3.1 6.7 3.3

Oak, white 7.1 13.0 1.1 7.9

All Moderates 27.5 24.6 27.7 26.9

Basswood sp.

Beech

Blackgum

Buckeye sp.

Hemlock sp.

Maple, red

Maple, sugar

Ail Tolerants

6.3 4.0 6.5

16.9 20.4 17.7

1.9 2.6

4.6 1.5

7.6 15.5

2.5 2.6 10.7

12.2 3.9 15.0

52.2 50.6 51.9

6.0

17.7

1.9

3.7

6.7

3.2

10.6

51.6

Other species 1.6 0.9 0.1 1.5

Total 100.0 100.0 100.0 100.0

B-l
Table C-l

Percentages of species composition (C), dead trees (D), and removed trees (R)

by forest type and shade tolerance for all combined ecological subregions.

Forest Type

Species Oak-Hickory Northern Hardwood Oak-Pine

n= % c % D % R n= % c % D % R n= % C % D % R

Ash sp. 1030 1.7 5.7 3.6 2 9 0.7 ___ ___

Aspen sp. 308

Birch sp. 625

Black cherry 1174

Black locust 1054 1.9 ‘:“$q‘,, 1.7 293 ‘” 1 .7 ?&$J 4.8 3 0.1 ___ ___

Black walnut 502

jja;l I

0.9 3RII,, j,: 4.6 104 0.6 4.7 @$$g 1 1 0.4 ___ _-_

Hickory sp. 4684 ‘.Q@$ :” ‘Q-7 2.3 345 1.8 6.7 7.1 179 5.5 s,_ ‘j#j&/ 3.5 0.7

Oak, scarlet 3103 “p‘@+g-& . 4.3 1 9 0.1 ___ ___ 155 3.7 e;;s . ( 0.9

Pine, loblolly 145 0.2 0.4 @;y 0 323 _’ .?‘7:2:* 1.1 6.1

. _ Pine, pitch 207 0.4 21.7“ 4.2 2 0.0 ___ -__ 7 2 2.0 kl,? 2.3

Pine, shortleaf 274 0.6 Tn;4 4,,ij$nt 0 324

Pine, Virginia 500 1.0 iO$l 9.4 1 6 0.1 ___ -_- 655

Sassafras 422 0.8 1899 0.6 4 0 0.3 l+gjl 0.0 1 0.0 --- ___

Sweetgum 213 0.4 4.5 8.9 2 0.0 --- --- 4 8 1.4 5.3 0.0

Yellow-poplar 8378 '13.4 2.7 3.4 613 3.0 2.6 >“,‘b~“~$~~~~l 289 :$Cj 2.7 2.0

All Intolerant 22619 38.4 7.3 3.6 7ggg 41 .o ~a~“8~l’“i”“,“:i,l,r~~~. _j ,a”gp$g&,~ ? . ’ 2140 59.4 6.4 2.9

Cucumbertree 580 0.9 3.4 1.8 183 0.9 a” I”j. N/L :.,>t, :;q$~g”*“y l&

,. j~b~~~l8bW 3 0.1 ___ ___

8 ~

___

Elm sp. 549 1.0 18;6’ 2.8 308. 1.7 !.$;,&&r 9.1 /I!/8 e>- 1 2 0.4 --- ---

Maanolia S D . 122 0.2 0.8 2.5 3 2 0.2 4.2 0.0 2 0.1 -__ ___

Oak, black

Oak, chestnut

Oak, northern red

Oak, other red

Oak, other white

Oak, white

Pine, white

4966 7.2 ’ ii:tj

jjj -- .

5.1 114 0.5 7.7 __, 1%&r 156 3.6 6.8 2.9

7800 12.1 4.0 3.3 111 o . 6 j8.j .Iliiigij.r& h/j,‘ . 203 4.9’ 0.1 1.8

6782 9.5 4.5 5.2 730 3.2 6.1 . ^ ,@ 114 2.9 1.7 0.0

402 0.7 3.0 2.2 9 0.0 ___ ___ 144 3.6 5.4 0.8

315 0.5 lOi9 2.3 7 0.0 ___ ___ 107 2.8 i2.3 2.4

7451 11.7 2.8 4.0 213 1.1 7.3 g#‘i’: 405 9.2 1.5 1.5

163 0.2 6.9 4.0 6 9 0.3 8.0 6.8 201 4.0 5.9 0.0

All Intermediate

Basswood sp.

Beech

Blackgum

Buckeye sp.

Hemlock sp.

