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U SD United States 


———— Department of 


Agriculture 
Forest Service 


Northeastern 
Area 


Morgantown, WV 


NA-TP-04-99 


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Changes in Composition 


Dar ie Sen 
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jiu; of is 
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_of the 
Mixed Mesophytic Forest: 





Effects of Succession 


and Disturbance 





About the Author 


Jim Steinman is a research forester with the USDA Forest Service Northeastern 
Research Station, Radnor, PA. He joined the Station’s Forest Health Monitoring 
Program in 1996 and has been conducting research on methods to quantify forest 
health. Dr. Steinman received his B.S. and M.S. degrees in forest biology from 
The Ohio State University in 1981 and 1983, respectively, and his Ph.D. in forest 
biology from The University of Maine in 1992. 


Acknowledgments 


I wish to thank Gerry Hertel and Dan Twardus of the USDA Forest Service for 
sponsoring this study and providing expert advice. I am deeply grateful to a 
number of my colleagues for their helpful reviews of the draft — from the Forest 
Service: Bob Anderson, Bill Burkman, Andy Gillespie, Will McWilliams, 
Manfred Mielke, and Dave Randall; from the Ohio Department of Natural Re- 
sources: Dan Balser; and from the West Virginia Department of Agriculture: 
Sherri Hutchinson and Clark Haynes. Special thanks to Tom Hall and John 
Quimby of the Pennsylvania Bureau of Forestry for their thorough critique. 


Jim Steinman 


Credits 


Editors: Lennie Eav and Brenda Wilkins 
Format and layout: Nancy Lough 


Printing and production: Patty Dougherty and Tinathan Coger 


Cover Photo: Courtesy of Dan Balser of Ohio Department of Natural Resources 








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USDA A United States 


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 


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1. Differences in Species Composition in the Last 50 Years ............cceee cece a 
Historical Changes in South@astern Kentucky ..........cccseeeeccecccneeecceccceeeecessenesees i 
FIA Data: Composition of Mixed MesophytiC FOrest ...........:ccccccseeeeccneeeeccesesenes 8 
PM ALCLOMMICCAVIOL CII MeeenmnnnT tet rec ccstieeotcc estes sccsccccssesssscircuesttesteneseracreeasesavas 9 

This section of the study ascertained whether trees were dying 
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List of Tables 


Table 1 — Characteristics of Ecological Subregions that Contain the Mixed Mesophytic Forest 
(McNab: and Avers:1 994). oon occ crscteeiioccacserececsettstcccecsavconenst oe cinteecee: Oeo te rnedur meen: cs aemie eit tte aaa 3 


Table 2 — Distribution of FIA Sample Plots with Trees at least 10 inches DBH (by Forest Type 
and Ecological Subregion). 2:.c.cscsteicesc.ceenptncesetecteecs coe sesasadcrese sas Sere cie reese: sence erent eet ee 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 ............::::0 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 mesopntyic forest (Braun 1950) and ecological subregions 
(McNab' and Avers 1994) ror. cerscte sige cen tet deen tcentae cere tees ne name ie cement 2 


Figure 3. Species composition in southeastern Kentucky during the 1940s and 1988 as estimated 
by Braun (1950) and from FIA sample plots; respectively: 22.c-c.:::.-:2---<-e2eeas--nceteche-- scp eee meee neen eee 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 


(FLAY Catal) Perr ereccererettrte rs tecentecererte tts tote serrate cc cetn tat gist ee te cnr nena eet tt eee 9 
Figure 6. Percentages of dead trees and removed trees in oak-hickory forest types by tolerance 

to competition (FIA data). ce sin -eitis ce eis ee aren a cee ee 10 
Figure 7. Percentages of dead trees and removed trees in northern hardwood forest types by 

tolerance :to'competition (FIA data). 2... cccc-seectencrcorecsane seeeeeeeeeene ty ete inate Sey Bi ean Ree ce 11 
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). «0.0.0.0... eee a 
Figure 9. Percentages of dead trees and removed trees in oak-hickory forest types by tolerance 

to competition;and ecological'subregion (FIA data) @ ie... .e.;srntec: corer ses eve yeeee oes ee 12 
Figure 10. Percentages of dead trees and removed trees in northern hardwood forest types by 

tolerance to competition and ecological subregion (FIA data). .............cccccccccccscsseseeeeeeecceeeeeeeenaaes Ve 
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 
withoutievidence of any removed) tlCGS ier cres.ecscetereti cess seer eee eee en 19 





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 descrip- 
tions of the many tree species observed in the 
overstory (see Appendix Table A-1), including 
counts of overstory trees along sampled transects. 
Summaries of Braun’s published counts (see Appen- 
dix 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 intoler- 
ant 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 composi- 
tion, 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 (Murr.) 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. How- 
ever, most studies to determine the effects of ambi- 
ent air pollutants on mature forest trees have shown 
inconclusive results due to the following factors: 


e Difficulty and expense involved in conducting 
controlled “cause and effect” field experiments. 


e Lack of reference or benchmark data with which 
to compare increases in air pollution 





data. 


e Reliance on evaluations that associ- 
ate tree mortality and ambient air 
pollution along spatial gradients 
(Shriner and others 1990). 


PA 








Studies of tree mortality and ambient 
air pollution in the northern Appala- 
chian region have only been able to 
demonstrate a relationship between 




















FON mortality of red spruce and concentra- 
oy 2? m tions of air pollutants at high eleva- 
The Mixed Mesophytic Forest — tions. Even so, two hypotheses exist 
Let nau Vo _ for the mixed mesophytic forest. 

Pe Field Sampling by Braun These hypotheses state that: (1) larger 

ers Ln trees of some species are dying at an 

y, |G Cumberland Mountains a2Ts accelerated rate, and (2) mortality is 

GA = Bre a mcine agi | due to the deposition of airborne 
10,140 | nitrogen and exposure to ozone, which 





Figure 1. States containing the mixed mesophytic forest and 


counties sampled by Braun (1950). 


predispose trees to root pathogens 
(Little 1995). 


Study Design 


The current analysis focused on testing the hypoth- 
esis that trees of some species are dying at acceler- 
ated 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 com- 
pare 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 
that deviate from forest-wide averages. These 
results were then used to infer historical differences 
in species composition between the 1940s and 
1980s, and the recent changes in species composi- 
tion and forest succession trends. 


While this study did not directly 


Methods 


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 231C of Alabama was 
retained for analyses, although it has a warmer 
climate and more loblolly pine than the other subre- 
gions. 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. 





address the hypothesis that air 
pollutants cause trees to die, 
FIA data were utilized to 
identify likely causes of mortal- 
ity 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 














wf 


The Mixed Mesophytic Forest | 
(Braun 1950) 











M221 B 


a 


221A 





Ecological Subregions 
(McNab and Avers 1994) 








American Maple Projects 
(Twardus and Mielke 1995). 


Figure 2. Delineations of the mixed mesophtyic forest (Braun 1 950) 
and ecological subregions (McNab and Avers 1994). 


298 


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 morpholo Materials Soil Taxa Vegetation ft Temp (F days Land Use 
212 Laurentian Mixed Forest Province 
212G dissected sandstone, __Ultisols Hemlock-N. 1000-2400 40-50 120-150 Fire; forestry, 
plateau siltstone, Inceptisols hardwoods 46-48 oil, 
shale Appal. oak- agriculture 
pine 
221 Eastern Broadleaf Forest Province 
221E dissected sandstone, __Ultisols Appalachian 660-1350 35-45 120-170 Fire, clearing; 
plateau siltstone, Inceptisols | hardwoods 39-55 agriculture, 
shale, coal urban 
221H dissected sandstone, __Ultisols Mixed; 650-980 46-55 175 Fire; forestry 
plateau shale, coal Inceptisols mesophytic, 
Appal. oak 
2211 Faulted/ sandstone, __Ultisols Appal. oak, 800-1000 46-55 175 Fire; forestry 
folded shale, Inceptisols Mixed 
monoclinal llimestone Mesoph. 
mountains 
231 Southeastern Mixed Forest Province 
231C Faulted/ stratified Ultisols Oak-hickory- 330-1330 50-55 200-210 Fire, drought; 
folded marine Inceptisols pine, 60-62 forestry 
monoclinal deposits Southeastern 
mountains mixed 
M221 Central Appalachian Broadleaf Forest - Coniferous Forest - Meadow Province 
M221A Faulted/ limestone, Inceptisols § Appal. oak, 300-4800 35-50 120-170 Fire; agric., 
folded sandstone, __Ultisols oak-hickory, 46-60 forestry, 
parallel shale pine urban 
ridges 
M221B Severely sandstone, Inceptisols Mixed 1000-4600 40-60 110-160 Fire; forestry 
dissected shale Ultisols hardwoods, 39-54 
plateau spruce-fir 
M221C Faulted/ sandstone, Inceptisols Mixed 2000-2600 40-47 140-160 Fire; 
folded shale Ultisols mesophytic, 45-50 agriculture, 
monoclinal Appal. oak forestry 
mountains 


ae 


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 | 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 cumula- 
tive 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), Tennes- 
see (1989), and West Virginia (1989). A few plots 
in the FIA data included sample plots from Mary- 
land (1986) and Virginia (1992). 


