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This is to certify that the
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Impacts of Beech Bark Disease on Stand Composition and
Wildlife Resources in Michigan.

presented by

Amy M. Kearney

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2/05 p:ICIRC/DateDue.indd-p.1

IMPACTS OF BEECH BARK DISEASE ON

STAND COMPOSITION AND WILDLIFE RESOURCES IN MICHIGAN.

By

Amy M. Kearney

A THESIS
Submitted to
Michigan State University

in partial fulfillment of the requirements
for the degree of

MASTER OF SCIENCE
Department of Entomology

2006

ABSTRACT

IMPACTS OF BEECH BARK DISEASE ON
STAND COMPOSITION AND WILDLIFE RESOURCES IN MICHIGAN.

By
Amy M. Kearney

Beech bark disease was discovered in Michigan in 2000 and will likely kill at
least 50% of mature American beech trees. Our study was designed to collect benchmark
data that will allow us, in future studies, to determine how understory composition,
wildlife resources, and stand productivity are affected by beech bark disease. We
assessed the potential impact of beech bark disease in forest stands with varying beech
densities (low, moderate, and high) and infestation of Cryptococcusfagisuga (absent,
light, and heavy). Variables were measured in 62 stands throughout the Upper and
Lower Peninsulas of Michigan in 2002 and 2003. Regenerating beech stem density did
not differ among stands and no current evidence suggests dense beech thickets will
regenerate in stands impacted by beech bark disease in Michigan. Beech snags were
significantly more abundant in stands with heavy C. fagisuga infestation than in stands
where C. fagisuga was absent or infestation was light, suggesting tree mortality from
beech bark disease had occurred. Coarse woody debris volume and beech nut mass
appeared to be highest in stands with heavy C. fagisuga infestation, although means were
not statistically different. Long-term monitoring of these stands will be necessary to

document the effects of beech bark disease as it progresses throughout Michigan.

Dedicated to
Walter Bartow and
Genevieve Hermann,
who taught the essentials.

iii

ACKNOWLEDGEMENTS

The research presented here has been a collaborative effort of many individuals.
Their contributions of expertise, field assistance, laboratory equipment, data, and support
made this project possible, for which I owe great thanks.

Deborah G. McCullough provides tremendous support to all of her students, yet it
is her unparalleled work ethic and passion for forest entomology that distinguishes her as
an outstanding adviser. I have been fortunate to develop my academic and professional
skills under her guidance. I am also grateful to Michael Walters and Therese Poland for
serving on my advising committee and contributing valuable, diverse perspectives on this
research.

With utmost gratitude, I thank William Kearney and Ivan Locke for providing
field assistance during winter leaf litter collections, a particularly cold and unpleasant
task. They also volunteered a great deal of woodworking and construction expertise to
the hardware used in this research. Above all, their encouragement and sense of humor
has been the greatest contribution to this project.

I had the assistance of two exceptional undergraduate student workers, Daniel
Armstrong and Christie Sampson. Their dedication to the research was demonstrated in
their diligence, accuracy, and good nature despite the ofien challenging work
environment. I also appreciate the field and laboratory assistance of Emily Carlson,
Deric Grain, Shannon Hannah, Laura Lazarus, Sarah C. Matteo, Lance J. Roberts, Abby

Sommers, Zsofia Szendrei, and Melanie DePoy. I was fortunate to share a lab with

iv

talented colleagues who offered valuable insight and assistance. Many thanks to Kirsten
Fondren, Nathan Seigert, Laura Lazarus, Andrew Klein, and Andrea Agius.

In addition to valuable discussions on forest ecology, Bruce Walker generously
offered access to his cabin in the Upper Peninsula, which was located conveniently near
many of our research stands. Paul Bloese and Randy Klavickas provided the drying oven
and workspace at the Tree Research Center on MSU’s main campus and kindly allowed
me to monopolize the oven schedule. Ray Miller and personnel at MSU Upper Peninsula
Tree Improvement Center in Escanaba, MI provided computer hardware and
WinDendro® for tree ring analyses. I specifically thank Brad Bender for his trouble-
shooting efforts and ultimately helping me acquire necessary data. Carol Graysmith
offered indispensable technical support in operating WinDendro® at the main campus.
Upon the insight and suggestion of William Kearney, Lesco, Inc. donated seed bags that
enhanced the quality and sanitary storage of leaf litter samples.

Research was conducted in conjunction with John Witter at the University of
Michigan, with particular thanks to Holly Petrillo and Erin Thompson for providing maps
and stand-level data. We received support and access to research stands from Michigan
Department of Natural Resources, Hiawatha Sportsmen’s Club, Shelter Bay, Longyear
Corporation, Pictured Rocks National Lakeshore, and Seney Wildlife Refuge. This

project was funded by a grant from the Michigan Department of Natural Resources.

TABLE OF CONTENTS

LIST OF TABLES ............................................................................................................. vii
LIST OF FIGURES ............................................................................................................ ix
INTRODUCTION ............................................................................................................... 1

CHAPTER 1. SPECIES COMPOSITION AND REGENERATION IN
HARDWOOD FORESTS OF MICHIGAN IMPACTED BY BEECH
BARK DISEASE ................................................................................................................. 5

CHAPTER 2. IMPACT OF VARYING STAGES OF BEECH BARK
DISEASE ON WILDLIFE RESOURCES IN MICHIGAN .............................................. 31

CHAPTER 3. RADIAL GROWTH AND LEAF BIOMASS OF
MICHIGAN BEECH FORESTS IMPACTED BY VARYING LEVELS

OF BEECH BARK DISEASE ........................................................................................... 54
APPENDICES ................................................................................................................... 75
BIBLIOGRAPHY ............................................................................................................ 102

vi

LIST OF TABLES

Table 1-1. Mean basal area (d: SE) of overstory species composition in stands with low,
moderate, and high beech density and absent, light, and heavy C. fagisuga infestation.
Means in rows (a-c) and columns (y-z) followed by different letters are significantly
different (p<0.05) ............................................................................................................... 20

Table 1-2. Spearman's correlation coefficients (rs) and significance value (p) for basal
area (mz/ha) among different overstory of 62 stands. Correlation coefficients with p<0.05
are significantly different ................................................................................................... 21

Table 1-3. Spearman's correlation coefficients (rs) and significance values (p) for basal
area (mz/ha) among overstory species and corresponding species of seedlings, saplings,
and recruits in 62 stands. Correlation coefficients with p<0.05 are significantly

different. ............................................................................................................................. 22

Table 1-4. Mean number of stems (i: SE) per m2 of American beech (Fagus grandifolia),
sugar maple (Acer saccharum), other hardwoods, conifers, and all species of seedlings,
saplings, and recruits in and recruits in stands with low, moderate, and high beech density
and absent, light, and heavy C. fagisuga infestation. Means followed by different letters
among rows are significantly different (p<0.05). .............................................................. 23

Table 2-1. Basal area, mean diameter, and number of trees examined and number of
cavities of three size classes by tree species) ..................................................................... 48

Table 2-2. Volume of coarse woody debris (m3/ha fl: SE) of 5 decay classes among stands
with low, moderate, and high beech density and absent, light, and heavy C. fagisuga
infestation ........................................................................................................................... 49

Table 2-3. Mean number (:l: SE) and mean basal area (mz/ ha 3: SE) of beech snags and

snags of all species among stands with low, moderate, and high beech density and among
stands with absent, light, and heavy C. fagisuga infestation. Means followed by different
letters among rows (a-b) and among columns (y-z) are significantly different ................. 50

Table 3-]. Mean annual increment ratios (mmzmm) for 5, 10, and 15-year intervals for
beech (n=304) and non-beech (n=62) trees among stands with absent, light, and heavy C.
fagisuga infestation and among stands with low, moderate, high beech density .............. 67

Table 3-2. Mean leaf mass (g/mz) of American beech (F agus grandifolia), sugar maple
(Acer saccharum), red maple (Acer rubrum), red oak (Quercus rubra), and other

hardwood species among stands with absent, light, or heavy C. fagisuga infestation ...... 68

Table 3-3. Spearman's correlation coefficients (rs) and significance value (p) for the
relationship between basal area (mz/ha) of overstory species and corresponding leaf

vii

biomass from 40 stands in 2002 and 54 stands in 2003. Correlation coefficients with
p<0.05 are significantly different ....................................................................................... 70

viii

LIST OF FIGURES

Figure 1-1. Distribution of beech bark disease research sites and Cryptococcusfagisuga
infestation indices throughout Michigan, 2003 ................................................................. 25

Figure 1-2. Intensive research plot design indicating arrangement of subplots 1-5,
overstory composition transects, and regeneration plots ................................................... 26

Figure 1-3. Number of stands with absent, light, and heavy Cryptococcusfagisuga
abundance among stands with low, moderate, and high beech densities .......................... 27

Figure 1-4. Interactive effects between beech density classification and C. fagisuga
infestation for basal area of overstory a) beech, b) sugar maple, and c) red maple ........... 28

Figure 1-5. Interactive effects between beech density classification and C. fagisuga
infestation for sugar maple a) seedlings, b) saplings, and c) recruits ............................... 29

Figure 1-6. Interactive effects between beech density classification and C. fagisuga
infestation for red maple a) seedlings, b) saplings, and c) recruits ................................... 30

Figure 2-1. Intensive research plot design indicating arrangement of subplots 1-5,
individual beech trees selected for long-term monitoring, beech nut collection traps, and
coarse woody debris and snag measurement transects ...................................................... 52

Figure 2-2. Beech nut mass (g/m2 i SE) among stands with low, moderate, and high
beech density and absent, light, and heavy C. fagisuga infestation in a) 2002 and
b)2003 ............................................................................................................................... 53

Figure 3-1. Intensive research plot design indicating arrangement of subplots 1-5,
overstory composition measurement transects, individual beech trees selected for long-
terrn monitoring, and leaf litter trap ................................................................................... 72

Figure 3-2. Age of beech trees (n=94) plotted against the ratio of mean annual increment
for intervals of a) 5-year, b) 10-year, and c) lS-years ....................................................... 73

ix

Figure 3-3. Composition of leaf biomass (g/mz) by species from a) 40 stands in of stands
in 2002 and from b) 54 stands in 2003 .............................................................................. 74

INTRODUCTION

Throughout history, beech trees (Fagus sp.) have supplied humans with a variety
of resources. Early Europeans and North American settlers used lumber for household
furnishings and water wheels, fed beech nuts to domestic hogs and turkeys, and stuffed
mattresses with dry, elastic beech leaves (Sand 1991, Tubbs and Houston 1990). Beech
wood imparts no taste and was a favored material for food containers and utensils, such
as ice cream paddles, cutting boards, and beer casks (Sand 1991). The dense wood
continues to be valued for firewood and harvested for pulp and lumber production (Peters
1997, Tubbs and Houston 1990).

American beech, Fagus grandifolia, primarily occurs in structurally diverse
stands of mixed species. Although previously spanning the Atlantic to Pacific coasts of
North America, the post-glacial range of American beech is delimited in the north by
Nova Scotia and the Upper Peninsula of Michigan, west by Indiana, Wisconsin and
eastern Texas, and south by northern Florida (Little 1971). Annual precipitation across
this area ranges from 760-1270 mm and mean annual temperatures vary from 4°—21° C
(40°-70° F), with lows of -42° C (-44° F) and highs of 38° C (100° F) (Tubbs and
Houston 1990). Beech stands are typically found on loose, dry mesic loams with a pH of
4.1-6.0 (Tubbs and Houston 1990).