Maple, red

Maple, sugar

All Tolerant

Other Species

All Species

29130 44.1 4.6 4.1 1776 8.5 j'l 9:jvb "i3:4" 1347 31.7 4.0 1.5

759 1.2 3.6 3.6 695 3.5 2.9 2.4 3 0.1 ___ ---

2529 3.1 4.9 1.5 1888 2 ,8.6~‘ 6.9. 5.7 2 9 0.7 --- ---

712 1.1 5.1 2.2 5 1 0.2 4.3 7.6 2 8 0.7 --- ---

231 0.4 3.0 1.6 4 6 0.2 0.0 3.9 3 0.1 --- ---

341 0.5 2.6 2.5 522 2.3 11.9 7.3 6 0 1.3 2.7 0.0

4245 7.4 3.1 1.7 3648 18.5' 3.7 5.5 154 3.9 4.2 0.9

1614 2.6 3.3 2.8 2984 ’ 15.4 4.3 4.3 2 6 0.7 ___ ___

10431 16.3 3.6 2.0 9834 48.6 4.8 5.0 303 7.4 4.1 2.9

707 1.2 19.0 1.3 314 1.6 12.3 7.0 5 4 1.4 9.7 0.0

62867 100 5.6 3.5 19923 100 6.6 6.1 3844 100 5.5 2.4

Highlighted values represent species that comprise at /east 5 percent of the composition or with percentages of dead or

removed trees significantly greater (p < 0.05) than percentages for all species.

Values for dead trees or removed trees are not shown for species with less than 30 sampled trees.

C-l
Table C-2

Percentages of dead trees and removed trees on FIA sample plots

bv diameter class.

Species

INumber of

sampled trees

10-15” I >15”

Ash sp. 1231 926

Aspen sp. 499 9 5

Birch sp. 1392 399

Black cherry 2963 2277

Black locust 933 417

Black walnut 406 211

Hickory sp. 3451 1757

Oak, scarlet 1632 1645

Pine, loblolly 246 222

Pine, pitch 231 5 0

Pine, shortleaf 502 9 6

Pine, Virginia 1006 165

Sassafras 379 6 4

Sweetgum 166 9 5

Yellow-poplar 4636 4442

4.6

6.9

3.2

17.1

6.1

9.2

0.9 1.0 g{#

11.6

2.3 2.9

5.3 7.0

0.6 0.0

All Intolerant 19877 12881 8.0 5.8 3.5 5.7

Cucumbertree 423 343

Elm sp. 572 297

Magnolia sp. 113 4 3

Oak, black 2296 2940

Oak, chestnut 4231 3863

Oak, northern red 3218 4406 4.8 4.3

4.2

4.6

2.0

3.7

2.7

5.0

3.1

6.4

2.0

Oak, other red

Oak, other white

Oak, white

Pine, white

All Intermediate

Basswood

sp.

Beech

Blackgum

Buckeye sp.

Hemlock

sp.

Maple red

Maple sugar

All Tolerant

298 257 3.9 3.1 1.6 2.4

263 166 11.7 9.1 2.3 2.1

4357 3712 2.9 2.5 2.6 i’ s 732 Ij (,”

161 252 r>~lrr!Ssg~al3iiage~~~~~~l % 4.4 0.5 , ~

I ‘$3

15952 16301 5.0 4.3 3.4 6.7

768 669 3.2 3.5 1.9 “: :,;, ‘~8.4 “ ’ ’

1636 2610 5.0 ‘:~&ga:~~Q~~~., :j;, lSBIIBiBI&iB1” I 1FBBIW 3.6 3.4

387 404 4.7 6.0 2.0 : s 3.6 ‘.,

149 131 2.7 1.9 0.5 L*: 4.9‘

537 ~“iis~~~~.;bUhiiii’s,,: ; * 6.5 3.4 j; 386 : ‘7.3 :

j. ” :. ,3’ j ‘.

5215 2632 3.4 3.4 2.9 ,4,8 J

2771 1653 4.0 3.6 3.2 54‘I ‘ _

11332 9238 4.0 4.7 2.9 4.6

Other Species 675 400 16.2 18.0 2.7 3.1

All Species 47838 38818 8.2 5.1 3.3 5.8

Highlighted values for a given diameter c/ass are significantly greater (p c 0.05) than percentages for the other class.

c-2
Table C-3

Percentages of species composition (C), dead trees (D), and removed trees (R)

by forest type and shade tolerance for ecological subregion 212G.

Birch sp.

Black cherry

Black locust

Black walnut

Hickory sp.

Oak, scarlet

Pine, loblolly

Pine, pitch

Pine, shortleaf

9 0.2 ___ _-_

2 6 0.8 --_ ___ 2 9 0.4 ___ ___

125 4.0 1;

I! 4 0.0 --- -__

2 2 0.7 ___ ___ 1 0.0 --- ___

Pine, Virginia

Sassafras

Sweetgum

1 0.0 ___ ___

Elm sp.

Magnolia sp.

Oak, black

Oak, chestnut

Oak, northern red

Oak, other red

Oak, other white

1 0 0.3 --_ ___ 5 0.1 ___ ___

174 ?

324 I

fi 8.7 8.3 1 3 0.1 ___ ---

j ,iB g3 3.8 2 1 0.3 _-- ___

1100 ;

\“:

z./ li’ 4.4 7.8 157 1.7 8.8 8.5

3 0.1 -_- ___ 2 0.0 ___ -__

Basswood sp. 7 0.3 ___ -__ 121 1.6 6.5 6.1

4 3 1.5

Blackgum 2 0.1 _-- ___ 4 0.1 -_- ___

Buckeye sp.