-4- 


The area of each sample plot was commonly 1/5 
acre, but varied among locations (1/6 to 1/4 acre) as 
a function of different sampling designs. Variables 
used from plot records included approximate loca- 
tions (latitude and longitude), forest type, stand size 
(trees dominated by saplings | to 5 inches dbh), 
poletimber (5 to 10 inches dbh), or sawtimber (>10 
inches dbh), and stand basal area (total cross-sec- 
tional 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. Vari- 
ables 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 facili- 
tated more robust comparisons among a variety of 
species groups. However, greater emphasis was 
placed on species that Braun described as predomi- 
nant in the overstory than on other species. Thus, 
analyses were confined to FIA plots within oak- 
hickory, 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 oak- 
hickory 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 


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 miscella- 
neous category. Each species was commonly 
represented by only one or two trees on a sample 
plot, and rarely by more than five trees. 


Analytical Procedure 


Analytical procedures to evaluate species composi- 
tion, 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 com- 
pared 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 were analyzed individually and in groups 
based on whether they were intolerant, moderately 
tolerant, or tolerant of competition (Burns and 
Honkala 1990a; 1990b). The tolerance ratings of 
species generally correspond to the successional 
stage in which they predominate, where tolerant 
species characterize a late successional forest. A 
previous study showed differences in stocking 


among species that vary in tolerance to competition 
and different successional stages of the forest 
(USDA Forest Service 1995). 


Species composition of the forest was evaluated by 
combining tree data from all FIA plots within a 
given forest type or ecological subregion. Percent- 
ages 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. 


Proportions of dead trees and removed trees were 
analyzed separately using FIA data. Percentages of 
all live, dead, and removed trees were used to 
express the amount of dead and removed trees 
among species in a given stratum. Expressing the 
amount of dead and removed trees for all species in 
each stratum allowed comparison of averages 
(higher or lower) among species. 


Table 2 
Distribution of FIA Sample Plots with Trees at least 10 inches DBH 
(by Forest Type and Ecological Subregion) 


Ecological Subregion 





Forest Type All 212G 221E 221H 2211 M221A M221B M221C 231C 
Seaaeesarenaqwescanwenancasnaneasomnsarmcnoana= Percent of plots -------------------------------------------00=---0"" 

Oak-Hickory 68.1 30.3 72.4 75.6 82.7 UPS) 63.4 89.8 49.3 

Northern 

Hardwood 18.5 62.6 16.5 Ze 5D 12.8 31.9 8.1 0.0 

Oak-Pine si 1a Zaft i paliees Ths 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 844 226 204 832 521 276 

Softwood 8.3 6.1 8.4 11.0 4.6 6.4 4.1 1.3 Clee 

and 

Other Types n= 490 34 181 104 11 14 36 7 103 

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


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 


Number of trees per sample ee eae aC TSoRE with at na eee 
Trees Be eee 
pone = 


Percent of 5404 plots 









Ash sp. 215%, 80.7 15.2 2.9 1.0 0.2 Ce ae 
Aspen sp. 594 95.4 3.0 Wer 0.2 0.1 246 82 
Basswood sp. 1457 89.5 6.9 2.6 0.8 0.2 568 193 
Beech 4446 Tie 122 5.8 3.7 121 1233 5/5 
Birch sp. 1791 86.7 9.4 ALTE 1.0 0.2 lig, 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 On, 0.1 601 153 
Black walnut 617 93.0 6.0 0.9 On 0.0 380 32) 
Blackgum 791 89.2 9.9 0.8 0.1 0.0 581 45 
Buckeye sp. 280 97.2 2.3 0.4 0.1 0.0 149 25 
Cucumbertree 766 91.8 6.8 led 0.2 0.0 441 73 
Elm sp. 869 91.9 6.3 iets) 0.3 0.0 440 100 
Hemlock sp. 923 93.5 4.0 eS) 0.6 0.1 350 133 
Hickory sp. 5208 60.7 26.5 9.7 PET 0.4 2124 692 
Magnolia sp. 156 98.3 eS 0.2 0.1 0.0 94 15 
Maple, red 8047 58.2 24.7 10.1 4.1 2.9 2259 925 
Maple, sugar 4624 73.1 16.7 5.9 2.9 3 1452 547 
Oak, black 5236 65.9 21.0 8.9 3.6 0.7 1845 711 
Oak, chestnut 8114 65.2 Set 10.4 6.6 2a, 1883 1066 
Oak, northern red 7626 56.2 29 11.4 4.6 1.9 2367 969 
Oak, other red 555 95.6 3.3 0.9 0.2 0.1 239 60 
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 47 
Oak, white 8069 56.4 23.2 12.6 5.8 1.9 2356 1100 
Pine, loblolly 468 97.7 1.0 Ona 0.4 0.1 124 68 
Pine, pitch 281 97.2 2.2 0.5 0.0 0.0 149 32 
Pine, shortleaf 598 95.4 3.3 0.9 0.3 0.1 247 70 
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 tS? 55 
Sassafras 463 95.4 3.9 0.5 0.1 0.1 249 38 
Sweetgum 263 98.0 1.3 0.5 0.1 0.0 107 35 
Yellow-poplar 9280 58.0 20.2 118 6.8 3.1 2267 1174 
Other species 1075 87.7 2.1 0.3 0.0 0.0 663 16 


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 percent- 
age 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 


Results and Discussion 


1. Differences in Species 
Composition in the Last 50 Years 


Historical Changes in Southeastern 
Kentucky 


In the 1940s, 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-1). Data 
collected from FIA plots in the same counties during 








compared. ‘Intolerant Species 
Poa. yellow-poplar 
To determine if dead trees Rcrorvcn 
and removed trees on the een oak 
same plot were associated, blackiocast 
each plot was designated as pine sp. 
having removals present or ash sp. 
absent depending on evidence ewaest birch Total 
of at least one removed tree Peermclone @ 1988: 38% 
of any species. To facilitate plac menery CO) 1940s: 18% 
statistical tests of association, - 5 "i i 0 
analyses were confined to percent of trees 
species represented on at least Moderately Tolerant Species 
50 sample plots. The percent- 
A chestnut oak 
ages of plots with dead trees ogee 
of individual species were 
d veto f black oak 
etermine or t 1€ groups O mn greed 
plots with and without Total 
: cucumbertree =< 
removals, and tests of inde- Herren (Absent in FIA Survey) Hl 1988: 37% 
pendence were used to Sine [1 1940s: 28% 
determine if percentages were 4 at 
ners . 0 5 10 15 20 
significantly different (p < pereentne rons 
0.05) for groups that were 
compared. Tolerant Species 
red maple 
Maps of plot locations and beech 
their corresponding attributes sugar maple 
were also used to show where basswood sp. Total 
dead trees were present or blackgum one! rea 
. ° 
absent for individual tree hemlock sp. 
doce ter came < Het ae cr) 1940s: 53% 
species. Spatial distributions yellow buckeye 
of plots containing dead trees : ; a is aS 
were visually interpreted to percent of trees 
























determine if plots occurred in 
groups or if they appeared to 
be randomly scattered. 