Beech is a shade tolerant species that regenerates by both seed and root-sprouting
(Ward 1961). Mature beech greater than 40 years of age produce periodic crops of beech
nuts every two to three years, while large-scale mast events occur in summers following a
drought year (Tubbs and Houston 1990, Piovsean and Adams 2001). Successful

germination of beech nuts is highly variable, ranging from 0-81% (Stalter 1982). In

northern regions, or where site conditions are poor, regeneration occurs primarily by root-
suckering (Graber 1992, Held 1983). Injury to trees or roots can stimulate trees to sprout
vigorous root-suckers, although these do not generally form desirable trees (Tubbs and
Houston 1990, Ward 1961). Beech grows in cycles of suppression and release, surviving
well in a shaded understory but not responding rapidly to sudden gaps (Kobe 1995,
Tubbs and Houston 1990). Beech trees are not preferentially browsed by deer, giving
them a competitive advantage in stands heavily impacted by herbivory (Kelty and
Nyland, 1981).

The bark of beech trees is composed of only a few layers of cork and is easily
penetrated by insects (Burns and Houston 1987). The beech scale insect, Cryptococcus
fagisuga Lind. (Homoptera:Cryptococcidae), has a feeding stylet of only 2 mm, which
cannot penetrate the relatively thin bark, but is instead used to feed on the bark
parenchyrna (Dubeler 1997, Shigo 1972). The scale insects feed on the sap in the inner
bark of beech, creating minute wounds that provide entry courts for at least one of three
Nectria fungus species, Neonectria galligena A., Bionectria ochroleuca A., or the exotic
Neonectria coccinea var. faginata A; resulting in the beech bark disease complex
(Houston and O’Brien 1994). The pathogen progressively kills areas of tissue and
eventually girdles the limbs and trunk of the tree. Eventually, secondary factors cause
structural weakening and trees often break in high winds during a condition known as
“beech snap” (Houston and O’Brien 1994).

Cryptococcusfagisuga disperses ahead of the Nectria spp., transported primarily
by wind currents, during a phase called the “Advancing Front” and typically colonizes

areas of rough bark on the trunks of large, old trees (Houston 1975, Mielke 1982).

Generally within a few years, Nectria sp. invades scale-infested trees and completes the
beech bark disease etiologic complex (Ehrlich 1934). Tree impact and mortality is high
during this phase known as the “Killing Front”. Mature trees that are not outright killed
by beech bark disease exhibit a substantial growth reduction (Twery and Patterson 1984).
Declining trees often produce dense root-sprouts that are genetically identical to the
parent trees and equally susceptible to beech bark disease (Houston 1975). In
northeastern stands impacted by beech bark disease, understory vegetation shifted to a
strongly dominant beech composition of both seed and sprout origin (Hane 2003). Trees
originating from root sprouts are generally stunted or deformed, contributing to the
characteristic “Aftermath” forest structure that has replaced much of the original beech
component of the northeastern United States (Houston 1994, Houston and Valentine
1987, Ostrofsky and McCormack 1986).

Beech bark disease first arrived in North America in Nova Scotia in 1890 on
infected nursery stock from Europe and has since spread to Maine, New Hampshire,
Vermont, New York, Massachusetts, Pennsylvania, Ohio, Tennessee, West Virginia,
Virginia, North Carolina, and Ontario (Houston and O’Brien 1994). The immature
crawlers are the mobile form of the insect and are transported primarily by wind, but
birds may also transport them over short distances or along migratory routes (Houston
and O’Brien 1994, McCullough et al. 2001). Beech bark disease infestations that occur
in close proximity to campgrounds suggests that firewood infested with crawlers serves
as another means of transporting C. fagisuga long distances.

Beech bark disease was discovered near camping areas in Mason and Luce

Counties in Michigan in 2000 and now infests at least two regions, one in the Upper

Peninsula and another in the Lower Peninsula (McCullough et a1. 2001)(Figure 1-1).
Both infestations were estimated to be at least 10 years old, based on observations by
Michigan Department of Natural Resources (MDNR) personnel and local residents
(O’Brien et al. 2001). The relatively recent occurrence of beech bark disease in Michigan
offers a unique opportunity to quantify characteristics of a forest stand before and after
infestation. We designed a study to monitor beech bark disease as it spreads in range and
intensity throughout Michigan. We established permanent plots to quantify current
conditions of stands including overstory composition and species regeneration,
availability of wildlife resources, radial growth, and leaf biomass production. Conditions
were compared among stands grouped by beech density and levels of C. fagisuga.
Although we are specifically addressing the impacts of beech bark disease, the broader
scope of this research will enhance our understanding of how non-native organisms
impact forested ecosystems.

The potential impact of beech bark disease is examined in three chapters; Current
stand composition and future regeneration are presented in Chapter One, Chapter Two
discusses wildlife resources, and Chapter Three focuses on stand—level productivity.
Details specific to the stands sampled in this study are detailed in Appendix One and
include location, beech density, and C. fagisuga infestation at the time of the study.
Occurrence of overstory and regenerating species are listed for each stand in Appendix
Two. Appendix three details stand stocking characteristics along with known
management histories. Individual tree measurements are listed in Appendix four and
include diameter, height, crown vigor, age, and radial growth trends. Data presented in

these appendices will be essential background measurements for future comparisons.

SPECIES COMPOSITION AND REGENERATION IN HARDWOOD FORESTS OF

MICHIGAN IMPACTED BY BEECH BARK DISEASE.

Introduction

Forests cover 19.3 million acres of the Michigan landscape over two distinct
landmasses, the Upper and Lower Peninsulas (Dickmann and Leefers 2003). The mesic-
deciduous forest type, also known as northern hardwoods, is most prevalent and is
comprised of beech (Fagus sp.), maple (Acer sp.), birch (Betula sp.), and hemlock (T suga
sp.) (Dickmann and Leefers 2003, Woods and Davis 1989). American beech (Fagus
grandifolia Ehrh.) is typically found in the beech-maple forest type dominant in
Michigan and typically occurs in association with sugar maple (Acer saccharum Marsh),
red maple (Acer rubrum L.), eastern hemlock (T suga canadensis), and northern red oak
(Quercus rubrum L.) (Tubbs and Houston 1990).

As mature beech trees are impacted by beech bark disease, the composition and
structure of the overstory may shift. Species composition may be further impacted by
beech bark disease if declining, mature trees produce root sprouts and create dense beech
thickets, subsequently limiting regeneration of other species (Hane 2003, Houston 1975,
Houston and Valentine 1987, Jenkins 1997, Twery and Patterson 1984).

We designed a study to address the impacts of beech bark disease on overstory
species composition and composition of regenerating vegetation in Michigan forests.
Our primary objective was to quantify differences in overstory and understory
composition among stands with varying levels of C. fagisuga infestation and among
stands with varying densities of beech. These data will be used to 1) examine the current
impact of C. fagisuga on species composition, 2) determine future composition of stands
impacted by beech bark disease, and 3) serve as a baseline for long-term monitoring and

firture comparisons as beech bark disease spreads and intensifies throughout Michigan.

Methods
Study Sites

Our research was conducted in the Upper and Lower Peninsulas of Michigan,
with study sites located within the infestation, along the periphery, and in areas not yet
impacted by the disease. During the time of this study in 2002-2003, known infestations
of C. fagisuga in Michigan occurred in only five counties (Chippewa, Manistee, Mason,
Luce, and Oceana) (Figure 1-1). Data were collected between May 2002 and December
2003 in 34 hardwood stands in the eastern Upper Peninsula and 28 stands in the western
Lower Peninsula of Michigan (62 stands total) (Appendix 1). In addition to beech,
overstory composition in our study stands included sugar maple, red maple, eastern
hemlock, and red oak, with lesser amounts of white ash (F raxinus americana), basswood
(T ilia americana), black cherry (Prunus serotina), birch sp., and aspen sp. (Populus
sp.)(Appendix 2). Soils in our study sites were primarily sands or sandy loams and
occurred on moraine, glacial outwash, lakebeds, or sand dunes (Veatch 1953).
Widespread logging occurred in Michigan from approximately 1840 to 1900 and stands
in our study have had variable harvest activities since (Appendix 3). We aimed to select
stands with no harvest activity since 1998 but were limited in some cases by the
availability of accurate management records.

Cooperators from the Michigan Department of Natural Resources (MDNR) and
the University of Michigan initially selected stands as part of the long-term Michigan

Beech Bark Disease Monitoring and Impact Analysis System (Thompson and Witter

2003). Stands were selected to represent three levels of beech stocking and three levels
of beech bark disease infestation.

We assigned classification indices to each stand based on beech basal area and C.
fagisuga infestation (Appendix 1). Beech density was classified for each stand based on
basal area data provided by University of Michigan. Stands were classified as low (<9
m2/ha), moderate (9-18 mZ/ha), or high (>18 mtha) beech density. Scale density was
classified by visually estimating C. fagisuga abundance on beech trees located within the
plot boundary. Stands were classified according to the percentage of beech trees having
greater than 50% coverage of C. fagisuga on at least one side of the main bole and
assigned to the following categories: absent (no infested trees), light (less than 50% of
beech trees infested), or heavy (greater than 50% of beech trees infested).

In each stand, University of Michigan field staff established five circular subplots
14.68 m diameter, one at center and one at 18.28 m from center in each of four azimuths
(Figure 1-3). Each subplot was marked with a permanent metal turf stake and geographic

positioning coordinates were recorded from the center subplot.

Overstory composition

Overstory species composition was measured in each stand along two
perpendicular transects (51 m long) established from subplot 2 to subplots 5 and 3
(Figure 1-3). Species and diameter at breast height, dbh, (1.3 m) were recorded for all

live trees > 24.5 cm dbh within 5 m of each transect.

Tree regeneration

Regeneration plots were established beyond the subplots to minimize the effects
of trampling likely to occur in the subplots. Four regeneration plots were established at
18 m from the center point of the center subplot, in each cardinal direction (Figure 1-3).
From the center of each regeneration plot, we identified number and species of seedlings
(< 30.5 cm tall) within a 2.4 m radius, saplings (> 30.5 cm tall and < 2.5 cm dbh) within a

3.5 m radius, and recruits (> 2.5 cm dbh and < 12.5 cm dbh) within a 7.3 m radius.

Statistical analyses

Normality was tested for all variables using the Shapiro-Wilk test (SAS Institute,
1985). Prior to analyses, mean number of stems per hectare of beech and mean number
of stems per hectare of all species combined were square root transformed to normalize
distribution. (N on-transformed means are referenced in tables.) Two-way ANOVA
[proc glm] was used to test the differences in mean number of stems among stands with
low, moderate, or high beech density and absent, light, or heavy C. fagisuga infestation
(SAS Institute 1985). When results of ANOVA were significant (or = 0.05), Tukey’s
Honestly Significant Difference was used to compare means (Tukey 1977).
Transformations did not normalize data distributions for mean basal area of overstory
trees (each species), number of stems of sugar maple, red maple, other hardwoods (red
oak, ash, aspen, basswood, birch, and black cherry), and conifers (eastern hemlock,
balsam fir, and white pine) for each stage of regeneration, and species richness, therefore,
these variables were ranked and analyzed with nonparametric tests. The nonparametric
Kruskal-Wallis test [proc rank] was used to determine if these variables differed among

stands with low, moderate, or high beech density and absent, light, or heavy C. fagisuga

infestation (SAS Institute 1985). For significant tests ((1 =0.05), means were compared
with nonparametric Tukey’s-type multiple comparison (Zar 1984). Spearrnan’s
nonparametric correlation [proc corr spearman] was used to evaluate the linear
relationship between basal area and regeneration of seedlings, saplings, and recruits of
selected species and both the correlation coefficient (rs) and significance value (p) are
reported (SAS Institute 1985). All analyses were conducted using SAS 9.1, except for
the nonparametric Tukey’s-type multiple comparison which was conducted with
Microsoft Excel software (Redmond, WA).