Hemlock sp.

All Tolerant 613 20.9 3.6 3.8 4331 55.6 5.0 5.6

Other Species 8 0.2 ___ _-_ 3 4 0.5 4.0 0.0

All Species 3264 100 6.9 5.6 7921 100 6.6 5.5

Highlighted values represent species that comprise at least 5 percent of the composition or with percentages of dead

or removed trees significantly greater (p c 0.05) than percentages for all species.

Values for dead trees or removed trees are not shown for species with less than 30 sampled trees.

c-3
Table C-4

Percentages of species composition (C), dead trees (D), and removed trees (F?) by

forest type and shade tolerance for ecological subregion 221 E.

Forest Type

Species Oak-Hickory Northern Hardwood

n= 1 %C 1 % D % R n= 1 %C 1 % D % R

4.2 400 cf”;“s’g;q”‘; ‘” 7.7 5.5

. . .

Ash sp. 589 2.6 7.3

Aspen sp.

Birch sp.

Black cherry

Black locust

Black walnut

Hickory sp.

Oak, scarlet

Pine, loblolly

Pine, pitch

Pine, shortleaf

Pine, Virginia

Sassafras

Sweetgum

103 0.5 6.8

605 2.7 7.0

460 2.2 .‘~-$f# l,la

314 1.5 y’,~;“;“&:

1880 9,l 9’: 6.7

,,

1152 4.4 _’ 8, qp’-

221 1.2

247 1.3

1 5 0.1 ---

1.0 5 6 1.4 11.5 0.0

2.5 9 5 1.9 1,:.jgJ $ 0.0

5.4 1056 “: ,:,gt& 8.3 7.9

2.7 145 3.2 &.3,.‘.:’

4.1 8 7 2.0 3.6

5.5 1 0.0 -__ ___

___

5.5 1 3 0.3 --- ___

1.1 3 2 0.8 q.fir 0.0

__.

Yellow-poplar

All Intolerant

Cucumbertree

Elm

sp.

Magnolia sp. 1 0.0 ___ --- 1 0.0 --- ___

Oak, black 2410 ’ ,~g;,~;~:;; 6.7 6.3 6 7 1.0 9.8 ,:.“j&,f$ .:

” Oak, chestnut 2299 . ..a.4 ;__ 5.0 4.8 2 9 0.6 ___ ---

Oak, northern red 1896 *l* ‘, ,;6;61:. 4.8 7.0 210 3.6 9.3 : 2$5

Oak, other red 2 4 0.1 _-- --- 7 0.1 --- ___

Oak, other white 4 5 o,2 i :

j ‘wll:j_ 22& i . 0.0 4 0.1 --- ___

Oak, white 3575 14:$i:!.;,’ 2.6 5.5 101 1.9 10.6~ ‘-lL§:.

Pine, white 3 7 0.2 11.3 0.0 3 3 0.5 13.1 10.4

All Intermediate 10640 42.6 5.4 5.5 737 13.7 ..:‘io;!& b :;I .‘15:7 __(

Basswood sp.

Beech

Blackgum

Buckeye sp.

Hemlock sp.

Maple, red

Maple, sugar

203 0.8 5.9 1.9 127

949 3.2 5.4 1.3 350

179 0.7 0.8 3.5 1 9

125 0.5 3.4 0.0 2 1

4 4 0.2 0.0 0.0 101

1371 ’ ,I( (.& 3.3 2.3 599

696 3.0 2.4 2.8 727

2.5 5.0 2.3

1.7 :,. i4.4’ 0.0

;i;ii::~;;j~~l,~: 4.0 3.7

w$b;6;;;: .‘, :‘ i: 3.0 3.5

All Tolerant 3567 14.7 3.6 2.1 1944 36.9 5.5 3.6

Other SDecles 360 1.7 13.1 1.6 9 3 1.6 19.6 2.4

All Species 23936 100 6.5 4.2 5077 100 6.2 6.6

Highlighted values represent species that comprise at /east 5 percent of the composition or with percentages of dead or

removed trees significantly greater (p < 0.05) than percentages for all species.

Values for dead trees or removed trees are not shown for species with less than 30 sampled trees.

c-4
Table C-5

Percentages of species composition (C), dead trees (D), and removed trees (R) by

forest type and shade tolerance for ecological subregion M221 A.

Species Oak-Hickory Northern Hardwood

n= % c % D % R n= 1 %C 1 % D % R

Ash sp. 3 7 1.7 0.0 6.6 6 3 ~~:~~~~~~~~~~,~*~‘~~ ’ &@PW81 -it*1 1.4 3.0

Asoen SD. 1 0.0 --- ___

.

Birch sp.