Gay 


Figure 3. Species composition in southeastern Kentucky during the 
1940s and 1988 as estimated by Braun (1950) and from FIA sample 
plots, respectively. 


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 com- 
prised 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 


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 Appen- 
dix Table C-1) shows that the oak-hickory types 
throughout the forest had a composition similar to 
southeastern Kentucky in 1988 (see Figure 3). 


event. However, correspond- 
ing 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 vat. 
faginata (Pers.:Fr.)] or hem- 
lock 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 out- 
competed by other species. It 
is more likely that distur- 
bances such as logging and 
land clearing created forest 
openings that were rapidly 
colonized by early succes- 
sional species well adapted to 
increased light, moisture, and 
nutrient availability. 


FIA data show that early 
successional species, includ- 
ing 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). 











Oak-hickory Forest Types 
Intolerant Species 

yellow-poplar 
hickory sp. 
scarlet oak 
black locust 
pine sp. 
ash sp. 
sweet birch 
black walnut 
black cherry 





Total: 38% 





0 5 10 15 20 
percent of trees> 10” dbh 


Moderately Tolerant Species 


chestnut oak 
white oak 
black oak 
north. red oak 
cucumbertree 
chestnut 

elm sp. 





(Absent in FIA Survey) 


Total: 44% 


——_+— —+— ++ 
0 5 10 15 20 
percent of trees > 10” dbh 








Tolerant Species 


red maple 
beech 

sugar maple 
basswood sp. 
blackgum 


hemlock sp. Total: 16% 


| 





yellow buckeye 





0 5 


10 
percent of trees > 10” dbh 








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


Ege 


Conversely, Figure 5 shows that the composition of 
northern hardwood forest types more closely re- 
sembled 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. Con- 
versely, high percentages of oak species in the oak- 
hickory 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 
species, and those in the 





Northern Hardwood Forest Types 


Intolerant Species 
yellow-poplar 
hickory sp. 
scarlet oak 
black locust 
pine sp. 
ash sp. 
sweet birch 


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 C- 
1). Pine species comprised 








black walnut Total: 41% about 40 percent of the oak- 
black cherry pine forest and consisted of 
0 5 10 15 20 Virginia pine, shortleaf pine, 
ec Sagi Sma loblolly pine, eastern white 
Moderately Tolerant Species pine, and pitch pine. Only 7 
chestnut oak percent of the trees in this type 
white oak were of species tolerant of 
black oak competition. 
north. red oak 
cucumbertree 2. Rate of Tree 
chestnut (Absent in FIA Survey) Total: 9% M ort al ity 
elm sp. ei ' —. 4 
: ape peryeymetgsgrigemy tH a This section of the study 


Tolerant Species 
red maple 
beech 
sugar maple 
basswood sp. 
blackgum 
hemlock sp. 
yellow buckeye 


0 5 10 








percent of trees > 10’ 


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 


Total: 49% 


20 


15 


* dbh 





Figure 5. Species composition of northern hardwood forest types by 


tolerance to competition (FIA data). 


as compared to 3 percent in the 
oak-hickory types. 


age 


Records of dead trees and removed trees in the FIA 
database represent cumulative amounts of mortality 
and cutting that occurred during the 10- to 15-year 
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 


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 percent- 
ages 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 


and consequent changes in 
species composition. In the 
oak-hickory forest type, 7 
percent of trees intolerant of 
competition were dead com- 
pared 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 competi- 
tion 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. 


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 
of competition and representa- 
tive of early stages of forest 
succession. Although elm is 
moderately tolerant of compe- 
tition, high percentages of 
dead elm were most likely due 





oak-hickory forest. Within the northern hardwood 





Oak-hickory Forest Types 


Intolerant Species 
yellow-poplar 
hickory sp. 
scarlet oak 
black locust 
pine sp. 
ash sp. 
sweet birch 
black walnut 
black cherry 







Mean 
@ Dead: 7% 
(11 Removed: 4% 








0 5 10 15 20 Zo) 
percent of trees> 10” dbh 


Moderately Tolerant Species 


chestnut oak 








white oak 
black oak 
north. red oak Mean 
cucumbertree @ Dead: 5% 
chestnut (Absent in FIA Survey) [C1 Removed: 4% 
es 
0 5 10 15 20 25 
percent of trees > 10” dbh 

Tolerant Species 
red maple 
beech 
sugar maple 
basswood sp. 
blackgum Mean 
hemlock sp. @ Dead: 4% 


oO - 90 
yellow buckeye Removed: 2% 


0 5 10 15 20 25 
percent of trees > 10” dbh 








Figure 6. Percentages of dead trees and removed trees in oak-hickory 
forest types by tolerance to competition (FIA data). 


“Nis 








forest, species moderately tolerant of competition 
were removed more frequently than the tolerant and 
intolerant species (see Figure 7 and Appendix Table 
C-1). For example, black oak, chestnut oak, north- 
ern 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 yellow- 
poplar) were also removed. 


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 signifi- 
cantly greater among larger trees of hickory, scarlet 
oak, shortleaf pine, Virginia pine, elm, magnolia, 


and beech. However, for all combined species 
intolerant of competition (Figure 8), there were more 
dead trees in the 10 to 15 inch class. 


10 





DBH class (inches) 
M10-15 [)>15 


percent of trees 





INT MOD TOL 
Dead 


Figure 8. Percentages of dead trees and 


INT MOD TOL 








Removed 








Northern Hardwood Forest Types 


Intolerant Species 
yellow-poplar 
hickory sp. 
scarlet oak 
black locust 
pine sp. 
ash sp. 
sweet birch 
black walnut 
black cherry 


@ Dead: 






[1 Removed: 6% 
+ —+ { 


removed trees by dbh class and tolerance to 
competition: Intolerant (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 


Mean 
8% 








0 5 10 15 


percent of trees> 10” dbh 
Moderately Tolerant Species 


20 


chestnut oak 
white oak 
black oak 
north. red oak 






cucumbertree 
chestnut 
elm sp. 


@ Dead: 








(Absent in FIA Survey) 





10 
percent of trees > 10” dbh 


0 5 15 20 


Tolerant Species 
red maple 
beech 
sugar maple 
basswood sp. 
blackgum 
hemlock sp. 
yellow buckeye 


@ Dead: 





= 
10 
percent of trees > 10” dbh 





0 5 15 20 





CO Removed: 13% 


[ Removed: 5% 
$+ 


me and structure are partly due to a dispropor- 


tionate removal of large trees of marketable 
species (Birch and others 1992). 


Mortality by Geographic Locations 





Mean 
° 


5 Significantly greater percentages of dead 


trees belonged to intolerant species within 
oak-hickory forests (Figure 9) at the Appala- 
chian 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 


25 


Mean 
5% 








25 





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


(FIA data). 


forests of the Appalachian plateau than in 
other subregions (Figure 10). 


Fite 








Oak-hickory Forest Types 
Intolerant Species 


212G 
221E 
M221A 
M221B 
M221C 
221H 
2211 
231C 


Mean 

@ Dead: 7% 
[J Removed: 4% 
<p 


Ecological Subregion 








a 


0 15 20 


10 
percent of trees> 10” dbh 
Moderately Tolerant Species 


212G 
221E 
M221A 
M221B 
M221C 
221H 
2211 
231C 


Mean 
® Dead: 5% 
[1 Removed: 4% 
+H 


Ecological Subregion 








+ “= 


tt) 5 15 20 


10 
percent of trees > 10” dbh 
Tolerant Species 


212G 
221E 
M221A 
M221B 
M221C 
221H 
2211 
231C 






Mean 
@ Dead: 4% 
tf Removed: 2% 


Ecological Subregion 


15 20 


0 5 


10 
percent of trees > 10” dbh 





Northern Hardwood Forest Types 
Intolerant Species 


212G 
221E 
M221A 
M221B 
M221C 
221H 
2211 
231C 






Mean 


@ Dead: 8% 
( Removed: 6% 
+ — 


15 20 


Ecological Subregion 


} (insufficient data) 





a 





+ 


0 10 
percent of trees> 10” dbh 


Moderately Tolerant Species 


212G 
221E 
M221A 
M221B 
M221C 
221H 
2211 
231C 





Mean 
® Dead: 9% 


} (insufficient data) 
(1) Removed: 13% 


Ecological Subregion 





————— + 


tt) 5 15 20 


10 
percent of trees > 10” dbh 
Tolerant Species 


212G 
221E 
M221A 
M221B 
M221C 
221H 
2211 
231C 





Mean 
Wi Dead: 5% 
t] Removed: 5% 


} (insufficient data) 


Ecological Subregion 





+ + 
0 5 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 yellow- 
poplar 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 subre- 
gions. Throughout the region, an average of 20 


Figure 10. 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 212G con- 
tained 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 221H 
also contained many species with dead trees on more 
than 20 percent of the plots (see Appendix Table 
D-1). 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 subre- 
gion 221H contained dead beech, hickory, black oak, 
and scarlet oak. 