Root injury during harvest activity has been shown to stimulate sprouting in beech
(Fowells 1965a). Although we attempted to select stands with no recent harvest activity,
we later found that two stands (stand ID 121 and 166) had been selectively harvested
between 1998 and 2000. To account for possibly confounding impacts of management
practices on species regeneration, we analyzed data with and without these two stands.
Inclusion of these two stands had no impact on results of analyses or our conclusions and

were retained.

Results

Cryptococcusfagisuga was absent in 18 stands with low beech density, 16 stands
with moderate beech density, and five stands with high beech density (39 stands total)
(Figure 1-3). Light C. fagisuga infestation was present in five stands with low beech
density, five stands with moderate beech density, and two stands with high beech density

(12 stands total). Heavy C. fagisuga infestation was present in two stands with low beech

10

density, three stands with moderate beech density, and six stands with high beech density

(1 1 stands total).

Overstory composition

Beech and sugar maple were the most prevalent species in the overstory, although
neither species was encountered in our transects in all 62 stands (Appendix 2). Beech
was present in 52 stands with basal area ranging from 1.39 to 39.04 mZ/ha. Sugar maple
was present in 50 stands with basal area ranging from 1.74 to 29.88 mZ/ha. Red maple
was also a common associate, present in 25 stands with basal area ranging from 1.43 to
15.65 mz/ha (Table 1-1). Aspen was only present in five stands yet comprised the
greatest basal area of any species with a maximum of 39.78 m2/ha in any one stand.
Eastern hemlock was encountered in 16 stands; only five of which were in the Lower
Peninsula. Red oak, white ash, basswood, black cherry, and birch sp. were also present,
but each species occurred in fewer than 20 stands. Balsam fir was encountered in a
single stand and white pine was not encountered in the overstory of any stands.

Basal area of overstory beech was not significantly different among stands with
absent, light, or heavy C. fagisuga infestation (p=0.060) (Table 1-1). There was
significantly greater beech basal area in stands classified as moderate beech density than
in stands classified as low beech density (dfi-2,59; F =19.05, p<0.001). The interactive
effect between beech density classification and C. fagisuga infestation was significant for
overstory beech (dfi4,57; F =10.65, p<0.001) (Figure 1-4). Basal area of overstory
beech was not significantly correlated with basal area of sugar maple, red maple, red oak,

other hardwoods, or conifers in the overstory (Table 1-2). Basal area of beech in the

11

overstory was not significantly correlated with mean number of beech saplings,
seedlings, or recruits (Table 1-3).

Basal area of overstory sugar maple, red maple, red oak, assorted hardwoods, and
conifers were not significantly affected by beech density (Table 1-1). Basal area of red
oak, assorted hardwoods, and conifers were not significantly affected by C. fagisuga
infestation while basal area of overstory sugar maple and red maple were significantly
affected by C. fagisuga infestation (Table 1-1). In stands with heavy C. fagisuga
infestation compared to stands with no C. fagisuga infestation, basal area of overstory
sugar maple was lower (dfi2,59;, F = 1.60, p<0.001) and overstory red maple was higher
(df=2,59; F = 7.33, p=0.001). Basal area of sugar maple in the overstory was inversely
correlated with basal area of red maple, red oak, and other hardwoods in the overstory
(Table 1-2). Basal area of overstory sugar maple and red maple were significantly
correlated with corresponding species regeneration, except regeneration of red maple
saplings (Table 1- 3). The interaction between beech density and C. fagisuga infestation
was significant for overstory sugar maple (dfi—4,57; F = 6.01 , p<0.001) overstory red

maple (($457; F=5.23, p=0.001) (Figure 14).

Tree regeneration

In the 11 stands with heavy C. fagisuga infestation, seedling regeneration was
entirely absent in two stands and another stand had only 2,073 stems per ha, compared
with 54,415 i 15,175.8 stems per ha for the remaining eight stands. Seedling and sapling
regeneration was present but highly variable in all 51 stands with absent or light C.

fagisuga. Number of seedlings ranged from 138 to 733,522 stems per ha and number of

12

saplings ranged from 129 to 52,455 stems per ha in these stands. Sapling regeneration
was entirely absent in four of the 11 stands, with an average of 12,569 d: 3,260.5 stems
per ha for the remaining seven stands.

Beech seedlings were encountered in 95%, saplings in 90%, and recruits in 89%
of all stands (Appendix 2). There were no significant differences in mean stems per ha of
beech seedlings, saplings, or recruits among stands with low, moderate, or high beech
density (seedlings: p=0.108) (saplings: p=0.681) (recruits: p=0.075) nor among stands
with absent, light, or heavy C. fagisuga infestation (seedlings: p=0.565) (saplings:
p=0.192) (recruits: p=0.593) (Table 1-4). The interaction between beech density
classification and C. fagisuga infestation was not significant for beech seedlings
(p=0.303), beech saplings (p=0.575), or beech recruits (p=0.415).

Sugar maple seedlings were encountered in 79%, saplings in 66%, and recruits in
80% of all stands (Appendix 2). Red maple seedlings were encountered in 60%, saplings
in 27%, and recruits in 32% of all stands (Appendix 2). Other hardwood species
encountered in regeneration plots included striped maple (Acer pensylvanicum), birch sp.,
aspen sp., black cherry, ash sp., red oak, American basswood, ironwood, serviceberry
(Amelanchier arborea), common witch-hazel (Hamamelis virginiana), and sassafras
(Sassafras albidum). Striped maple stems were most prevalent and represented 53%,
34%, and 30% of all assorted hardwood seedling, sapling, and recruit regeneration
respectively. Regenerating conifer species included eastern hemlock, balsam fir, and
white pine, collectively comprising less than 3.9% of total seedling, sapling, and recruit
regeneration of all stands combined. Maximum stand densities of all conifer species

combined were 599 seedlings/ha, 2455 saplings/ha, and 134 recruits/ha.

l3

Regeneration of sugar maple, red maple, other hardwoods, and conifers was not
significantly affected by beech density, except for sugar maple recruits, which were
significantly more abundant in stands with low beech density than in stands with
moderate beech density (off—2,59; F =3.99, p=0.024) (Table 1-4). Regeneration of sugar
maple and red maple stems was significantly affected by C. fagisuga infestation (Table 1-
4). Significantly fewer sugar maple seedlings, saplings, and recruits occurred in stands
with heavy C. fagisuga infestation than in stands without C. fagisuga infestation
(seedlings: df—=2,59; F =7.61, p=0.001) (saplings: djE2,59; F :62], p=0.004) (recruits:
df=2,59; F =6.84, p=0.002). The interaction between beech density classification and C.
fagisuga infestation was significant for sugar maple seedlings (dfi4,57; F =3.80,
p=0.008), saplings (dj-L—4,57; F =3.06, p=0.024), and recruits (df=4,57; F =5.17, p=0.001)
(Figure 1-5). Red maple sapling and recruit stems were significantly greater in stands
with heavy C. fagisuga infestation than in stands with no C. fagisuga infestation
(saplings: df=2,59; F =4.28, p=0.01 8) (recruits: dfi2,59; F = 26.39, p<0.001). The
interaction between beech density classification and C. fagisuga infestation were
significant for red maple saplings (df=4,57; F =2.5 5, p=0.049) and red maple recruits
(df=4,57; F =16.39, p<0.001) (Figure 1-6). Regeneration of red oak, other hardwoods,
and conifers were not significantly affected by C. fagisuga infestation (Table 1-4). Red
oak regeneration was not encountered in any of the 62 stands surveyed.

On average, species richness of understory regeneration was relatively low.
There were 3.2 :1: 0.02 seedling species, 2.6 at 0.02 sapling species, and 2.5 :1: 0.01 recruit
species per stand. Mean number of seedling, sapling, or recruit species did not differ

among stands with absent, light, or heavy C. fagisuga infestation (seedlings: p=0.163)

14

(saplings: p=0.564) (recruits: p=0.3 53) nor among stands with low, moderate, or high

beech density (seedlings: p=0.830) (saplings: p=0.792) (recruits: p=0.179).

Discussion
Overstory composition

In all but one of the stands, a maple species, either sugar or red, was also present
in the overstory. In the northeastern United States, beech bark disease killed 85% of the
overstory beech and created gradual gaps (Krasny 1992). If beech bark disease were to
progress through our study area with a similar effect on mature beech, we could expect
sugar maple and red maple to dominate the forest structure following the “Killing Front”.
The formation of gaps in the overstory may also enhance the growth of sugar maple,
which responds to rapidly to sudden gaps by increasing incremental and terminal growth
(Canham 1988). Both species of maple were present at similar levels in the overstory of
stands with varying densities of beech but their basal area was inversely related to each
other. This indicates one or the other species of maple will likely dominate stands
impacted by beech bark disease.

Factors that predispose a stand to beech bark disease are uncertain but likely
include density and composition of overstory species (Twery and Petterson 1984). The
presence of eastern hemlock in a stand may enhance shading and moisture retention,
conditions which have been correlated with C. fagisuga colonization and survival (T wery
and Patterson 1984). In a New England study, stands impacted by beech bark disease
had a greater component of eastern hemlock in the overstory prior to infection than stands

that were not impacted by beech bark disease (Twery and Patterson 1984). Results from

15

our study do not support this association; the basal area of eastern hemlock in the
overstory was relatively low and similar among stands with absent, light, and heavy C.
fagisuga infestations.

In our study, the presence of C. fagisuga was not related to the density of beech in
the stand, contrary to other studies where beech mortality was correlated with beech basal
area (Runkle 1990). During the time of our study, the distribution of C. fagisuga was
geographically limited and therefore, the actual impact of stand composition on beech
bark disease susceptibility may not yet be realized. As C. fagisuga spreads to additional
stands with varying overstory components, comparisons could be made among stands

with varying mortality to determine characteristics of the most susceptible stands.

Tree regeneration

Mature beech trees that are diseased or growing in stressful conditions are known
to generate root-suckers and create dense thickets (Houston 1975). Studies conducted in
northeastern stands impacted by beech bark disease documented increases in beech
regeneration in the “Aftermath” forest (Hane 2003, Houston 1994, Houston and
Valentine 1987, Ostrofsky 1986). Our results, however, suggest that beech bark disease
has not yet altered the overall abundance of beech seedlings, saplings, or recruits in
Michigan forests. Although beech bark disease was evident in at least three stands with
heavy C. fagisuga infestation, as indicated by “beech snap” and beech tree mortality, the
disease complex has not yet progressed to the “Aftermath” stage. Future comparisons of
beech regeneration with baseline data will account for potential lag time in the response

of understory regeneration. There are documented instances in the northeast, however, in

16

which density of regenerating beech did not change despite impacts of beech bark disease
(Twery and Patterson 1984).

The overstory basal area of beech in our stands was not correlated with beech
regeneration, suggesting that beech regeneration may not be directly impacted by
mortality of beech in the overstory. Results from our study demonstrated no reduction in
beech regeneration in stands that had been impacted by C. fagisuga. A similar study
conducted in Wisconsin found no correlation between beech basal area of overstory
beech and beech seedlings (Ward 1961).