Black cherry

Black locust

6 2 2.7 2.5 0.0 0.0

3 7 1.3 0.0 0.0 5.3

166 I>~“::a:r~~~~jl:~~~e; ~8, . ;.t 1 / 8.6 0.0 8.5

Black walnut

Hickory sp.

Oak, scarlet

Pine, loblolly

Pine, pitch

Pine, shortleaf

2 1 1.0 0.0 0.0 6 1.1 ___ --_

149 %?Iar.L j ,$/iB’/ 4.9 3.3 1 9 3.8 ___ ___

9 5 4.1 7.6 4.0

2 8 1.1 8.2 1.7

Pine, Virginia 1 3 0.6 --- ___ 1 0.4 ___ ___

Sassafras

Sweetgum

Yellow-poplar

8 0.4 --- _-.

171 ‘1:;. ii 71!!L’:“, 0.0 0.0 9 1.6 ___ --_

Cucumbertree

Elm sp.

Magnolia sp.

Oak, black

Oak, chestnut

Oak, northern red

Oak, other red

Oak, other white

Oak, white

Pine, white

All Intermediate

Basswood sp.

Beech

Blackgum

Buckeye sp.

Hemlock sp.

Maple, red 131 ; (> 5.9~ ) rj 2.1 0.0 4 3 0.0

Maple, sugar 6 3 2.5 0.0 0.0 109 4.2

All Tolerant 328 13.3 4.8 0.0 255 49.7 7.8 2.3

Other Species 1 9 0.8 --- ___ 2 3 4.8 8.1 0.0

All Species 2557 100 3.0 1.2 519 100 9.0 3.8

Highlighted values represent species that comprise at least 5 percent of the composition or with percentages of dead or

removed trees significantly greater (p e 0.05) than percentages for all species.

Values for dead trees or removed trees are not shown for species with less than 30 sampled trees.

c-5
Table C-6

Percentages of species composition (C), dead trees (D), and removed trees (R) by

forest type and shade tolerance for ecological subregion M221 B.

Species

Ash sp.

Aspen sp.

Birch sp.

Black cherry

Black locust

n=

101

2 0

233

Forest Type

Oak-Hickory Northern Hardwood

% c 1 %D 1 % R n= 1 %C ( % D % R

1.0 4.9 5.8 182 3.5 5.8 5.4

0.3 __- ___ 1 4 0.3 _-- ___

Black walnut

Hickory sp.

Oak, scarlet

Pine, loblolly

Pine, pitch

Pine, shortleaf

293 2.8 5.1 4.6 2 0.0 T-v ___

1 4 0.1 ___ ___

Pine, Virginia 8 0.1 ___ --_

Sassafras 7 2 @ 0.0 4 0.1 --- ---

Sweetgum 1 0.0 ___ ___

Elm sp.

Magnolia sp.

Oak, black

Oak, chestnut

Oak, northern red

Oak, other red

Oak, other white

Oak, white

Pine, white

1 4 0.1 ___ ___ 3 3 0.7 5.1 0.0

5 8 0.6 0.0 0.0 2 7 0.5 --- ___

297 2.8 7.2 4.1 2 2 0.3 --_ ---

6.0 4.2 4 0

4.4 5.5 284

_-- ___

1 4 0.1 ___ ___

Blackgum 9 7 0.8 2.8 3.8 1 7 0.3 --- ___

Buckeye sp. 9 0.1 ___ ___ 9 0.1 --_ ___

Hemlock SD. 8 4 0.7 1.2 4.3 175 2.9 4.0 6.4

Maple, red 1238 ~~~~~~~~~~~~ v n”.‘a 1 ‘%il 2.4 1 .3 752 1.8 8.4 . . . Maple, sugar 296 3.0 0.7 3.5 727 4.9 4.1

All Tolerant 2206 22.1 2.3 2.5 2574 46.2 4.5 5.9

Other Species 8 1 0.9 9.7 0.0 149 2.6 9.1 14.0

All Species 10124 100 4.7 3.7 5239 100 5.5 6.5

Highlighted values represent species that comprise at least 5 percent of the composition or with percentages of dead or

removed trees significantly greater (p c 0.05) than percentages for all species.

Values for dead trees or removed trees are not shown for species with less than 30 sampled trees.

C-6
Table C-7

Percentages of species composition (C), dead trees (D), and removed trees (R) by

forest type and shade tolerance for ecological subregion M221 C.

Species

Forest Type

Oak-Hickory Northern Hardwood

n= % c % D % R n= % c % D 1 %R

Ash sp.

Aspen sp.

Birch sp.

Black cherry

Black locust

Black walnut

Hickory sp.

Oak, scarlet

Pine, loblolly

Pine, pitch

Pine, shortleaf

Pine, Virginia

Sassafras

Sweetgum

7 3 1.1 1.3 1.1 3 0 3.5 6.5 0.0

1 0.0 --_ ___

107 1.7 6.2 0.0 1 6 2.9 ___ ___

2 5 0.4 ___ ___ 9 1.1 --_ --_

0.0 1 1 1.7 0.0 0.0

6.2 3 --_ -_-

1.6 3 7 p *&liI 9.2 0.0

4.1 2 --_ ___

1 8 0.3 ___ ___

6 0.1 ___ -__

1 5 0.3

5 8 0.9 1 0.0 3 0.6 ___ ___

1 7 0.4 ___ ___

Cucumbertree

Elm sp.