2102 








Ash species 


\ 
Ecological Number Percent with eT. Ecological Number Percent with 
Subregion ofPlots Dead Trees / Subregion of Plots Dead Trees 
54 28 


231C 
M221A 
M221B 





%\ 
Sten Be Boat toe 
Subregion ofPlots Dead Trees Subregion of Plots Dead Trees 
23 
27 ae 
34 
28 
0 
20 \ 
21 M221B 
19 M221C 





Dead Trees greater than 10 inches DBH: 
© Present ® Absent 


Figure 11. FIA plots containing at least three trees greater than 10 inches dbh. 


=13: 


Black Cherry 


Ecological Number Percent with 
Subregion of Plots Dead Trees 








Ecological Number Percent with 
Subregion ofPlots Dead Trees 





212G 8 
221E 12 
221H 

2211 

231C 

M221A 

M221B 

M221C 





Dead Trees greater than 10 inches DBH: 


® Present 


wT AS 


® Absent 


Figure 12. FIA plots containing at least three trees greater than 10 inches dbh. 


Ecological Number Percent with 
Subregion of Plots Dead Trees 


Ecological Number Percent with 
Subregion of Plots Dead Trees 





212G 
221E 
221H 
2211 
231C 
M221A 
M221B 
M221C 








Ecological Number Percent with 
| Subregion of Plots Dead Trees 
40 
33 








Ecological Number Percent with 
| Subregion of Plots Dead Trees 


212G 
221E 
221H 
2211 
231C 
M221A 
M221B 
M221C 





Dead Trees greater than 10 inches DBH: 


® Present 


Hickory species 


NGS 





® Absent 


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


Ecological Number Percent with 
Subregion of Plots Dead Trees 








Ecological Number Percent with 
Subregion of Plots Dead Trees 





212G 137 
221E 185 
221H 28 
2211 

231C 

M221A 

M221B 

M221C 


Ecological Number Percent with 
Subregion of Plots Dead Trees 





Northern Red Oak 


Ecological Number Percent with 
Subregion ofPlots Dead Trees 





212G 27 
221E 352 221E 
221H 154 221H 
2211 27 2211 
231C 231C 
M221A M221A 
M221B M221B 
M221C 


All 








White oak 


Ecological Number Percent with Ecological Number Percent with 
Subregion of Plots Dead Trees Subregion ofPlots Dead Trees 


212G 36 212G 70 26 
221E 23 221E 499 13 
221H 15 221H 237 14 
2211 18 2211 38 21 
231C 1 . 231C 62 3 
M221A 10 M221A 43 0 
M221B 22 M221B 105 11 
M221C 14 M221C 46 7 


All 18 All 1100 13 

















Dead Trees greater than 10 inches DBH: 
® Present ® Absent 


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


6: 


Scarlet Oak 


Number Percent with 
Subregion of Plots Dead Trees 





Dead Trees greater than 10 inches DBH: 
© Present ® Absent 


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


he 


Ecological Number Percent with 
Subregion of Plots Dead Trees 


212G 
221E 
221H 
2211 
231C 
M221A 
M221B 
M221C 





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 charac- 
teristic 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 geo- 
graphic locations as a function of inherent differ- 
ences 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 north- 
ern 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 


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 propor- 
tion 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 mid- 
successional 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 competi- 
tion (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 evi- 
dence 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. 


Bie 








Intolerant Species 


Virginia pine 
ARS RS ie ere 


black locust 7X7 
scarletoak [a ee 
sweet birch [asta et | 
hickory sp. Fema ene 

ash sp. ee ee 


black cherry [eo 
yellow-polla 


0 10 20 30 40 50 
percent of plots with dead trees present 


Moderately Tolerant Species 





elm sp. 

black oak 
north. red oak 
chestnut oak 
cucumbertree 


(Insufficient data) 


white oak 





0 10 20 30 40 50 
percent of plots with dead trees present 


Tolerant Species 
beech 
red maple 
hemlock sp. 





sugar maple 
basswood sp. 
0 10 20 30 40 50 


percent of plots with dead trees present 


Evidence of Removals: @ Present © Absent 








Figure 16. Percentages of FIA 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 non- 
forested 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. 


tee 


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 distur- 
bances 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-1). 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). 


Air Pollution 


Tree species of the mixed mesophytic forest that are 
sensitive to sulfur, nitrogen and/or ozone air pollut- 
ants 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 relation- 
ships 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 addi- 
tion, 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 composi- 
tion 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 deter- 
mined 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 compo- 


sition 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 publica- 
tion was written (1998). The FHM plots are less 
numerous than FIA plots, but include more measure- 
ments of forest health including conditions of tree 
crowns and evidence of tree damage. The abun- 
dance 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 distur- 
bances 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: 


LY 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. 


L) 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. 


L) 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. 


0: 


L) 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. 


L) 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 de- 
scribed by Braun. 


L) 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. 


CL) 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. 


oD Rae EAP tae eas 


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 post- 
logging 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. 


Burns, 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., comps. 1994. Ecological 
subregions of the United States: Section 
descriptions. Administrative Publication WO- 
WSA-S5. 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. 


=p 


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: 313-315. 


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, 
ING 2709 m3 2500: 


USDA Forest Service. 1995. Shifts in stocking reveal 
forest health problems. NA-TP-07-95. Radnor, 
PA: USDA Forest Service, Northeastern Forest 
Experiment Station. 9 p. 


Bye 

















Appendices 


List of Appendices 


Table A-1 
Common and Scientific Names of Trees in the Overstory of the Mixed Mesophytic Forest..................+ A-1 


Table B-1 
Species Composition of Different Geographic Areas as Summarized from Data by Braun (1950) .......... B-1 


Table C-1 
Percentages of species composition (C), dead trees (D), and removed trees (R) 
by forest type and shade tolerance for all combined ecological SUDreGiONS. ..............cceceeeeeeeeeeeeeeeeeeeeeeees C-1 


Table C-2 
Percentages of dead trees and removed trees on FIA sample plots by diameter class. ..............:.::::::e0+ 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 SUDregiOn 212G. ...........cccccccccesseeeeeeeeeeeeeeeeeeeeeeaeeeeeeeees 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 221E. ................cccccssssessseeeccccccessecescecccennsees 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 M221A. .............ccccccccccsseeeeccecceeneeeeseeseeaeners C-5 


Table C-6 
Percentages of species composition (C), dead trees (D), and removed trees (R) 
by forest type and shade toierance for ecological Subregion M221B. ..............::::ssssesseeceeeeeceeeeeeeeeeeeeeaeaees 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 M221C. ..............ccccccssssssseeseeceecenseecseeseeeneess 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 221H. .................csssecereeceeeeccecsesssssneceassensns 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 221L. ....................sssssceessescocssccccceceseseeseenees C-9 


Table C-10 

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

by forest type and shade tolerance for ecological subregion 231C. ..................ssssssesssssssssscccccececeeeeeeeees C-10 
Table D-1 

Percentages of FIA plots with dead tree species by ecological SUDreGiON. ...............cccescssecesssesceeeeeteeeeees D-1 


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. .........ccccc6e..6.. D-3 


Table D-4 
Percentages of FIA plots with dead tree species on plots with and without tree removals. ..............00.06+. D-4 











Table A-1 


Common and Scientific Names of Trees in the Overstory 


Common Name 


Ash, Green 

Ash, White 

Aspen, Bigtooth 
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-Poplar 


of the Mixed Mesophytic Forest 





Scientific Name 





Fraxinus pennsylvanica Marsh. 
Fraxinus americana L. 