Sugar maple was the most common associate of beech in our stands. Like beech,
it is shade-tolerant and capable of responding to overstory gaps with rapid growth
(Canham 1985, Pulson and Platt 1996). Declines in sugar maple regeneration
corresponding to increased beech regeneration have been documented for northeastern
forests impacted by beech bark disease (Hane 2003, Jenkins 1997, Houston 1975). Sugar
maple seedlings decline when competing with dense beech for understory light resources
(Hane 2003). Competition for underground resources and beech leaf leachate have also
been also shown to reduce sugar maple regeneration (Hane 2003, Hane et. a1. 2003,
Huntley et.al. 1989). In our stands, sugar maple stems were less abundant in the
understory of stands with heavy C. fagisuga. Beech regeneration, however, was not
related to sugar maple sapling abundance, suggesting direct competition was not the
limiting factor. Rather, the low density of sugar maple saplings was likely due to the
lower component, relative to beech, of sugar maple basal area in the overstory of stands
with heavy C. fagisuga. Sugar maple regeneration was directly correlated with overstory

composition and therefore, it is likely that future impacts of beech bark disease would

17

increase sugar maple regeneration more dramatically in areas with higher sugar maple
canopy basal area.

Red maple is often an early colonizer but may be dominated by sugar maple after
approximately 80 years in northern hardwood stands (Fowells 1965b). Red maple is a
vigorous sprouter but also regenerates from the light, wind—disseminated seeds that are
produced annually. In our stands, regeneration of red maple was highest in stands with
heavy C. fagisuga infestation, which is likely attributed to the greater basal area of red
maple in the overstory of stands with heavy C. fagisuga infestation than in stands with
light to no C. fagisuga. As the canopies of mature beech impacted by beech bark disease
become sparse, shade-intolerant red maple seedlings are expected to benefit from
increased light levels on the forest floor. No current evidence suggests that abundant
beech regeneration will compete with red maple in the understory and consequently, a
compositional shift to a increased dominance by red maple may occur.

In addition to beech bark disease, other factors likely affected regeneration in the
seven stands we studied in Mason County (Appendix 1). Regardless of C. fagisuga
infestation, stands in these counties had dramatically fewer stems in the understory than
stands sampled in other counties (Appendix 2). Three of the stands were in close
proximity to a popular campground and regeneration may have been limited by heavy
foot traffic. Nonetheless, other stands in our study, including some with heavy C.
fagisuga, were located near campgrounds with presumably similar public use, yet
regeneration was present. Stand-level factors inhibiting regeneration in these counties are
not conclusively explained by beech bark disease or proximity to campgrounds and

warrant further investigation.

18

The data presented in this study represent the conditions of beech in Michigan
forests as of 2002-2003 but are largely intended as a baseline for future comparisons. As
the range of beech bark disease expands and additional stands are exposed, the
significance of overstory composition as a predisposing factor to the severity of infection
will be determined. Likewise, the effects of beech bark disease on firture species
composition will require that stands be revisited to assess regeneration over the long-

term.

19

Table 1-1. Mean basal area (i SE) of overstory species composition in stands with
low, moderate, and high beech density and absent, light, and heavy C. fagisuga
infestation. Means in rows (a-c) and columns (y-z) followed by different letters are

significantly different (_p<0.05).

 

 

C. fagisuga Beech density
infestation low moderate high all stands
American beech
Fagus grandifolia
absent 3.70 :1: 1.056 9.41 i: 1.468 18.78 i 6.237 7.97 :1: 1.320y
light 8.19 :1: 3.699 13.01 i 2.421 18.66 i 7.604 11.95 :1: 2.258y
heavy 3.14 :1: 3.141 8.67 d: 3.410 19.36 i 3.129 13.50 i: 2.819y
all stands 4.55 d: 1.094” 10.07 i 1.174 19.03 :I: 2.760“
Sugar maple
A cer saccharum
absent 10.78 :t 1.940 14.53 :1: 1.760 13.11 i 3.838 12.62 d: 1.249y
light 4.94 i 2.430 4.74 :t 2.441 8.75 a: 6.226 5.49 :t 1.606’rz
heavy 5.85 i 5.847 1.07 :t 1.070 3.14 :h 1.690 3.06 i 1.3092
all stands 9.22 :t 1.579a 10.81 i 1.6813 7.84 i 2.165a
Red maple
Acer rubrum
absent 1.97 i 0.645 1.86 d: 1.114 0.00 d: 0.000 1.67 :1: 0.5462
light 0.96 :1: 0.956 4.36 d: 2.762 0.00 :1: 0.000 2.21 :1: 1.2662
heavy 8.67 :t 0.076 7.15 i 3.203 4.87 i 1.730 6.18 :1: 1.269y
all stands 2.30 i 0.6293 3.04 i 1.037a 2.25 :t 1.033a '
Red oak
Quercus rubra
absent 2.85 i 1.682 1.22 i 0.943 0.00 i 0.000 1.82 :h 0.870y
light 5.56 d: 4.923 0.36 :1: 0.362 0.00 i 0.000 2.47 i 2.077y
heavy 10.43 i 10.429 2.36 :1: 2.356 2.91 :1: 2.540 4.13 d: 2.220y
allstands 4.00 :t 1.6773 1.19 i 0.6822' 1.34 i 1.190a
Other hardwood sp.l
absent 7.72 i 2.605 3.60 i 0.811 7.86 :t 0.905 6.05 :l: 1.276y
light 4.00 d: 1.706 0.98 d: 0.630 1.19 :t 1.188 2.27 d: 0.847y
heavy 16.97 1 14.797 2.41 d: 1.755 3.45 :t 1.664 5.63 :t 2.790y
all stands 7.72 :h 2.1653 2.91 d: 0.619a 4.80 :1: 1.0923
Eastern hemlock
T suga canadensis
absent 2.27 i 1.227 2.04 i 1.237 0.00 :t 0.000 1.89 :t 0.757y
light 2.96 i 2.274 0.00 i 0.000 0.00 :t 0.000 1.23 :1: 0.989y
heavy 0.83 i 0.828 4.33 i 2.224 3.37 i 2.602 3.17 a: 1.506y
all stands 2.30 :1: 0976‘“ 1.90 i 0.886a 1.55 d: 1.2408

 

lOther hardwood species included ash (F raxinus sp.), aspen (Populus sp.), basswood
( T ilia sp.), birch (Betula sp.), and black cherry (Prunus serotina ).

20

 

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21

Table 1-3. Spearman's correlation coefficients (rs) and significance values (p) for

basal area (mZ/ha) among overstory species and corresponding species of seedlings,
saplings, and recruits in 62 stands. Correlation coefficients with p<0.05 are
s_i_gnificant1y different.

Seedlings Saplings Recruits
American beech
rs 0.216 0.072 0.115
p 0.092 0.579 0.373
Sugar maple
rs 0.552 0.517 0.290
p <0.001 <0.001 0.022
Red maple
rs 0.229 0.253 0.434
p 0.074 0.047 <0.001
Other hardwood sp.a
rs -0.258 -0.024 0.029
p 0.043 0.855 0.821
Conifer sp. b
rs 0.049 0.092 0.182
p 0.704 0.476 0.157

 

aOther hardwood species included red oak (Quercus rubra) ash (Fraxinus sp.),
aspen (Populus sp.), basswood (T ilia sp.), birch (Betula sp.), and black cherry
(Prunus serotina ).

bOther conifer species included hemlock (Tsuga canadensis ), balsam fir (A bies
balsamea) , and white pine (Pinus monticola ).

22

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23

Figure Legends
Figure 1-1. Distribution of beech bark disease research sites and Cryptococcusfagz’suga

infestation indices throughout Michigan, 2003.

Figure 1-2. Intensive research plot design indicating arrangement of subplots 1-5,

overstory composition transects, and regeneration plots.

Figure 1-3. Number of stands with absent, light, and heavy Cryptococcusfagisuga

abundance among stands with low, moderate, and high beech densities.

Figure 1-4. Interactive effects between beech density classification and C. fagisuga

infestation for basal area of overstory a) beech, b) sugar maple, and c) red maple.

Figure 1-5. Interactive effects between beech density classification and C. fagisuga

infestation for sugar maple a) seedlings, b) saplings, and c) recruits.

Figure 1-6. Interactive effects between beech density classification and C. fagisuga

infestation for red maple a) seedlings, b) saplings, and c) recruits.

24

1.1 _°

 

 

 

 

 

 

 

Manistee

 

 

 

 

Mason

Oceana
Cryptococcusfagisuga densities:
0 Absent (n = 39) f.) ,

® Light (n = 12)
O

 

 

 

 

 

 

Heavy (n= 11)

 

Counties with BBD indicated by text.

5
I:
:4

25

 

Key to plot diagram:

Regeneration plot

__ Overstory basal area sampling transect

Subplot

26

Number of stands

20
18
l6
14
12

ONAO‘OO

bw

 

mode rate

Beech density

27

C. fagisuga
infestation

E] absent

I light
I heavy

 

high

Basal area (ml/ha)

Basal area (mz/ha)

Basal area (mZ/ha)

a) Overstory beech

 

 

 

 

 

 

 

 

absent light heavy

C. fagisuga infestation

b) Overstory sugar maple
16.0

 

14.0 ‘
12.0 -
10.0 -.
8.0 ~
6.0 -
4.0 —

 

 

 

2.0 a
0.0 I 1

 

 

 

 

absent light heavy

C. fagisuga infestation

c) Overstory red mple
10.0

 

 

 

 

 

 

 

absent light heavy
C. fagisuga infestation

28

Beech density

 

low
- - ‘- -— moderate
...- '— high

Beech density

 

low
~--—moderate
———'—high

Beech density
bw

—- -———-moderate

---high

 

Number of stems Number Of stems

Number of stems

3) Sugar maple seedlings

 

200,000

150,(X)0 ~

100,(X)0 a -

50,(X)0 -

 

 

 

 

 

 

light

C. fagisuga infestation

absent

b) Sugar maple saplings

 

7,000

6,000 “'

5,(I)0 ‘
4,000 J

3,000 l

2,(I)O r
l,(I)0 1

 

 

 

 

absent

light

C. fagisuga infestation

c) Sugar maple recruits

 

Beech density
low

- ‘- -* -— moderate
"r"hgh

 

Beech density
low

 

—' r r —' moderate
''''' high

 

600

500 «A

400 -
300 ~
200 -
100 -

 

 

 

 

4 Beech density

 

low

-' -— — - moderate

""'""hlg'1

 

 

light

C. fagisuga infestation

absent

29

Number of stems Number of stems

Number of stems

a) Red maple seedlings

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

35,000 .
Beech densny
30,000 - -— - low
25,000 ‘ C — ———--— -— moderate
20,000 -1 .. o -—H ...-.-th
15,000
10,000 ~- ___
5,000 r" ——o
0
absent light heavy
C. fagisuga infestation
b) Red maple sapl'ngs
Beech density
bw
" " ’ ’ moderate
**** high
absent light heavy
C. fagisuga infestation
c) Red maple recru'ts
250 Beech density
I
200 ’v.’/ low
I” r " r ’ moderate
150 ’1” """high
100 , ’ ,,
I ” I .” ’
50 ’ ’ , .J'
0 A] ’ ’ T “ J ' ’ 1
absent light heavy

C. fagisuga infestation

30

IMPACT OF VARYING STAGES OF BEECH BARK DISEASE ON WILDLIFE

RESOURCES IN MICHIGAN.