Magnolia sp.

Oak, black

Oak, chestnut

Oak, northern red

Oak, other red

Oak, other white

Oak, white

Pine. white

169 2.4 1.5 1.4 1 8 2.3 ___ ___

1 6 0.2 ___ ___ 1 0.3 -_- ___

2 5 0.5 ___ ___ 4 0.9 _-_ ___

498 5.0 4.3 6 0.8 ___ __-

965 1.6 1.7 1 2 2.0 ___ _--

562 3.0 1.3 2 5 3.3 ___ __-

8 0.1 ___ ___

4 ___ ___ 2 0.4 ___ -_-

371 2.5 1.3 8 1.7 ___ ___

Beech

Blackgum 103 1.4 2.2 1.1 7 0.8 ___ -_-

Buckeye sp. 4 2 0.7 0.0 0.0 7 0.8 _-- ___

Hemlock sp. 6 5 0.9 6.7 1.0 1 7 2.5 ___ --_

All Tolerant 1432 20.1 3.0 0.5 432 60.9 2.7 3.6

Other Species 5 5 1.0 18.4 0.0 6 2.4 ___ -_-

All SDecies 6967 100 3.6 2.1 663 100 3.8 5.0

Highlighted values represent species that comprise at /east 5 percent of the composition or with percentages of dead or

removed trees significantly greater (p e 0.05) than percentages for all species.

Values for dead trees or removed trees are not shown for species with less than 30 sampled trees.

c-7
Table C-8

Percentages of species composition (C), dead trees (D), and removed trees (I?) by

forest type and shade tolerance for ecological subregion 221 H.

I Forest Type

Species Oak-Hickory Oak-Pine

n= % c % D % R n= % c % D % R

Ash sp.

Aspen sp.

Birch sp.

Black cherry

Black locust

Black walnut

Hickory sp.

Oak, scarlet

Pine, loblolly

Pine, pitch

Pine, shortleaf

Pine, Virginia

Sassafras

109 1.1 2.4 2.7

4 0.1 --- --_

2 7 0.3 --- ---

1 7 0.1 --- ---

_._

4 5 0.4

167 2.0

152 1.8

24 0.3 --- ---

3

3

1

4

5 8

8 0

4 5

2 5

227

253

0.2 --- --_

Cucumbertree 64 0.5 0.0 0.0 1 0.1 --- ---

Elm sp.

Magnolia sp.

Gak, black

Oak, chestnut

Oak, northern red

Oak, other red

Oak, other white

Oak, white

110 1.1 5.8

---

3.4

3.3

3.0

6.1

3.1

2.8

4 0.4 ---

---

1 0.2 ---

59 4.0 9.0

70 4.2 0.3

28 2.1 ---

-__

0.0

3.0

___

0.0

3.0

0.9

Pine, white

All Intermediate

Basswood sp.

Beech

Blackgum

Buckeye sp.

Hemlock sp.

Maple, red

Maple, sugar

All Tolerant

5 4 0.4 5.4 7.8 3 1 2.2 0.0 0.0

5165 46.3 4.9 3.2 451 30.8 5.6 1.1

101 0.9 7.5 3.4

557 3.5 7.1 1.4 4 0.1 --- -_-

211 1.9 7.7 0.7 1 7 1.2 --- ___

2 9 0.3 --- ---

8 3 0.6 0.4 1.1

,; ; , 1’.( .:;,:: , I pi

464 p~::~#~~;~$~ 4.4 1.8 :; ~~~~~~~~~~ ;y5 ;yo

214 1.9 5.3 2.5 4 0.3 --- _--

1659 13.8 5.7 1.8 115 7.4 6.8 0.0

Other Species 125 1.3 43.6 2.7 15 1.2 --- _--

All Species 10805 100 s)a;yj$%b “*:I *.“$; .ii@&T&~,i 2.9 1359 100 ~~~~~~~~~~~~~ 1 .g F“ ( r:t:, .;

Highlighted values represent species that comprise at least 5 percent of the composition or with percentages

of dead or removed trees significantly greater (p c 0.05) than percentages for all species.

Values for dead trees or removed trees are not shown for species with less than 30 sampled trees.

C-8
Table C-9

Percentages of species composition (C), dead trees (D), and removed trees (R) by

forest type and shade tolerance for ecological subregion 221 L.

Species

Forest Type

Oak-Hickory Oak-Pine

n= % c % D % R n= % c % D % R

Ash sp.

Aspen sp.

Birch sp.

Black cherry

Black locust

Black walnut

Hickory sp.