Populus gandidetata Michx. 
Populus tremuloides Michx. 
Tilia americana L 

Tilia heterophylla Vent. 

Fagus grandiflora Ehrh 

Betula lenta L. 

Betula allehaniensis Britton 
Nyssa sylvatica Marsh. 
Aesculus glabra Wild 
Aesculus octandra Marsh. 
Prunus serotina Ehrh. 
Castanea dentata (Marsh.) Borkh. 
Magnolia acuminata L. 

UlImus americana L. 

UlImus rubra Muhl. 

Tsuga caroliniana Englem. 
Tsuga canadensis (L.) Carr. 
Carya cordiformis (Wangenh.) K. Koch 
Carya tomentosa (Poir.) Nutt. 
Carya glabra (Mill.) Sweet 
Carya ovata (Mill.) K. Koch. 
Carya laciniosa (Michx. f.) Loud. 
Robinia psuedoacacia L. 
Magnolia fraseri Walt. 

Acer rubrum L. 

Acer saccharum Marsh. 
Quercus velutina Lam. 
Quercus prinus L. 

Quercus rubra L. 

Quercus coccinea Muenchh. 
Quercus shumardii Buckl. 
Quercus falcata Michx. 
Quercus alba L. 

Diospyros virginiana L. 

Pinus strobus L. 

Pinus taeda L. 

Pinus rigida Mill. 

Pinus echinata Mill. 

Pinus virginiana Mill. 
Sassafras albidum (Nutt.) Nees 
Liquidambar styraciflua L. 
Juglans nigra L. 

Liriodendron tulipifera L. 





A-1 








Table B-1 
Species Composition of Different Geographic Areas 
as Summarized from Data by Braun (1950) 





Cumberland Appalachian Allegheny All Areas 
Species Mountains Plateau Mountains (n=10140) 


(n=7027) (n=2250) (n=863) 




















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 
All Intolerants 18.5 23.7 20.3 19.8 
Chestnut 9.5 1.5 13.4 8.1 
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 PAE 0.9 4.8 
Oak, n. red 3.0 3.1 6.7 Koa} 
Oak, white 7.1 13.0 1.1 7.9 
All Moderates 27.5 24.8 27.7 26.9 
Basswood sp. 6.3 4.0 8.5 6.0 
Beech 16.9 20.4 17.7 17.7 
Blackgum 1.9 2.8 1.9 
Buckeye sp. 4.8 1.5 3.7 
Hemlock sp. 7.6 15.5 8.7 
Maple, red 2.5 2.6 10.7 3.2 
Maple, sugar 12.2 3.9 15.0 10.6 
All Tolerants 52.2 50.6 51.9 51.8 
Other species 1.8 0.9 0.1 1.5 
Total 100.0 100.0 100.0 100.0 





Table C-1 
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 %R n= %C 

Ash sp. 1030 iro 3.6 1098 56 69 5 29 0.7 ae. 
Aspen sp. 308 0.6 14.0 WZ 280 1.7 e242 2.9 6 0.2 --- --- 
Birch sp. 625 12 7.7 1.4 1156 6.5 12.1 4.7 10 0.3 --- 

Black cherry 1174 2.0 55 4.9 4031 19.7 LS} 2 5.9 35 0.9 5.1 0.0 
Black locust 1054 1.9 20.6 et 293 1.7 24.9 4.8 3 0.1 --- --- 
Black walnut 502 0.9 9.3 4.6 104 0.6 4.7 9.9 if 0.4 --- --- 
Hickory sp. 4684 8.4 6.7 2.3 345 1.8 Si7/ es 179 Sys) CHS 0.7 
Oak, scarlet 3103 4.8 10.3 4.3 19 0.1 --- --- 155 3.7 9.9 0.9 
Pine, loblolly 145 0.2 0.4 24.9 0 323 7.7 1.1 6.1 
Pine, pitch 207 0.4 21-7 4.2 2 0.0 --- --- 72 2.0 FAT7 Pe! 
Pine, shortleaf 274 0.66 12.4 10.1 0 324 9.9 10.1 4.5 
Pine, Virginia 500 1.0 20.1 9.4 16 0.1 --- --- 655 19.6 7.9 2.6 
Sassafras 422 0.8 18.9 0.6 40 0.3 17.2 0.0 1 0.0 --- 
Sweetgum 213 0.4 4.5 8.9 2 0.0 --- --- 48 1.4 5.3 0.0 
Yellow-poplar 8378 © 13.4 PT 3.4 613 310 ZAG 9.6 289 7.0 PT 2.0 
All Intolerant 22619 38.4 7.3 3.6 7999 41.0 8.1 5.7 2140 59.4 6.4 2.9 
Cucumbertree 580 0.9 3.4 1.8 183 0.9 145 10.7 3 0.1 --- --- 
Elm sp. 549 1.0 18.6 2.8 308 1.7 11.9 97 12 0.4 --- --- 
Magnolia sp. 122 0.2 0.8 Zao 32 0.2 4.2 0.0 2 0.1 --- --- 
Oak, black 4966 7.2 6.5 Sat 114 0.5 TT NTA 156 3.6 6.8 2.9 
Oak, chestnut 7800 Wel2it 4.0 3:3 ili 0.6 16.1 12.1 203 4.9 0.1 1.8 
Oak, northern red 6782 9.5 4.5 BZ 730 3:2 6.1 17.3 114 2.9 Wet 0.0 
Oak, other red 402 0.7 3.0 PP 9 0.0 --- --- 144 3.6 5.4 0.8 
Oak, other white SiS) 0.5 10.9 2.3 i 0.0 --- --- 107 2.8 Vai2ts 2.4 
Oak, white 7451 11.7 2.8 4.0 213 ill 7.3 BASE 405 9.2 HES eS 
Pine, white 163 0.2 6.9 4.0 69 0:3 8.0 6.8 201 4.0 5.9 0.0 
All Intermediate 29130 44.1 4.6 4.1 1776 8.5 9.1 13.4 1347 831.7 4.0 ude 
Basswood sp. 759 1.2 3.6 3.6 695 SHS 2.9 2.4 3 0.1 --- --- 
Beech 2529 Sal 4.9 lds 1888 8.6 6.9 bai 29 0.7 --- --- 
Blackgum 712 el Sal (Pe 51 0.2 4.3 7.6 28 0.7 --- == 
Buckeye sp. 231 0.4 3.0 1.6 46 02 0.0 3.9 3 0.1 ace 2a 
Hemlock sp. 341 0.5 2.6 Zo 522 2.3 wae9 fs} 60 1.3 Xf 0.0 
Maple, red 4245 7.4 oul 7, 3648 818.5 Shii/ BS) 154 3.9 4.2 0.9 
Maple, sugar 1614 2.6 3.3 2.8 2984 15.4 4.3 4.3 26 0.7 --- 

All Tolerant 10431 16.3 3.6 2.0 9834 48.8 4.8 5.0 303 7.4 41 2.9 
Other Species 707 1.2 19.0 ies 314 16 12.3 7.0 54 1.4 9.7 0.0 


All Species 62887 100 5.6 3.5 19923 100 6.6 6.1 3844 100 5.5 2.4 


Highlighted values represent species that comprise at least 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. 