31

Introduction

American beech (F agus grandifolia Ehrh.) provides important food and habitat
resources for diverse bird, mammal, and amphibian species. Beech, a shade tolerant
species, is characteristic of late successional communities. It is a major component of
Michigan hardwood forests, typically associated with sugar maple (Acer saccharum),
eastern hemlock (T saga canadensis), red maple (Acer rubrum), red oak (Quercus rubra),
black cherry (Prunus serotina), and birch (Betula sp.).

Mature beech trees, greater than 40 years of age, periodically produce crops of
highly nutritious nuts (Dix 1975, Forcier 1973, Fowles 1965, Piovesan 2001, Tubbs and
Houston 1990, but see Statler 1982). Afier one year of maturation, beech nuts ripen
between September and October and fall after the first heavy frost (Tubbs and Houston
1990). Blue jays (Cyanocitta cristata) are among the many animals that utilize this
source of hard mast and, through caching behavior, contribute to long-range dispersal of
beech (Held 1983, Johnson and Adkisson 1985). Many wildlife species rely on
concentrated fat resources in the fall prior to hibernation and beech nuts, which have a
high fat content, are particularly valuable in stands without other mast-producing species
such as oak and hickory (Carya sp.) (Piovesan 2001, Tubbs and Houston 1990). Fertility
of black bears (Ursus americanus), for example, is influenced by females attaining a
threshold weight prior to hibernation and fecundity has been shown to cycle
synchronously with hard mast production (Lariviere 1994, Rogers 1987). Populations of
white-footed mice (Peromyscus leucopus), deer mice (P. maniculatus), and red-backed
voles (Clethrionomys gapperi) also cycle in synchrony with beech mast years (Schnurr et

012002)

32

Mature beech trees are susceptible to many cavity-forming pathogens and these
cavities provide shelter, nesting, and hibernation resources for a diverse array of wildlife
species that rely upon them (Allen 1990, Conner et al. 1975, Goodburn and Lorimer
1998, Mannan et al. 1980, Paragi 1996, Raphael and White 1984,Robb and Bookhout
1995, Runde and Capen 1987, Sadlon, Steele 1995, Stribling 1990, Welsh et. al. 1992);
In addition, as beech trees decline and die, they provide critical habitat for a variety of
wildlife species. Standing dead trees, termed “snags”, create important forage, hunting,
and roosting habitat for birds (Conner et. a1 1975, Mannan et. a1 1980, Raphael and White
1984, Paragi 1996, Stribling 1990). Fallen trees and branches accumulate on the forest
floor as coarse woody debris, which provides shelter, travel corridors, and forage
substrate for many small mammals, reptiles, and amphibians (Bowman et.al. 2000,
Dueser I978, Harmon et.al. 1986, Jaeger 1980).

As beech bark disease spreads throughout hardwood forests of Michigan, beech
nut production, cavity abundance, snag density, and coarse woody debris characteristics
will likely be altered. Beech trees killed by beech bark disease may be replaced by other
species that produce less energetically valuable seeds, such as maple, or by dense thickets
of stunted, beech root-sprouts that fail to reach nut-producing maturity (Houston 1994).
Availability of suitable wildlife cavities may decline as mature beech are removed from
the overstory and replaced by species less prone to cavity-formation. As mature beech
decline in vigor, changes in the abundance of snags and coarse woody debris may occur,
potentially increasing habitat for certain wildlife species, for at least a period of time.

We initiated a long-term study to address how wildlife resources will be affected

as beech bark disease spreads in range and intensity throughout Michigan. Our specific

33

objectives were to i) collect baseline data related to beech nut masting and abundances of
cavities, snags, and coarse woody debris in stands not yet affected by beech bark disease
infestation; ii) compare current abundance of wildlife resources among stands with
varying levels of beech bark disease and C. fagisuga infestation; and iii) establish long-

term research plots to monitor changes in wildlife resources over time.

Methods
Study Sites

Study site selection was described in detail in Chapter One.

Beech nut production

Beech nuts were collected using four leaf litter/seed traps set between subplots 2
and 5 in each stand (Figure 2-1). Traps were constructed from industry standard 1.9 cm
diameter (% in) pvc pipe assembled into a horizontal one m2 square frame. The
horizontal frame was elevated approximately 70 cm from the ground and supported by
legs of varying length (to account for variable terrain). Shade cloth (40%) was attached
to the frame to collect beech nuts , as well as acorns (red oak mast), that fell from
overstory trees. Traps were placed in the 40 stands established in 2002 by 15 August and
retrieved by 5 December. In 2003, traps were placed in 54 of the 62 total stands by 15
August and were retrieved by 7 December, afier leaf drop was complete. Traps were not
set in eight stands in 2003 because of heavy timber harvesting (three stands), seasonal
inaccessibility (two stands), and vandalism (three stands). Due to these limitations and

because of damage to traps, seed was collected from only five stands with heavy C.

34

fagisuga infestation. Leaves and seeds from each trap were bagged and returned to the
Forest Entomology laboratory at Michigan State University. All seed, beech nut, and
acorn samples were sorted to species, oven-dried at 38°C (100° F) for 72 hours, and
weighed. To test viability, beech nuts were floated in de—ionized water for 40 hours
(Williams and Hanks 1976). Beech nuts remaining in suspension were not considered
viable seeds. Preliminary dissection tests indicated that this method accurately separated
green, viable seeds from the nonviable. Mean values for number of beechnuts, number of
viable beechnuts, and mass of beechnuts were calculated from the number of successful

traps in each stand.

Cavities

Cavity abundance in standing live trees was assessed after leaf senescence in all
stands between 21 November and 7 December 2003. We recorded species, dbh, and
number of cavities of each of three size classifications for all trees greater than 25.4 cm
dbh in the five subplots in each stand. Trees less than 23.4 cm were not sampled because
previous studies have shown cavities occur predominantly in large diameter trees
(Goodbum and Lorimer I998, Gysel 1961, Welsh and Capen 1992). Cavity diameter,
visually estimated from the ground, was classified as small (<7.0 cm diameter), moderate
(7030.0 cm diameter), or large (>30.0 cm diameter). The main hole and limbs of trees
were examined for cavities when leaves did not obstruct the view of the crown.

Cavity abundance and size was also assessed for individual beech trees selected
for long-term monitoring (Appendix 4). In each stand, 12 beech trees were selected 10.7

m from the center of each of the four outer subplots in each cardinal direction, excluding

35

the azimuth leading toward the center subplot, and marked with round blue tree tags
(Figure 2-1). When more than one beech tree fell within the designated distance and
azimuth, we preferentially selected a tree by the following prioritized criteria: 1) the
largest tree with a canker, 2) any tree (pole-sized or larger) with a canker, 3) large tree
with C. fagisuga present, 4) any tree (pole-sized or larger) with C. fagisuga, or 5) any
large tree (closest to the azimuth used). During full leaf expansion in the summer,
canopy dieback was visually estimated for each of the 12 trees as low (<30%), moderate

(30-60%), and high (>60°/0).

Coarse woody debris and snags

Abundance and volume of coarse woody debris and basal area and number of
snags were measured in each stand along two transects, 51 m long by l m wide, running
between subplots 2 and 5 and subplots 2 and 3 (Figure 2-1). Each piece of coarse woody
debris greater than 7.6 cm diameter and occurring within 0.5 m of the transects was
measured and decay recorded as 0 (freshly downed, no discoloration of sapwood, bark
intact, no visible decay), 1 (discoloration of sapwood, bark intact, no visible decay), 2
(form maintained under impact, bark peeling, visible decay), 3 (crushes under impact,
bark removed, mosses and advanced visible decay), or 4 (little structural integrity, mixing
with soil)(Daniels 1997). Volume of coarse woody debris was calculated as cylinders
(length x 1t(1/diameter)2) for pieces in decay classes 0-3 and as rectangles (length x
width x height) for pieces in decay class 4. All standing dead trees, within five meters of
each transect, greater than 12.7 cm diameter at breast height (dbh) (1.3 m) were counted,

identified to species, and dbh was measured.

36

Statistical analyses

Normality was tested for all variables using the Shapiro-Wilk test (SAS Institute,
1985). Mean oven-dried beech nut masses and mean volume of coarse woody debris (all
decay classes pooled) were log transformed to achieve normality. Mean number of snags
(all species combined) and mean number of beech snags were square—root transformed to
achieve normality. (N on-transfonned means are referenced in tables.) Two-way
ANOVA [proc glm] was used to determine if mean beech nut mass, number of beech
nuts collected in 2002 and 2003, volume of coarse woody debris in each decay class,
number of all species of snags, and number of beech snags differed among stands with
low, moderate, or high beech density and absent, light, or heavy C. fagisuga infestation
(SAS Institute 1985). When ANOVA results were significant (oc=0.05), means were
separated by Tukey’s Honestly Significant Difference (Tukey 1977). Transformation did
not normalize data distributions for mean number of viable beech nuts per stand, number
of cavities in each size class, number of cavities in trees of each dieback rating, volume
of coarse woody debris in each decay class, basal area of all species of snags, and basal
area of beech snags. These variables, therefore, were ranked and the nonparametric
Kruskal-Wallis test [proc rank] was used to determine if these variables differed among
stands with low, moderate, or high beech density and absent, light, or heavy C. fagisuga
infestation (SAS Institute 1985). For significance tests ((1 =0.05), means were compared
with nonparametric Tukey’s-type multiple comparison (Zar 1984). Spearman’s
nonparametric correlation analysis was used to assess the relationship between basal area

of beech and red oak in the overstory with the mass of beech nuts and acorns,

37

respectively (SAS Institute 1985). Spearman’s nonparametric correlations were also used
to determine if total overstory basal area and beech basal area were related to basal area
of snags (all species combined) and beech snags, respectively. All analyses were done
using SAS 9.1, except for the nonparametric Tukey’s-type multiple comparison which
was done with Microsoft Excel software (Redmond, WA).

Three stands with recent selective harvest activity were omitted from coarse
woody debris analyses to avoid confounding effects of logging slash and beech bark

disease impacts (stand ID 104, 107, 121) (Appendix 3).

Results
Beech nuts

In 2002, beech nuts were collected from all 36 stands we surveyed. Mean beech
nut mass in 2002 averaged 18.4 :1: 2.31 g/m2 and did not significantly differ among stands
with absent, light, or heavy C. fagisuga infestation (p=0.234), nor among stands with
low, moderate, or high beech density (p=0.162) (Figure 2-2). Mean beech nut mass
varied considerably among stands, ranging from 0.1 to 51.8 g/mz. Mean beech nut mass
within stands was also variable. In one stand, for example, mean beech nut mass among
the four traps varied from 0.38 g/m2 to l 19.00 g/mz. Basal area of beech in the overstory
was significantly correlated with beech nut mass produced in 2002 (r,=0.393, p=0.018).
Mean number of beech nuts collected in 2002 averaged 98 i 11.3 /m2 and was not
significantly affected by C. fagisuga infestation (p=0.614), nor beech density (p=0.490)

(Figure 2-2). An average of 51 % of the beech nuts collected in 2002 were viable.