Oak, scarlet

Pine, loblolly

Pine, pitch

Pine, shortleaf

Pine, Virginia

Sassafras

Sweetgum

46 1.7 4.7 1.2 2 1.4

23 1 .O --- ---

25 1.1 --- _-_

48 2.6 9.7 1 .o

1 0.0 -_- _-_

1 7 0.8 --- ---

2 0.9

1 0.4

---

_-_

_--

---

-_-

-_-

10.4

---

_--

_--

---

-_-

---

--_

0.0

--_

Yellow-poplar 51 6 ‘:,‘.’ l$$,i( 2.0 3.6 1 8 ;:; j gyp::; ___ ---

All Intolerant 1092 43.7 5.8 3.3 139 71.1 8.3 2.3

Cucumber-tree

Elm sp.

Maanolia S D .

21 0.8 --- ---

1 6 0.5 --- ---

1 5 0.8 --- ---

Oak, black 192 ,~:gj;@~:lr;, 4.2 2.0 12 I;:(; I’;&$;;;’ ___ ---

Oak, chestnut 446 : “‘j$&:. j

.+;:

2.7 0.4 5 2.2 --- -_-

Oak, northern red 202 6!6 ~2 4.8 2.0 12 ~-i;;&::. --- --_

Oak, other red 36 1.1 2.3 1.8 3 1.4 --- ---

Oak, other white 1 2 0.7 --- _-- 4 1.9 --- _--

Oak, white 236 8:2 2.6 2.6 28 “ .&#: ___ -_-

Pine, white 4 0.1 --_ -__ 1 0.1 --- ---

All Intermediate 1182 38.9 3.8 1.5 65 23.9 8.7 0.0

Basswood sp. 43 1.4 2.1 ‘:‘jg;#;;j; 1 :

Beech 104 2.8 6.3 0.0 2 0.6 --- ---

Blackgum 63 1.9 ’ 164 0 3.3 1 0.2 --- _--

Buckeye sp. 1 4 0.5 --- _-_

Hemlock sp. 24 0.9 --- __-

Maple, red 138 5.2 4.8 0.0 4 1.9 --- ---

Maple, sugar 9 8 3.4 2.3 5.1 1 0.5 --- -_-

All Tolerant 484 18.1 5.3 3.5 8 3.2 --- ---

Other Species 38 1.4 26.7 0.0 3 1.9 --- ---

All Species 2798 100 5.2 2.8 215 100 8.0 1.6

Highlighted values represent species that comprise at /east 5 percent of the composition or with percentages of dead or

removed trees significantly greater (p c 0.05) than percentages for all species.

Values for dead trees or removed trees are not shown for species with less than 30 sampled trees.

c-9
Table C-IO

Percentages of species composition (C), dead trees (D), and removed trees (R) by

forest type and shade tolerance for ecological subregion 231C.

Species Oak-Hickory

Forest Type

Oak-Pine

n= % c 1 %D ( % R n= % c % D % R

Ash sp.

Aspen sp.

Birch sp.

Black cherry

Black locust

Black walnut

Hickory sp.

Oak, scarlet

Pine, loblolly

Pine, pitch

Pine, shortleaf

Pine, Virginia

Sassafras

Sweetgum

Yellow-poplar

3 0

7

4

327

120

141

6 2

5 3

1.2 1.9 0.0 7 0.7 --- -__

5 0.6 --- -__

0.4 --- ___ 3 0.3 ___ ___

0.1 --_ ___ 1 0.2 --_ -__

1.8

0.0

3.6

a)

3.6 B 6.6

2.7 1

?

1 5.9

107

187

Cucumbertree 1 0.0 --- ___

Elm sp. 1 7 0.7 --- ___ 5 0.5 ___ -__

Magnolia sp. 1 0.0 --- ___ 1 0.1 ___ -_-

Oak, black 133 1 5.5 4.3 3 0 2.7 3.3 3.4

Oak, chestnut 401 1 0.5 0.8 6 8 ~~~*~~~~~~~~~~~ 0.0 0.0

Oak, northern .’ red 8 4 3.2 0.8 2.5 2 2 2.1 ___ ---

Oak, other red 0.6 1.8

Oak, other white 98 4.0 4.1 2.7 4 5 4.1 3.4 1.9

Oak, white

Pine, white

1.7

All Intermediate

Basswood sp.

Beech

Blackgum

Buckeye sp.

Hemlock sp.

Maple, red

Maple, sugar

All Tolerant

1237 49.7 1.9 1.7 343 31.5 2.3 1.4

2 0.1 --_ ___ 1 0.1 ___ ___

4 7 1.1 0.0 7.4 5 0.4 --- ---

4 3 2.0 0.7 3.2 7 0.8 ___ __-

1 0.2 ___ ___

4 3 1.7 5.5 1.5 9 1.0 ___ __-

7 0.3 --- ___

142 5.2 2.0 3.3 2 3 2.4 --- ___

Other Species 2 1 0.8 --- ___ 14 1.5 ___ -__

All Species 2438 100 3.1 4.3 1037 100 3.4 2.7

Highlighted values represent species that comprise at /east 5 percent of the composition or with percentages of dead or

removed trees significantly greater (p c 0.05) than percentages for all species.