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






Percent 
Removed 


Number of Percent 
sampled trees Dead 
10-15" | S15") 





2 OSS ee 
Ash sp. 1231 926 6.9 4.8 3.8 5.6 
Aspen sp. 499 95 18.5 18.0 2.2 0.0 
Birch sp. 1392 399 11.1 6.9 Bus 3.0 
Black cherry 2963 2277 6.1 3.2 5.0 7.0 
Black locust 933 417 22.5 Ul 2S 2.9 
Black walnut 406 211 7.8 10.0 4.0 10.1 
Hickory sp. 3451 1757 6.1 8.2 2.1 4.1 
Oak, scarlet 1632 1645 9.2 12.1 3.0 6.3 
Pine, loblolly 246 222 0.9 1.0 13.4 8.0 
Pine, pitch 231 50 20.5 7.0 3.2 7.2 
Pine, shortleaf 502 96 10.7 15.8 6.6 11.2 
Pine, Virginia 1006 165 13.5 15.5 5.3 7.0 
Sassafras 379 84 19.6 11.6 0.6 0.0 
Sweetgum 168 95 4.7 By. 9.0 0.0 
Yellow-poplar 4838 4442 3.0 2.0 27f 5.8 
All Intolerant 19877 12881 8.0 5.8 3.5 5.7 
Cucumbertree 423 343 6.6 4.4 4.2 6). 
Elm sp. 572 297 15.2 19.8 4.8 6.4 
Magnolia sp. 113 43 0.9 3.8 2.0 2.0 
Oak, black 2296 2940 6.9 6.1 Gh.7/ 7.6 
Oak, chestnut 4231 3883 4.4 3.5 PAT 4.8 
Oak, northern red 3218 4408 4.8 4.3 5.0 8.2 
Oak, other red 298 PASY/ 3.9 3.1 1.8 2.4 
Oak, other white 263 166 Wt 9.1 2.3 2.1 
Oak, white 4357 3712 2.9 2.5 2.8 7.2 
Pine, white 181 252 8.3 4.4 0.5 5.5 
All Intermediate 15952 16301 5.0 4.3 3.4 6.7 
Basswood sp. 788 669 3.2 3.5 1.9 5.4 
Beech 1636 2810 5.0 6.8 3.6 3.4 
Blackgum 387 404 4.7 6.0 2.0 3.6 
Buckeye sp. 149 131 Tf ue) 0.5 4.9 
Hemlock sp. 386 537 8.8 6.5 3.4 7.3 
Maple red 5215 2832 3.4 3.4 2.9 4.8 
Maple sugar 2771 1853 4.0 3.8 3.2 5.2 
All Tolerant 11332 9236 4.0 4.7 2.9 4.6 
Other Species 675 400 16.2 18.0 2.7 3.1 
All Species 47836 38818 6.2 5.1 3.3 5.8 





Highlighted values for a given diameter class are significantly greater (p < 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. 


Forest Type 























Oak-Hickory Northern Hardwood 















%D | %R 
Ash sp. 45 le 11.9 0.0 401 5.1 7.6 ae 
Aspen sp. 56 2.1 24.0 2.9 210 3.1 28.4 3.9 
Birch sp. 70 2.6 Onn 0.0 556 8.0 13.8 5.6 
Black cherry 98 3.2 3:7, 0.3 1972 22.9 4.5 B27 
Black locust 
Black walnut 9 0.2 = 223 
Hickory sp. 26 0.8 --- --- 29 0.4 ao _ 
Oak, scarlet 125 4.0 8.1 12.1 4 0.0 --- ae 
Pine, loblolly 
Pine, pitch 22 0.7 --- --- 1 0.0 --- ee 
Pine, shortleaf 
Pine, Virginia 
Sassafras 1 0.0 --- Bes 
Sweetgum 
Yellow-poplar 77 ee 37.2 148) 55 0.6 5.9 0.0 
Elm sp. 10 0.3 --- --- 5 0.1 one = 
Magnolia sp. 
Oak, black 174 5.1 8.7 8.3 ie 0.1 --- --- 
Oak, chestnut 324 10.9 9.3 3.8 21 0.3 --- --- 
Oak, northern red 1100 29.5 4.4 7.8 157 Hlev/a 8.8 8.5 
Oak, other red 3 0.1 --- --- 2 0.0 --- --- 
Oak, other white 
Oak, white 449 14.3 Bye 6.0 40 0.5 13.0 3.7 
Pine, white 15 0.4 --- --- 36 0.4 3.4 3.6 
All Intermediate 2115 61.8 6.0 6.5 327 3.7 11.0 6.3 
Basswood sp. ii 0.3 --- --- 121 1.6 6.5 6.1 
Beech 43 15 9.8 9.9 604 7.4 Bus 7 
Blackgum 2 0.1 =-= --- 4 0.1 --- --- 
Buckeye sp. 
Hemlock sp. 23 0.7 --- --- 206 P26} 14.4 13.1 
Maple, red 505 17.3 2.8 Pf 2209 28.3 ShI/ bi 
Maple, sugar 33 lit 12.6 7.2 1187 16.0 5.4 Ne 
All Tolerant 613 20.9 3.6 3.8 4331 55.6 5.0 5.6 
Other Species 8 0.2 --- --- 34 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 < 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 (R) by 
forest type and shade tolerance for ecological subregion 221E. 


Forest Type 


Oak-Hickory Northern Hardwood 
Line aes 
589 2.6 








Ash sp. Tas 4.2 400 8.1 elt BS 
Aspen sp. 226 eZ 10.6 1.0 56 1.4 eS 0.0 
Birch sp. 103 0.5 6.8 2.5 95 1.9 15.9 0.0 
Black cherry 605 Pall 7.0 5.4 1056 21.7 8.3 7.9 
Black locust 460 2.2 27.4 D7 145 3.2 34.2 6.3 
Black walnut 314 1S 8.9 4.1 87 2.0 3.6 10.1 
Hickory sp. 1880 9.1 6.7 Bath 134 2.8 4.5 8.6 
Oak, scarlet 1152 4.4 9.7 4.8 8 0.2 --- --- 
Pine, loblolly 

Pine, pitch 79 0.4 27.7 Bus} 1 0.0 --- --- 
Pine, shortleaf 22 0.1 --- --- 

Pine, Virginia 221 2 23.7 5) 15) 13 0.3 --- --- 
Sassafras 247 leo 22.4 1.1 32 0.8 18.4 0.0 
Sweetgum 15 0.1 --- --- 

Yellow-poplar 3256 13.6 3.0 4.1 276 5.1 2.4 10.6 
All Intolerant 9169 40.9 8.4 3.7 2303 47.6 9.7 7.0 
Cucumbertree 110 0.4 7.6 1.8 21 0.3 --- --- 
Elm sp. 443 Onl 20.2 2.6 264 5.6 11.4 10.6 
Magnolia sp. 1 0.0 oH oor 1 0.0 --- --- 
Oak, black 2410 9.1 6.7 6.3 67 1.0 9.8 19.3 
Oak, chestnut 2299 9.4 5.0 4.8 29 0.6 --- --- 
Oak, northern red 1896 6.6 4.8 7.0 210 3.6 9.3 20.5 
Oak, other red 24 0.1 --- --- i 0.1 --- --- 
Oak, other white 45 0.2 22.3 0.0 4 0.1 --- --- 
Oak, white 3575 14.6 2.6 5:5 101 1.9 10.8 19.6 
Pine, white 37 0.2 11.3 0.0 33 0.5 oat 10.4 
All Intermediate 10840 42.8 5.4 5.5 737 13.7 10.2 15.7 
Basswood sp. 203 0.8 5.9 1.9 127 7248) 5.0 2.3 
Beech 949 3.2 5.4 1.3 350 5.0 7.5 5.8) 
Blackgum 179 0.7 0.8 3.5 19 0.4 --- --- 
Buckeye sp. 125 0.5 3.4 0.0 21 0.5 --- --- 
Hemlock sp. 44 0.2 0.0 0.0 101 Wet 24.4 0.0 
Maple, red 1371 6.2 3.3 28 599 12.1 4.0 3.7 
Maple, sugar 696 3.0 2.4 2.8 VEU 14.6 3.0 3:5 
All Tolerant 3567 14.7 3.6 2.1 1944 36.9 5.5 3.6 
Other Species 360 1.7 13.1 1.6 93 1.8 19.6 2.4 
All Species 23936 100 6.5 4.2 5077 100 8.2 6.8 


Highlighted values represent species that comprise at least 5 percent of the composition or with percentages of dead or 
removed trees significantly greater (p < 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 M221A. 




