38

In 2003, beech nuts were collected from 27 of the 51 stands we surveyed. Only
four stands in 2003 produced more than 1.0 g/m2 beech nut mass; three of the four were
in stands classified as low beech density. Mean beech nut mass in 2003 averaged 0.3 :t
0.09 g/m2 and number of beech nuts only averaged 2.0 3: .06 /m2. Neither mass nor
number of beech nuts was significantly affected by C. fagisuga infestation level
(p=0.053, p=0.221, respectively) or beech density (p=0.771, p=0.534, respectively)
(Figure 2-2). Only 10% of beech nuts collected in 2003 were viable. There was no
significant correlation between beech basal area and beech nut mass produced in 2003
(p=0.205). For the 32 stands sampled for two consecutive years, total beech nut
production decreased by an overall mean of 99.5 d: 2.31 beech nuts/m2 and viable beech
nut production decreased by a mean of 52.8 :i: 1.35 beech nuts/m2.

Red oak was encountered in the overstory of 13 stands with basal area ranging
from 1.47 to 26.32 mz/ha. In 2002, acorns were collected from six of the 36 stands, all in
the Lower Peninsula. Mean acorn mass in 2002 averaged 1.0 :i: 0.50 g/mz. In 2003,
acorns were collected from seven of the 51 stands, all in the Lower Peninsula except for
one stand. Mean acorn mass in 2002 averaged 0.9 :t 0.54 g/mz. Basal area of oak in the
overstory was significantly correlated with acorn mass in 2002 (r,=0.647, p<0.001) and

in 2003 (r, =0.407, p<0.001).

Cavities
Cavity size and number were recorded for a total of 683 trees representing 12
species (Table 2-1). At least one tree of each species, except black cherry, contained

small cavities (<7.0 cm diameter). Small cavities occurred in trees ranging from the

39

smallest tree sampled (10.0 cm dbh) to the largest tree sampled (29.0 cm dbh). There
were significantly more small cavities in American beech trees than in any other species
sampled (df=8,673; F =1 7.01, p<0.001) (Table 2-1). Tree age was significantly
correlated with diameter (r,=0.494, p<0.001) and number of cavities (r,=0.215,

p=0.03 8). Diameter and number of cavities were also significantly correlated (r,=0.336,
p=0.001).

Beech trees contained significantly more moderately sized (7.0 to 30.0 cm
diameter) cavities than sugar maple trees (df——8,673; F =3.64, p<0.001) but differences
between beech and other species were not significant. A beech tree that was 48.1 cm dbh
contained the maximum number of moderately sized cavities encountered (11) and
occurred in a stand with light C. fagisuga infestation. A total of 89 trees comprised of
red oak, black cherry, ash, hemlock, and basswood trees were examined; none had
moderate seized cavities.

Large diameter cavities (>30.0 cm diameter) occurred on only six of the 683 trees
and did not significantly differ among species, likely due to the limited sample size
(p=0.697). Five of the large cavities were on the main bole of beech treesand one was on
the main bole of a sugar maple; no single tree had more than one large cavity. The six
trees with large cavities ranged from 13.3 cm to 28.4 cm in dbh with an average of 21 .3 i
2.85 cm.

Individual trees selected for long-term monitoring (n=94) from the perimeter of
subplots had a mean diameter of 3 1 .8 cm and were 82 years old on average (see Chapter
3). The number of cavities per tree was significantly correlated with age (r,=0.215,

p=0.038), diameter (r,=0.336, p=0.001), and canopy dieback (r3: 0.156, p=0.134) (Table

40

2-2). Age was significantly correlated with diameter (r,=0.394, p=0.000) and dieback
(r,=0.255, p=0.013). Dieback was significantly correlated with diameter (r,=0.339,

p=0.001) (Table 2-2).

Coarse Woody Debris and Snags

Coarse woody debris was encountered in every stand and ranged from 2.5 to
235.0 m3/ha. A total of 746 pieces were encountered on the transects. Size of pieces
ranged from 008—122 m length; 49% of the pieces less than 1 m length, 48% were
between 1 and 3 m length, and 3% were greater than 3 m long. Only 64% of the coarse
woody debris pieces sampled retained characteristics that allowed identification to
species. Identified pieces were predominantly beech (24% of total) and white birch
(Betula papyrifera) (12% of total) and a lesser component of sugar maple, hemlock,
aspen (Populus sp.), and black cherry.

Volume of coarse woody debris did not vary among stands with low, moderate, or
high beech density (p=0.061) and absent, light, or heavy C. fagisuga infestation
(p=0.192) (Table 2-2). The interaction between beech density classification and C.
fagisuga infestation level was not significant for coarse woody debris volume (p=0.314).

Volume of coarse woody debris in decay classes 0, l, 2, 3, and 4 was not
significantly affected by C. fagisuga infestation level (decay class 0: p=0.226) (decay
class 1: p=0.557) (decay class 2: p=0.467) (decay class 3: p=0.735) (decay class 4:
p=0.153) (Table 2-2). Only 1% of total coarse woody debris volume was of decay class

0. These relatively fresh pieces were encountered only in seven stands, three of which

41

had heavy C. fagisuga infestation. The majority of total coarse woody debris volume
(62%) was assigned to decay classes 2 and 3.

Snags were present in 45 of the 62 stands we sampled, but only 12 stands
contained beech snags. A total of 112 snags with a mean dbh of 31 :t 1.6 cm were
examined. Snags were predominantly beech (24), sugar maple (17), white birch (l 7), and
black cherry (1 1). Basal area of snags of all species combined (including beech) did not
significantly differ among stands with low, moderate, or high beech density (p=0.528)
and absent, light, or heavy C. fagisuga infestation (p=0.155) (Table 2-3). Mean number
of snags of all species also was not significantly affected by beech density (p=0.113) or
C. fagisuga infestation (p=0.100) (Table 2-3). Basal area of all species of trees in the
overstory was not correlated with basal area of all species of snags (p=0.273).

Mean basal area of beech snags was significantly higher in stands with high beech
density (dfi2,59; F =4.57, p=0.014) and in stands with heavy C. fagisuga infestation
(df=2,59; F =4.65, p=0.013) than in other stands. Similarly, significantly more beech
snags occurred in stands with high beech density (dfi-2,59; F=5.17, p=0.009) and in
stands with heavy C. fagisuga infestation (dfi2,59; F =4.08, p=0.022) than in other
stands. The interactive effect between beech density classification and C. fagisuga
infestation was significant for basal area of beech snags (off—2,59; F =2.34, p=0.083) and
number of beech snags (df=2,59; F =3.57, p=0.012). Basal area and number of beech
snags were at least two times more abundant in stands with high beech density and heavy

C. fagisuga infestation than in other stands.

Discussion

42

Beech nut production

The effect of beech bark disease on mature beech trees will eventually reduce
beech nut abundance and may affect the animals that nutritionally depend on them. Our
study results, however, indicate that stand-level C. fagisuga infestation had not yet
impacted nut production. Stands with heavy C. fagisuga infestation were in heavily used
recreation areas where leaf litter traps were often disturbed by humans or damaged by
falling trees and limbs. Consequently, our results might have been affected by the limited
sample size (n=5 stands) of intact beech nut collections. Beech bark disease may,
however, progressively stress mature trees and reduce the overstory component of
mature, nut-producing trees as it reaches the “Aftermath” forest structure in Michigan.
Therefore, continued monitoring will be needed to quantify effects of beech bark disease
on nut production.

Beech nut production can fluctuate annually by up to 65% and the phenomenon of
even year mast events has been well documented (Gysel 1971, Piovesan 2001 , Schnurr
2002). Our results indicate a mast event occurred in 2002 followed by a substantially
reduced intermast crop in 2003. Levels of beech nut production we documented in the
mast event of 2002 (98 beech nuts/m2) are similar to a previous study in Michigan. Gysel
(1971) reported beech nut production in mast years averaged 40 beech nuts/m2 among
stands, with production in one stand exceeding 250 beech nuts/m2. The viability of beech
nuts in our study also varied between mast and intermast years, ranging from 58% in
2002 to less than 10% in 2003. Viability in 2002 was markedly higher than that reported

by Gysel (1971) who found only 10% of nuts were viable during mast years.

43

Our methods did not account for the possible consumption of beech nuts by blue
jays in early August before seed drop and often before seeds are fully ripe. (Johnson and
Adkisson 1985). Blue jays select only viable seeds, potentially increasing the proportion
of non-viable seeds in our traps. We collected beech nuts throughout the caching period
but from near the forest floor and consequently, may have underestimated actual

production (Johnson and Adkisson 1985).

Cavities

Our results demonstrate that cavities of all diameter classes were more abundant
in beech than in other overstory species, except white birch. Other studies conducted in
hardwood forests found that beech had more small cavities than other species (Gysel
1961, Robb and Bookhout 1995). Tree diameter was correlated with number of cavities,
indicating that declines in mature beech from beech bark disease will likely reduce the
abundance of cavities available for wildlife. Gysel (1961) found that cavities greater than
10 cm diameter occurred more often in trees of declining vigor. This suggests that
mature trees impacted by beech bark disease may contain an abundance of moderate to
large size cavities. Our data indicated no difference in cavity abundance among stands
with varying levels of beech bark disease nor among trees with varying degrees of
dieback.

Both size and location of cavities can influence habitat suitability for specific
species. For example, downy woodpeckers (Picoides pubescens L.) prefer to nest in 20-
cm diameter cavities located 6 m above ground, whereas large pileated woodpeckers

(Dryocopus pileatus L.) prefer 56-cm diameter cavities 18 m aboveground (Sadlon

44

1992). In addition to external cavity dimensions, internal dimension can also be critical,
as in the case of wood ducks (Aix sponsa) that prefer entrances of 7 x 9 cm with bottom
dimensions of at least 12 x 17 cm (Robb and Bookhout 1995). In our study, five of the
six large cavities we encountered were in beech trees. Eventual loss of large diameter
beech and the corresponding loss of large cavities may have negative consequences for

certain wildlife species.

Coarse woody debris and snags

Coarse woody debris volume was consistently greater in stands with heavy C.
fagisuga infestation, although means were not statistically different because of the limited
number of stands with heavy C. fagisuga infestation. Studies conducted in northeastern
states impacted by beech bark disease showed a consequent increase in coarse woody
debris volume of 20-30% with volumes exceeding 166 at 42 m3/ha, whereas stands
unaffected by beech bark disease contained 15-45 m3/ha (Carbonneau 1986, McGee
2000). Mean volumes of coarse woody debris from our stands with heavy C. fagisuga
exceeded the maximum volume measured in uninfested stands, but, this was likely due to
stand-level variations in productivity.

Decay characteristics of coarse woody debris can determine species-specific
values for wildlife (Bowman 2000, Jaeger 1980). For example, red-backed voles (C.
gapperi) use highly decayed logs for shelter and fresh, solid form logs for runways,
thereby requiring a diversity of decay classes (Bowman et. al. 2000, Maser and Trappe
1994). Highly decayed logs also serve as critical nutrient and moisture sinks on the forest

floor (Covington 1981, Fraver et al. 2002). In our study, coarse woody debris volumes in

45

 

stands with heavy infestations of C. fagisuga were evenly distributed among decay
classes, indicating no recent accumulation of freshly downed material, as might be
expected from declining, breaking trees.

The greater number and basal area of beech snags in stands with heavy C.
fagisuga suggests beech bark disease has begun to affect individual trees and that stands
have progressed to the “Killing Front”. Abundance of snags of other species remained
constant among stands, further supporting that beech bark disease was specifically
contributing to greater snag abundance. The great diversity of wildlife using snags may
benefit from this alteration of forest structure for some period of time, although specific
characteristics, such as decay stage and remaining branches, influence actual value.
Hardwood and conifer stands in which snags were retained supported a greater diversity
of wildlife in eastern and western states, although surpassing minimum density thresholds
may not proportionately benefit wildlife (Gunn and Hagan 2000, Raphael and White
1984, Paragi 1996, Robb and Bookhout 1995, Scott 1979, Stribling 1990, Yahner 1987).