Values for dead trees or removed trees are not shown for species with less than 30 sampled trees.

c-10
Table D-l

Percentages of FIA plots with dead tree species by ecological subregion.

Species

All 212G

n = % n = %

221 E

n = %

Ecological Subregion

221H 221 I

n= % n = %

231 C M221 A M221 B M221 c

n = % n = % n = % n= %

ash sp.

basswood sp.

beech

birch sp.

black cherry

black locust

cucumber-tree

u

1, elm sp.

hemlock sp.

hickory sp.

maple, red

maple, sugar

oak, black

oak, chestnut

oak, northern red

oak, scarlet

oak, white

pine, Virginia

221 23.1

193 10.9

575 24.5

211

571 21.2

153

73 19.2

100

133 24.8

692 23.1

925 15.7

547

711

###

969 19.5

429

###I 12.6

152

54 27.8

19

86 23.3

75

193 24.4

0

a

2

35

7

272 18.8

137

27 29.6

44

130 26.9

17

70 25.7

0

13

105 25.7

50 10.0

175

27

201

78

1 2

88

21 33.3

279

255 15.7

185 14.1

352 25.0

308 22.7

301 21.3

145

499 12.8

60 ^-"" "

481 13.3

a

14

80

1

0

3

5

1

16

145

65 15.4

28 21.4

3

7

la

2

3

5

1

2

3

217 14.8

75 25.3

126

237 13.9

52 28.9

169 6.5

1 1

ia

27 11.1

60 18.3

33 15.2

1 5

38 21.1

11

71 4.2

2 1 2

0 a

7 11

1 11

0 9

0 23 26.1

0 6

1 0

0 5

59 a.5 22 13.6

5 19

1 17

1 9 22 13.6

70 1.4 71 9.9

9 59 11.9

14 13

62 3.2 43 0.0

23 30.4 3

30 3.3 20 0.0

33 18.2

52 9.6

128 21.1

79 27.9

162 11.1

33 27.3

1 8

5

39 la.0

65 21.5

262 13.4

122 10.7

39 20.5

165 21.8

281 17.8

42 14.3

105 10.5

1

192 7.8

4

43 7.0

70 18.6

15

3

11

23 31.5

1

14

82 14.6

36 8.3

39 la.0

71 16.9

131 13.7

a1 11.1

57 21.1

46 6.5

2

yellow-poplar ### 9.4 198 6.6

Average ### 19.6 63 27.3 191 21.8 73 22.6 19 16.0 16 6.5 20 10.0 96 15.7 49 12.8

Highlighted values represent species with dead trees found on a percentage of plots significantly greater (p < 0.05) than the overall average of 19.6 percent.

Percentages are only shown for species represented by at /east 20 sample plots within a given forest type.
Table D-2

Percentages of FIA plots with dead tree species on plots with and without tree removals.

Species

Ash sp.

Basswood sp.

Beech

Birch sp.

Black cherry

Black locust

Cucumbertree

u

63 Elm sp.

Hemlock

Hickory sp.

Maple, red

Maple, sugar

Oak, black

Oak, chestnut

Oak, northern red

Oak, scarlet

Oak, white

Pine, Virginia

Yellow-poplar

All Types Northern Hardwood Oak-Hickory Oak-Pine

With Without With Without With Without With Without

Removals Removals Removals Removals Removals Removals Removals Removals

n= % n= % n= y0 n= % n= y0 n= y0 n= % n= %

4 9 30.6 172 20.9 29 ~~~~~~~~ 1o7 ~~~~~~~~-~,~ 2.

s+“‘-:<sf * =I 8’ 5.0 64 18.8 0 1 __ “.,ii

2 2 9.1 171 11.1 12 8 9 10.1 10

111 30.6 464 23.1 5 6 30.4 189 23.8 54

3 1 35.5 180 31.1 2 6 30.8 125 35.2 5

2 1 52.4 132 37.1

1 1 6 2 19.4

2 6 53.9 74 47.3

19 36.8 114 22.8

5 2 9 37.9 16

3 14 8

12 2 5 36.0 14

14 6 4 31.3 5

8 2 4 16.7 109

8 2 12.2

29.6 273 22.7

5 4 20.4

17.9 111 14.4

103 36.9

4 8 14.6

4 8 5;. 1

4 3 11.6

30.3 531 22.0

128 26.6 583 24.2

169 23.7 800 18.6

5 3 43.4 376 31.1

182 15.9 918 12.0

2 5 40.0 127 33.1

6 8 119

2 8 138

3 1 12.9 60 21.7 138

0 0 4 9

9 18 169

0 1 1 1

22.6 154 10.4

27.7 558 24.2

16.6 844 12.1

4 3 44.2

0 0

1 2

0 1

0 4

0 0

0 0

0 1

0 7

2 18

2 21 9.5

1 2

3 17

1 3 3 3.0

0 12

4 13

4 5 6 8.9

14 8 3 26.5

Highlighted values represent species with dead trees found on a percentage of plots significant/y greater (p c 0.05) than the overall average ofl9.6percent.