! epecles Oak-Hickory Northern Hardwood 

oo Cina. Dene eR 
Ash sp. 37 ille/a 0.0 6.6 63 12.0 1.4 3.0 
Aspen sp. 1 0.0 --- — 
Birch sp. 62 26. 25 0.0 37 7.4 13.9 0.0 
Black cherry 37 1.3 0.0 0.0 29 5.6 5.3 D3 
Black locust 166 6.8 8.6 0.0 34 7.0 17.3 8.5 
Black walnut 21 1.0 0.0 0.0 6 ile} --- --- 
Hickory sp. 149 6.2 4.9 oi56) 19 3.8 --- --- 
Oak, scarlet 95 441 TAS) 4.0 
Pine, loblolly 
Pine, pitch 28 ei 8.2 het! 
Pine, shortleaf 
Pine, Virginia ic 0.6 --- --- 1 0.4 ost ae 
Sassafras 8 0.4 ses ve 
Sweetgum 
Yellow-poplar 171 7A 0.0 0.0 9 1.6 --- =o5 
All Intolerant 788 33.1 4.2 1.5 198 38.9 11.3 5.2 
Cucumbertree 45 Nez 3.4 0.0 5 0.8 --- one 
Elm sp. 2 0.1 --- --- 2 0.5 --- --- 
Magnolia sp. 1 0.2 --- --- 
Oak, black 143 5.4 1.4 4.1 2 0.3 --- oe 
Oak, chestnut 528 19.9 1.8 1.5 1 0.2 --- eer 
Oak, northern red 408 14.7 1.9 0.7 31 4.4 6.8 13.1 
Oak, other red 1 0.0 --- --- 
Oak, other white 2 0.1 --- --- 
Oak, white 249 9.3 We 0.9 2 0.3 --- --- 
Pine, white 39 i128} 1.2 0.0 
All Intermediate 1424 52.8 1.8 1.3 43 6.6 6.2 8.8 
Basswood sp. 45 1.6 3.4 0.0 26 4.2 --- --- 
Beech 43 Wt 6.6 0.0 59 12.6 ES 0.0 
Blackgum 14 0.6 --- --- 1 0.1 --- --- 
Buckeye sp. We 0.4 --- --- 1 0.2 --- 
Hemlock sp. 18 0.6 --- --- 16 Pall --- --- 
Maple, red 131 5.9 Zl 0.0 43 9.3 25.5 0.0 
Maple, sugar 63 728) 0.0 0.0 109 20.6 4.6 4.2 
All Tolerant 326 13.3 4.6 0.0 255 49.7 7.8 2.3 
Other Species 19 0.8 --- --- 23 4.8 6.1 0.0 
All Species 2557 100 3.0 ee 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 < 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 M221B. 


























Forest Type 
Species Oak-Hickory Northern Hardwood 
D 
Ash sp. 101 1-0 49 5.8 182 3:9 5.8 5.4 
Aspen sp. 20 0.3 --- --- 14 0.3 ane oee 
Birch sp. 233 2.7 11.7 2.6 436 9.5 9.7 5.1 
Black cherry 360 Shit 4.5 6.1 958 18.1 Zo 3.9 
Black locust 196 2.3 16.9 2.4 98 2.2 16.9 2.2 
Black walnut 32 0.3 8.6 21.9 Uf 0.1 --- === 
Hickory sp. 452 4.9 6.7 0.5 103 2.0 7.9 6.3 
Oak, scarlet 293 2.8 mal 4.6 2 0.0 --- --- 
Pine, loblolly 
Pine, pitch 14 0.1 --- --- 
Pine, shortleaf 
Pine, Virginia 8 0.1 --- --- 
Sassafras 72 0.9 17.0 0.0 4 0.1 --- --- 
Sweetgum 1 0.0 --- coe 
Yellow-poplar 1504 14.1 2.3 4.3 173 3.2 2.6 US 
All Intolerant 3286 33.4 6.2 3.7 1977 39.0 5.8 4.6 
Cucumbertree 130 eS 23 4.9 81 oe 19.4 15.4 
Elm sp. 14 0.1 --- --- 33 0.7 al 0.0 
Magnolia sp. 58 0.6 0.0 0.0 27 0.5 --- --- 
Oak, black 297 2.8 UL 44 22 0.3 --- --- 
Oak, chestnut 1308 13.0 6.0 4.2 40 0.8 26.2 14.4 
Oak, northern red 1996 18.5 4.4 BS 284 5.0 2.8 19.7 
Oak, other red 13 0.1 --- --- 
Oak, other white 1 0.0 --- --- 1 0.0 --- --- 
Oak, white 718 7.0 2.6 2.2 51 1.0 0.0 10.7 
Pine, white 14 0.1 --- --- 
All Intermediate 4549 43.6 4.6 4.4 539 10.0 7.8 15.2 
Basswood sp. 138 1.4 3.9 10.0 263 4.9 0.9 0.2 
Beech 346 2.9 3.4 2.4 631 11.6 9.1 6.8 
Blackgum 97 0.8 2.8 3.8 Ud 0.3 --- --- 
Buckeye sp. 9 0.1 --- === 9 0.1 --- --- 
Hemlock sp. 84 0.7 1.2 4.3 WAS 2.9 4.0 6.4 
Maple, red 1238 13.2 2.4 1.8 752 14.5 1.8 8.4 
Maple, sugar 296 3.0 0.7 3:5 727 13.9 4.9 44 
All Tolerant 2208 22.1 2.3 2.5 2574 48.2 4.5 5.9 
Other Species 81 0.9 9.7 0.0 149 2.8 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 < 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 M221C. 















Forest Type 
























Oak-Hickory Northern Hardwood 

jn= | xe | po [| Rm |[ n=] xe | 
Ash sp. 73 tel ile) lol 30 3.5 6.5 0.0 
Aspen sp. 1 0.0 sen — 
Birch sp. 107 ile, 6.2 0.0 16 2.9 --- === 
Black cherry 25 0.4 --- =e 9 1.1 vo - 
Black locust 132 Zeal 11.7 0.0 11 ites 0.0 0.0 
Black walnut 39 0.6 20.1 8.2 3 0.5 --- sce 
Hickory sp. 558 8.5 6.0 1.6 37 5.9 9.2 0.0 
Oak, scarlet 367 5.5 eS 4.1 2 0.4 Sas _ 
Pine, loblolly 
Pine, pitch 18 0.3 --- seo 
Pine, shortleaf 6 0.1 --- oth 
Pine, Virginia 15 0.3 
Sassafras 58 0.9 14.6 0.0 3 0.6 --- so 
Sweetgum VW 0.4 --- — 
Yellow-poplar 1446 §=21.9 1.8 3.1 56 8.5 3.2 16.7 
Elm sp. 16 0.2 --- --- 1 0.3 --- = 
Magnolia sp. 25 0.5 --- --- 4 0.9 wee tes 
Oak, black 498 6.4 5.0 4.3 6 0.8 --- -- 
Oak, chestnut 965 = 13.0 1.6 ilet 12 2.0 --- sec 
Oak, northern red 562 6.7 3.0 os} 25 Sra cee ails 
Oak, other red 8 0.1 --- --- 
Oak, other white 4 0.1 --- --- 0.4 — af. 
Oak, white Bilal 5.6 2.5 es at --- =e 
Pine, white 
All Intermediate 2618 35.0 2.6 2a 76 11.6 5.0 8.7 
Basswood sp. 220 3.2 0.3 0.2 We 16.8 1.4 5) 
Beech 440 4.8 2.2 0.1 122 16.6 2.9 6.4 
Blackgum 103 1.4 2.2 ile u 0.8 --- --- 
Buckeye sp. 42 0.7 0.0 0.0 7 0.8 --- --- 
Hemlock sp. 65 0.9 6.7 1.0 We PS) --- -- 
Maple, red 355 6.1 2.6 0.6 25 SLI --- --- 
Maple, sugar 207 Sy Ut lei 137% 19.7 0.7 SUS 
All Tolerant 1432 20.1 3.0 0.5 432 60.9 20 3.6 
Other Species 55 1.0 18.4 0.0 8 2.4 se === 
All Species 6967 100 3.8 2.1 683 100 3.8 5.0 


Highlighted values represent species that comprise at least 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-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 221H. 