To date, beech bark disease has likely contributed to an increase in snag
abundance in stands with heavy infestation of C. fagisuga, but other wildlife resources
have not been significantly altered. More cavities were found in beech trees than in any
other species occurring in the stands we examined. Beech was typically the only hard
mast-producing species except in seven stands where red oak also occurred. As mature
beech succumb to beech bark disease and are replaced by other species, the abundance of
hard mast and cavities will be reduced. Snags and coarse woody debris, however, will
likely continue to increase as beech bark disease progresses and trees die. The data

presented here provide a valuable baseline to compare with data acquired from long-term

46

monitoring in the same plots as beech bark disease expands in range and intensity

throughout Michigan.

47

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48

Table 2-2. Volume of coarse woody debris (m3/ha i SE) grouped by decay class
among stands with low, moderate, and high beech density and absent, light, and

heavy C. fagisuga infestation.

 

 

Beech C. fagisuga infestation

densifi absent light heavy all stands
Decay Classes Poolec

low 46.2 i: 3.21 29.4 d: 3.44 50.6 :1: 4.82 43.1 i 1.95
moderate 62.3 :1: 3.78 77.6 i 15.51 125.3 3: 19.69 74.8 i 1.89
high 43.8 i 6.01 40.8 :t 22.33 70.2 i 7.82 55.5 i 3.06
allstands 52.4 d: 1.41 51.4 :t 4.59 81.7 d: 4.68

Decay Class 02'

low 0.3 :I: 0.08 0.4 d: 0.17 0.0 i 0.00 0.3 :1: 0.05
moderate 0.1 d: 0.03 0.0 3: 0.00 0.4 :t 0.25 0.1 :t 0.01
high 0.2 i 0.09 0.0 i 0.00 5.3 :t 2.06 2.5 d: 0.65
all stands 0.2 i 0.03 0.2 :1: 0.04 3.0 i 0.83

Decay Class 1

low 3.9 :t 0.32 0.4 :1: 0.17 4.5 i 3.18 3.3 :I: 0.21
moderate 10.2 i 0.61 16.9 d: 4.61 4.7 :t 1.00 11.0 :1: 0.40
high 7.1 3: 1.85 5.3 i 3.76 8.5 d: 2.26 7.5 :1: 0.81
all stands 6.8 d: 0.21 8.1 d: 1.35 6.7 :1: 0.92

Decay Class2

low 13.0 :t 0.68 10.0 i 1.73 6.2 d: 0.28 11.8 i 0.44
moderate 20.5 i 1.17 13.0 d: 3.13 57.8 :t 15.69 23.9 :t 0.78
high 17.1 i 1.99 20.3 i 13.55 17.1 :t 3.09 17.6 31.19
all stands 16.5 i 0.38 13.0 :t 1.17 26.2 i 2.94

Decay Class3

low 10.9 i 0.70 13.2 :t 1.93 11.0 :t 0.41 11.4 :1: 0.45
moderate 19.7 i 2.14 22.6 :t 5.32 52.3 3 11.04 24.8 :t 0.95
high 15.5 :t 2.26 13.7 :t 4.87 9.9 d: 2.22 12.6 :t 0.88
all stands 14.9 :I: 0.57 17.2 :t 1.50 21.7 :1: 2.40

Decay C1ass4

low 6.6 i 0.71 5.3 :t 1.60 28.9 :t 8.13 8.2 :I: 0.53
moderate 12.5 :1: 1.92 25.1 i 8.37 10.0 i 3.36 15.0 i 0.90
high 3.8 i 1.71 1.4 i 0.15 29.5 i 9.21 15.3 i 2.96
all stands 8.5 i 0.52 12.9 :t 2.32 24.1 i 3.70

 

a Decay classes recorded as 0 (freshly downed, no discoloration of sapwood, bark
intact, no visible decay), 1 (discoloration of sapwood, bark intact, no visible decay), 2
(form maintained under impact, bark peeling, visible decay), 3 (crushes under impact),
and 4 (little strucutral integrity, mixing with soils).

49

Table 2—3. Mean number (i SE) and basal area (mz/ ha :t SE) of beech snags and
snags of all species among stands with low, moderate, and high beech density and
among stands with absent, light, and heavy C. fagisuga infestation Means
followed by different letters are significantly different among rows (a-b) or among

columns (y-z).

Number of snags /ha

 

 

 

 

All species C. fagisuga abundance
Beech density absent light heavy all stands
low 15 i 1.0 25 i 3.0 5 i 3.5 16 :1: 0.7
moderate 12 :1: 0.8 14 :t 3.0 23 :I: 7.5 13 :h 0.4
high 24 :t 3.8 15 :t 10.4 36 i 4.6 28 :1: 1.8
all stands 15 i 0.4 19 :t 1.3 27 :1: 2.3
Beech
Beech density absent light heavy all stands
low 1 :1: 0.1 4 d: 1.1 0 a: 0.0 1 i 0.12
moderate 1 3: 0.2 0 i: 0.0 3 :1: 1.9 1 at 0.1Z
high 6 d: 2.6 10 i 6.9 21 :1: 4.6 14 :1: 1.6y
all stands 2 i 0.1b 3 :1: 0.5ab 12 i 1.53

Basal area of snags (ml/ha)
All species
Beech density absent light heavy all stands
low 1.0 d: 0.08 1.2 :I: 0.26 0.6 d: 0.42 1.0 :1: 0.05
moderate 0.8 i 0.07 2.0 i 0.56 4.8 :t 1.46 1.6 i 0.10
high 44.8 :1: 0.34 1.5 d: 1.07 2.5 d: 0.58 1.9 i 0.20
all stands 1.0 i 0.03 1.6 :1: 0.17 2.8 :1: 0.32
Beech
Beech density absent light heavy all stands
low 0.1 i 0.03 3.5 i 1.09 0.0 :t 0.00 0.8 d: 0.112
moderate 0.8 i 0.14 0.0 i 0.00 2.2 i 1.28 0.8 :1: 0.092
high 1.6 :1: 0.70 5.6 :1: 3.97 12.5 i: 2.62 7.2 d: 0.91y
all stands 0.6 :t 0.05b 2.4 a: 0.39ab 7.4 i 1.16“

 

50

Figure legends
Figure 2-1. Intensive research plot design indicating arrangement of subplots 1-5,
individual beech trees selected for long-term monitoring, beech nut collection traps, and

coarse woody debris and snag measurement transects.

Figure 2-2. Beech nut mass (g/m2 i SE) among stands with low, moderate, and high

beech density and absent, light, and heavy C. fagisuga infestation in a) 2002 and b) 2003.

51

 

X
El
X X
Key to plot diagram:
X Sample tree X

LEI Leaf litter trap

Coarse woody debris and snag

sampling transect

Subplot

52

Mass of beechnuts (g/mz)

Mass of beechnuts (g/mz)

a) 2002

 

 

El bw beech density
——: nndemte beech density

I high beech density

J

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

1——1

  

 

 

 

 

 

 

 

 

 

 

 

 

 

absent light heavy
C. fagisuga abundance

 

 

 

 

 

 

 

 

 

 

b) 2003
4
3 ..L._. ...... ___
2 _, B A#__, # _7 A #__ ,,_
1 2 ____- _ g __ __ ____
O _ 3...... HM.
absent light heavy

C. fagisuga abundance

53

 

RADIAL GROWTH AND LEAF BIOMASS OF MICHIGAN BEECH FORESTS

IMPACTED BY VARYING LEVELS OF BEECH BARK DISEASE.

54

Introduction

Beech bark disease has been advancing throughout the range of beech in North
America and, in 2000, was discovered in Mason County in the northwest Lower
Peninsula Michigan and Luce County in the eastern Upper Peninsula Michigan
(McCullough et.al. 2001, O’Brien et.al. 2001). In 2001, we initiated a study to collect
baseline data for long-term comparisons of forest productivity as beech bark disease
spreads and intensifies throughout Michigan. The primary objective of this project was
to determine the impacts of beech bark disease on total biomass production in stands by
quantifying radial grth and foliage production of beech in stands with varying levels of
beech bark disease and beech densities. Our research addressed the following questions:
i) Does infestation of C. fagisuga impact radial growth of individual trees and if so, are
stand-level trends evident? ii) Is there a difference in foliar biomass of beech and other
species produced among stands with varying levels of beech bark disease?

Although beech bark disease can kill trees, some beech trees survive decades after
infection, persisting in a state of decline and contributing limited biomass production to
the stand (Hane 2003, Shigo 1972, Twery and Patterson 1984). Reductions in radial
growth lead to lower timber and fiber production in a stand. Predicting the impact of
beech bark disease, therefore, is important for long-term management of sustainable

forests.

Methods

Study Sites

55

 

These data were collected from the study sites described in Chapter One and

classified by the same beech density and C. fagisuga infestation criteria.

Radial growth

We selected 12 beech trees, 10.67 m from the center of each of the four outer
subplots in each cardinal direction, excluding the azimuth leading toward the center
subplot (Figure 3-1) for long-term monitoring. Each tree was marked with a round, blue
tree tag nailed into the base of the tree facing the subplot center. When more than one
beech tree fell within the designated distance and azimuth, we preferentially selected a
tree by the following prioritized criteria: 1) the largest tree with a canker, 2) any tree
(pole-sized or larger) with a canker, 3) large tree with C. fagisuga present, 4) any tree
(pole-sized or larger) with beech scale, or 5) any large tree (closest to the azimuth used).

Abundance of C. fagisuga on each of the 12 trees was estimated by overlaying a
transparent, acetate grid containing 50 cells, each 1.9] cm2, at 1.3 m aboveground on the
north and south azimuths of the main bole. Each cell was assigned a rating based on the
area occupied by C. fagisuga as absent, light (<10%), or heavy (>10%). Overall values
were then assigned to each tree as 1) the percentage of the main bole with C. fagisuga
present and 2) the percentage of the main bole heavily covered by C. fagisuga. CanOpy
dieback was visually estimated for each tree and, based on the percentage of mortality in
the crown, classified as absent (0), low (< 30%), moderate (30-60%), and high (>60%).

To analyze radial grth of beech trees before and afier the onset of beech bark
disease, increment cores were extracted from the 12 beech trees in each stand between

May and August 2002 or June and August 2003. Because trees were actively growing

56

 

when cored, current year increments were not included in analyses to eliminate partial
increments. Increment cores were extracted at approximately 1 meter above ground
perpendicular to the main bole of each of the 12 beech trees using a cordless drill and
case-hardened coring bit. Coring tools were dipped in alcohol and flamed before
sampling each tree to prevent possible artificial inoculation of Nectria sp. to uninfected
trees. Increment cores were mounted on wooden holders, sanded four times with
progressively finer paper (100 grit, 220 grit, 600 grit, and emery cloth) to create a flat
surface, then scanned into a digital image for annual increment measurements with
WinDENDRO® (Regent Instruments, Inc.). To distinguish between effects of site
conditions and beech bark disease, cores were also extracted from two non-beech trees
within each stand. Approximately half the cores were either damaged in the preservation
and measurement process or did not have sufficiently visible annual growth rings and
were excluded from analyses. To eliminate the effect of crown position on growth
pattern, only trees in the dominant and codominant dominance classes were included in
the analyses. Consequently, a total of 90 trees with intermediate crown class and 15
suppressed trees were removed from analyses. A total of 376 cores were analyzed; 304
were from beech and 72 were from non-host trees.