Percentages are only shown for species represented by at least 20 sample plots within a given forest type.
Table D-3

Percentages of FIA plots with dead tree species on plots with and without tree removals.

Species

212G

With Without

Removals Removals

n= % n= %

Ecological Subregion

221 E 221 H 221 I

With Without With Without With Without

Removals Removals Removals Removals Removals Removals

n= % n= % n= % n= % n= % n= %

Ash sp.

Basswood sp.

Beech

Birch sp.

Black cherry

0 Black locust

G

Cucumber-tree

Elm sp.

Hemlock

Hickory sp.

Maple, red

Maple, sugar

Oak, black

Oak, chestnut

Oak, northern red

Oak, scarlet

Oak, white

Pine, Virginia

Yellow-poplar

12

2

2 8

19

5 0

0

3

0

10

3

6 5

3 5

6

4

2 0

3

13

0

4

4 2

17

28.6 58

5 6

34.0 143

0

5

2

2 5

4

26.2 207

34.3 102

2 1

4 0

30.0 110

14

5 7

0

9

23.8

20.7

39.3

21.0

36.0

16.4

28.4

28.6

32.5

26.4

26.3

5

12

6 9

1

0

2

4

1

14

42.9 124

5 5

2 2

40.9 132

21.9 185

6 4

114

11.8 203

4 5

13.6 147

0

2

33.3 3

0

1

0

0

0

0

26.6 7

16.4 2

18.2 3

29.6 4

13.5 4

25.0 5

45.6 1

14.3 7

28.9 3

5.4 12

3

5

15

2

2

5

1

2

3

2 6 26.9

9

15

23 13.0

5 6 17.9

2 8 14.3

14

3 1 19.4

8

5 9 5.1

2 4 29.2 8 1 24.7 3

6 4 4 11.4 2

3 4 29.4 141 26.2 11

3 2 4 29.2 0

46 0

14 6 4 51.6 1

1 1 1 1

2 6 53.9 62 45.2 0

1 2 0 35.0 2

6 2 25.8 217 25.8 21

5 0 205 10

4 3 142 6

7 4 27.0 278 24.5 22

6 0 248 3 2

7 4 25.7 227 19.8 11

2 3 39.1 122 30.3 12

Average 15 30.3 4 8 24.3 3 8 28.7 152 20.0 1 0 24.4 6 3 21.6 3 16 14.8

Highlighted table values denote a significantly greater percentage of plots (p c 0.05) with both dead and removed trees than adjacent values for plots with dead

trees and no removed trees. Percentages are only shown for categories with at least 20 sample plots.
Species

Table D-4

Percentages of FIA plots with dead tree species on plots with and without tree removals.

Ecological Subregion

231C M221A M221B M221 c

With Without With Without With Without With Without

Removals Removals Removals Removals Removals Removals Removals Removals

n= % n= % n= % n= % n = % n = % n= % n= %

Ash sp.

Basswood sp.

Beech

Birch sp.

Black cherry

Black locust

u

A Cucumbertree

Elm sp.

Hemlock

Hickory sp.

Maple, red

Maple, sugar

Oak, black

Oak, chestnut

Oak, northern red

Oak, scarlet

Oak, white

Pine, Virginia

Yellow-poplar

0

0

3

0

0

0

0

0

0

1 1

1

0

4

5

2

1

1 0

6

6

2

0

4

1

0

0

0

1

0

48

4

1

15

65

7

1 3

52

17

24

2

1

0

0

1

1

0

0

0

a.3 2

0

4

2

1.5 4

3

1

3.9 2

0

4.2 0

10

7

11

11

a

22

6

0

5

20

19

13

20

67

56

12

4 1

3

20

22.7

10.0

15.0

9.0

12.5

0.0

0.0

7 26 19.2

7 45 11.1

24 29.2 104 19.2

a 7 1 28.2

27 11.1 135 11.1

4 29 24.1

5 13

0 5

4 35 17.1

7 58 19.0

23

7

17.4 99 9.1

32 la.8

47 23.4 234 16.7

5 37 13.5

1 3 92 10.9

0 1

29 10.3 163 7.4

1

2

a

1

0

1

1

0

2

6

3

2

9

1 2

7

7

5

0

1 9

3

41

62

14

3

10

22

1

1 2

76

33

37

62

119

74

50

41

2

179

4.9

16.1

13.6

13.2

9.1

16.2

19.4

10.9

12.2

20.0

7.3

Average 3 13 4.2 1 18 9.4 15 21.8 81 14.6 5 44 11.2

Highlighted table values denote a significantly greater percentage of plots (p c 0.05) with both dead and removed trees than adjacent values for plots with dead

trees and no removed trees. Percentages are only shown for categories with at least 20 sample plots.