Forest Type 


Species Oak-Hickory 
%C | %D jn= | *e | %o | %R | 














Ash sp. 109 ie 2.4 2.7 3 0.2 --- --- 
Aspen sp. 4 0.1 --- --- 

Birch sp. AE 0.3 <<- --- 

Black cherry lr 0.1 --- --- 3 0.2 --- --- 
Black locust ae 0.6 45.2 0.0 1 0.1 --- --- 
Black walnut 58 0.6 8.3 0.0 4 0.3 --- --- 
Hickory sp. 1057 #114 8.7 ae 58 5.4 iss) 0.0 
Oak, scarlet 856 7.8 13.7 1.8 80 5.0 15.4 0.0 
Pine, loblolly 4 0.0 --- --- 45 3.6 ie 19.1 
Pine, pitch 45 0.4 18.2 4.2 25 2.0 --- --- 
Pine, shortleaf 167 2.0 13.0 9.6 Zen 18.9 10.7 Sa 
Pine, Virginia 152 1.8 19.2 9.8 253  =20.1 5.0 iO 
Sassafras 24 0.3 --- --- 

Sweetgum 63 0.6 3.4 15.7 2 0.1 --- --- 
Yellow-poplar 1221 11.6 a2 1.9 rad 4.9 2.6 0.0 
All Intolerant 3856 38.5 8.9 3.0 778 60.7 15 2.6 
Cucumbertree 64 0.5 0.0 0.0 1 0.1 --- --- 
Elm sp. 31 0.3 15.3 8.8 4 0.4 --- --- 
Magnolia sp. 15 0.2 --- --- 1 0.2 --- --- 
Cak, black 1119 9.6 7.5 3.4 59 4.0 9.0 0.0 
Oak, chestnut 1527 @313:5 3.5 oo 70 4.2 0.3 3.0 
Oak, northern red 534 4.2 8.3 3.0 28 2.1 --- --- 
Oak, other red 110 Ted 5.8 6.1 67 4.8 4.5 0.0 
Oak, other white 153 fe 11.5 3.1 55 4.2 20.3 3.0 
Oak, white 1558 Get5 3.0 2.8 135 8.6 2.1 0.9 
Pine, white 54 0.4 5.4 7.8 31 2.2 0.0 0.0 
All Intermediate 5165 46.3 49 3.2 451 30.8 5.6 1.1 
Basswood sp. 101 0.9 U5) 3.4 

Beech 557. OD eal 1.4 4 0.1 --- o-- 
Blackgum 211 1.9 Tite O7, 17 Ah --- --- 
Buckeye sp. 29 0.3 -- --- 

Hemlock sp. 83 0.6 0.4 Wa WE 0.9 --- es 
Maple, red 464 4.8 4.4 1.8 7p} 4.9 6.5 0.0 
Maple, sugar 214 1.9 53 25 4 O13 --- oe 
All Tolerant 1659 13.8 5.7 1.8 115 7.4 6.8 0.0 
Other Species 125 1.3 43.6 2.7 15 Aad --- --- 
All Species 10805 100 7.1 2.9 1359 100 7.0 1.9 


Highlighted values represent species that comprise at least 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-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 221L. 





























Forest Type 
Species Oak-Hickory Oak-Pine 

nani Can COCR n= | *C | %D | %R 
Ash sp. 46 ive 4.7 ee 2 1.4 --- oe 
Aspen sp. 
Birch sp. 23 1.0 --- a 
Black cherry 25 ae => aos 2 0.9 --- --- 
Black locust 48 2.6 2 PTh 1,0 
Black walnut 25 0.9 --- --- 1 0.6 --- eee 
Hickory sp. 235 9.4 7.9 0.9 10 4.8 --- --- 
Oak, scarlet 95 Bhd 16.1 11.3 if 3.0 --- --- 
Pine, loblolly 8 4.0 a pe 
Pine, pitch 1 0.0 --- --- 
Pine, shortleaf 17 0.8 --- --- 29 17.3 a 3% 
Pine, Virginia 38 1.6 18.2 2.8 61 32.7 10.4 0.0 
Sassafras 13 1.0 --- --- 
Sweetgum 10 0.4 --- --- 1 0.4 aoe eee 
Yellow-poplar B18 19.4 2.0 3.6 18 6.0 --- --- 
Elm sp. 16 0.5 --- --- 
Magnolia sp. 15 0.8 --- --- 
Oak, black 192 6.0 4.2 2.0 12 4.9 --- --- 
Oak, chestnut 448 13.9 If 0.4 5 2.2 --- ae 
Oak, northern red 202 6.6 4.8 2.0 12 4.5 --- --- 
Oak, other red 36 el 2.3 1.8 3 1.4 --- was 
Oak, other white 12 OnG --- --- 4 1.9 --- --- 
Oak, white 236 8.2 2.6 2.6 28 8.9 --- --- 
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 2a 15.9 
Beech 104 2.8 6.3 0.0 2 0.6 --- --- 
Blackgum 63 1.9 16.9 3.3 1 0.2 --- --- 
Buckeye sp. 14 0.5 --- --- 
Hemlock sp. 24 0:9 --- = 
Maple, red 138 5.2 4.8 0.0 4 1.9 --- --- 
Maple, sugar 98 3.4 23 Dal 1 0.5 --- --- 
All Tolerant 484 16.1 5.3 3.5 8 3.2 -— --- 
Other Species 38 1.4 26.7 0.0 3 1.9 “= --- 
All Species 2796 100 5.2 2.6 215 100 8.0 1.6 


Highlighted values represent species that comprise at least 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-9 


Table C-10 
Percentages of species composition (C), dead trees (D), and removed trees (R) by 
forest type and shade tolerance for ecological subregion 231C. 














Forest Type 
Ash sp. 30 1.2 1.9 0.0 7, 0.7 --- --- 
Aspen sp. 
Birch sp. 5 0.6 a= --- 
Black cherry 7 0.4 --- --- 3 0.3 --- --- 
Black locust 
Black walnut 4 0.1 --- --- 1 0.2 --- --- 
Hickory sp. 327 15.0 1.9 2.5 78 8.4 1.4 1.8 
Oak, scarlet 120 5.3 12.6 1.9 31 2.8 4.5 0.0 
Pine, loblolly 141 5.1 0.0 25.0 270 22.2 0.9 3.6 
Pine, pitch 
Pine, shortleaf 62 3.6 5.9 15.9 61 7.2 5.6 6.6 
Pine, Virginia 5g) 2.7 13.3 32.5 104 13.1 8.7 5.9 
Sassafras 
Sweetgum 107... #45 6.1 18 45 £48 5.6 0.0 
Yellow-poplar 187 6.5 1.3 0.0 52 4.2 0.0 0.5 
All Intolerant 1038 44.3 4.4 7.3 657 64.6 3.8 3.4 
Cucumbertree 1 0.0 --- --- 
Elm sp. ly 0.7 --- --- 5 0.5 --- --- 
Magnolia sp. 1 0.0 --- --- 1 0.1 --- --- 
Oak, black 133 5.3 B59 4.3 30 a 33 3.4 
Oak, chestnut 401 17.1 0.5 0.8 68 6.9 0.0 0.0 
Oak, northern red 84 Oe 0.8 Paks) 22 2.1 --- --- 
Oak, other red 207 8.1 1.6 0.6 71 6.2 7.0 1.8 
Oak, other white 98 4.0 44 ala 45 441 3.4 1.9 
Oak, white 295 11.2 1.6 ez 101 8.9 0.0 0.7 
Pine, white 
All Intermediate 1237 49.7 1.9 1.7 343 31.5 2.3 1.4 
Basswood sp. 2 0.1 --- --- 1 0.1 --- --- 
Beech 47 flea 0.0 7.4 5 0.4 --- --- 
Blackgum 43 2.0 0.7 3.2 if 0.8 --- --- 
Buckeye sp. 
Hemlock sp. 1 0.2 --- --- 
Maple, red 43 1.7 5:5 15 9 1.0 --- --- 
Maple, sugar 7 0.3 =a= --- 
All Tolerant 142 5.2 2.0 3.3 23 2.4 ae es 
Other Species 21 0.8 oe =-- 14 1.5 ee = 
All Species 2438 100 3.1 4.3 1037 100 3.4 2.7 


Highlighted values represent species that comprise at least 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. 


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