Mean distances between annual growth rings (mm/yr) were calculated for each
tree in 5, 10, and 15-year intervals. To identify any changes in growth in the last 5, 10, or
15 years, ratios were established for mean annual grth during 1997~2001 and 1992-
1996 (5 years), 1992-2001 and 1982-1991 (10 years), and 1987-2001 and 1986-1972 (15
years). Intervals were established around the last 5, 10, and 15-year periods because, as

of 2000, beech bark disease had likely been present in Mason and Luce Counties for at

57

 

least 10 years (O’Brien et al. 2001). The established intervals were also intended to
account for the expansion of beech bark disease into previously uninfected stands over
time. Although trees cored in 2003 included a complete increment for 2002, all analyses
were initiated from 2001 to maintain consistency. Annual increments (mm/yr) were
plotted for each tree and distinct reductions in growth were identified. To distinguish
regularly oscillating fluctuations growth patterns from distinct reductions in growth, only
annual increments that decreased below the average growth increment and did not
recover to previous growth increments for at least 10 years were considered growth
reductions.

We were not able to determine the age of 210 beech trees from which cores were
extracted due to light annual rings that were not detectable by WinDENDRO® or
because the heartwood was decayed with no pith remaining as a reference point. Tree
age was quantified for 94 beech trees from 45 stands. Some stands had no beech cores
successfully extracted and aged while others had up to six cores with complete analysis.
Fifty five beech cores were aged from stands with absent C. fagisuga infestation (27 low,
23 moderate, and five high beech density), 24 stands with light C. fagisuga infestation
(11 low, 10 moderate, and three high beech density), and 15 stands with heavy C.
fagisuga infestation (one low, two moderate, and 12 high beech density).

Age was determined from 28 of the 72 non-host cores extracted, representing 28
stands, with a maximum of one core successfiilly aged from each stand. Non-host cores
were aged from 18 stands with absent C. fagisuga infestation (six low, eight moderate,

and four high beech density), four stands with light C. fagisuga infestation (zero low,

58

three moderate, and one high beech density), and six stands with heavy C. fagisuga

infestation (two low, two moderate, and two high beech density).

Leaf biomass

Leaves were collected in the 1m2 pvc traps described in Chapter Two. Traps
were not set in eight stands in 2003 because of heavy harvesting (three stands), seasonal
inaccessibility (two stands), and vandalism (three stands). Due to these limitations and to
site damage to traps, leaf collections were only made fi'om five stands with heavy C.
fagisuga infestation; none of these five stands had a heavy infestation of C. fagisuga.
Leaves from each trap were bagged and returned to the forest entomology lab at
Michigan State University. All leaf samples were sorted to species, oven-dried at 38°C

(1000 F) for 72 hours, and weighed.

Statistical analyses

Normality was tested for all variables using the Shapiro-Wilk test (SAS Institute,
1985). Data distributions for growth ratios for 5, 10, and lS-year intervals and overstory
basal area by species could not be normalized by transformation, thus these variables
were ranked and analyzed with nonparametric tests. The nonparametric Kruskal-Wallis
test was used to determine if growth ratios for 5, 10, and lS-year intervals differed among
stands with low, moderate, or high beech density and absent, light, or heavy C. fagisuga
infestation (SAS Institute 1985). Spearman’s correlations were used to determine if the
percent of the main hole with C. fagisuga or with heavy C. fagisuga correlated with

annual growth ratios for 5, 10, and 15-year intervals. Spearman’s nonparametric

59

correlation coefficients were also calculated to assess linear associations among age,
dieback, and annual growth ratios for 5, 10, and lS-year intervals.

Leaf biomass for each stand was calculated as the mean of all traps collected in
each stand. A paired t-test was used to test the differences in mean leaf mass produced in
2002 and 2003. Two-way ANOVA was used to test the differences in mean leaf mass of
(1) all species and (2) beech among stands with low, moderate, or high beech density and
among stands with absent, light, and heavy C. fagisuga infestation. When results of
ANOVA were significant (01 = 0.05), Tukey’s Honestly Significant Difference was used
to compare means (Tukey 1977). Basal area of overstory species were not normalized by
transformation, thus were ranked and analyzed with nonparametric tests. Spearman’s
correlation coefficients were calculated for basal area of each overstory species (beech,
sugar maple, red maple, red oak, other hardwoods, and all species combined) and their
respective leaf mass. All data analyses were conducted using the Statistical Analysis

System (a = 0.05) (SAS Institute 1985).

Results
Radial growth

A total of 304 beech cores from 59 stands were successfully extracted and
measured, an average of 5 :t 0.3 cores per stand (Appendix 4). Continuous, detectable
annual growth rings from bark to pith were present for only 94 beech trees, which
averaged 81i 2.5 years of age. Mean increments calculated for all 304 beech trees
averaged 2.2 d: 0.06 mm in the last 5 years, 2.1 :1: 0.05 mm in the last 10 years, and 2.1 i:

0.05 mm in the last 15 years (Table 3-1). Of the 304 beech trees examined, only 29

60

exhibited a reduction in radial growth, with onsets occurring between 1956 and 1997.
The 29 trees exhibiting growth reductions were from 21 different stands. Only four
stands contained more than one tree exhibiting growth reductions; none of these trees had
C. fagisuga.

Cryptococcusfagisuga was present on 73 of the 304 individual beech trees
measured. The percentage of the main bole covered with C. fagisuga varied greatly.
Five of the 73 infested trees exhibited a reduction in radial growth, with onsets of decline
occurring between 1956 and 1993. Two of the beech trees exhibiting decline (trees 776
and 820) had trace amounts of C. fagisuga present on the main bole (Appendix 4). The
other three declining beech trees had a heavy infestation of C. fagisuga on virtually all of
the main bole, with 40-55% of the main bole heavily covered. Each of the five heavily
infested trees that exhibited growth reduction were from separate stands, thus no
consistent reduction in radial growth was evident at a stand level.

Of the 304 beech trees cored, 101 exhibited no canopy dieback, 159 exhibited low
dieback, 35 exhibited moderate dieback, and nine exhibited heavy dieback.
Cryptococcusfagisuga was present on the main bole of ten trees exhibiting no dieback
(10%), 40 trees exhibiting low dieback (25%), 18 trees exhibiting moderate dieback
(51%) and five trees exhibiting high dieback (56%).

Ratios for 5, 10, and 15-year intervals of mean annual growth in beech trees did
not significantly differ among stands with absent, light, or heavy C. fagisuga infestation
(S-yr interval: p=0.966) (10-yr interval: p=0.076) (IS-yr interval: p=0.338), or among
stands with low, moderate, or high beech density (Table 3-1) (S-yr interval: p=0.884)(10-

yr interval: p=0.696)(15-yr interval: p=0.873). The percentage of the main bole with C.

61

fagisuga present was not correlated with growth ratios of 5, 10, or 15-year intervals (5-yr
interval: p=0.52) (IO-yr interval: p=0.948) (15-yr interval: p=0.842), nor was the
percentage of the main bole heavily covered by C. fagisuga (5-yr interval: p=0.982) (10-
yr interval: p=0.477) (1 5-yr interval: p=0.584). Tree age was not correlated with the
annual growth ratios for 5, 10, and lS-year intervals (5-yr interval: p=0.127) (10-yr
interval: p=0.774) (IS-yr interval: p=0.687) (Figure 3-2). Dieback was not correlated
with the annual growth ratios for 5, 10, and 15-year intervals (S-yr interval: p=0.173)

(10-yr interval: p=0.981) (IS-yr interval: p=0.377).

 

Non-beech tree cores were extracted to serve as a baseline for comparison with
grth reduction in beech. Sixty-two cores from non-beech trees from 45 stands were
successfully extracted and measured, an average of 1 :t 0.9 non-beech cores per stand.
Non-beech trees that were cored were primarily sugar maple (44 trees from 32 stands)
and red maple (15 trees from 12 stands). Continuous, detectable annual growth rings
from bark to pith were present for 25 non-beech trees and averaged 78 :t 4.8 years of age.
Mean increments for individual non-beech trees averaged 2.1 i 0.13 mm in the last 5
years, 2.10 :1: 0.12 mm in the last 10 years, and 2.0 d: 0.11 mm in the last 15 years (Table
3-1). As no reductions in beech growth were detected, no further analyses were

conducted on the non-beech cores.

Leaf biomass
Mean leaf biomass of all species combined was 289 i 15.0 g/m2 in 2002 and 311
i 16.0 g/m2 in 2003 (Table 3-2). Although differences in leaf biomass were not

substantial, they differed significantly among years (of/E30, t=-2.81, p=0.008). Beech and

62

sugar maple comprised the greatest portion of leaf litter in both 2002 and 2003 (Figure 3-
3).

In 2002, mean leaf biomass of all species combined was significantly greater in
stands with no C. fagisuga infestation than in stands with heavy C. fagisuga (djE2,33;
F=4.60, p=0.017), but not different among stands with low, moderate, or high beech
density (p=0.304). Mean leaf biomass of beech did not significantly differ among stands
with absent, light, or heavy C. fagisuga (p=0.2l3), but was greater in stands with high

beech density than in stands with low beech density (df=2,33; F =0.12, p=0.014).

 

In 2003, mean leaf biomass of all species was not significantly affected by C.
fagisuga infestation (p=0.45 8), nor beech density (p=0.238). Mean leaf biomass of beech
did not significantly differ among stands with varying levels of C. fagisuga (p=0.437),
but was significantly greater in stands with moderate and high beech density than in
stands with low beech density (df=2,48; F =7.64, p=0.001).

Basal area of beech, sugar maple, red maple, red oak, and other hardwoods was
significantly correlated with species leaf biomass of each corresponding in 2002 and

2003 (Table 3-3).

Discussion
Radial growth

Examination of radial growth trends for all beech trees (n=304) did not elicit any
marked reductions in growth attributable to beech bark disease. Only five of the 73 cores
from beech trees infested with C. fagisuga exhibited distinct reductions in growth. Each

was from a different stand, indicating no stand-level reduction in growth. In at least three

63

of our 11 stands with heavy C. fagisuga, signs and symptoms of beech bark disease were
evident (including perithecia of Nectria sp.) and mortality and breakage of mature beech
had begun by the onset of our study. We extracted cores from only the remaining live
trees with C. fagisuga in these stands and likely included the effects of the changing
canopy structure on radial growth in our analyses. Mature beech trees increase radial
growth when growing in open gaps, such as those created by the mortality of a
neighboring tree (Canham 1990, Poage and Peart 1993). The availability of increased
light may have outweighed the effects of beech bark disease on radial growth. Such an
occurrence was documented in New York stands affected by beech bark disease, where
radial growth of surviving beech was highest in stands with greatest mortality (Runkle
1990)

Beech bark disease occurs with greatest frequency on poor sites where trees are
subject to greater physiological stress, particularly when moisture is limited (Lonsdale
1980, Shigo 1964). Under poor conditions, beech may have “missing” rings (Peters
1997). We did not cross-date our cores and therefore did not account for this possible
discrepancy among trees growing under various site conditions.

Beech bark disease had not yet impacted the radial grth patterns of beech trees
sampled from our study stands. Other studies demonstrated a 25-40% reduction in radial
growth of beech that had been impacted by beech bark disease for 8 to 30 years (Gavin
and Peart 1993, Mize and Lea 1979, Runkle 1990). Our study accounts for roughly 10
years of cumulative impact of C. fagisuga. Because C. fagisuga cannot live on dead
tissue, trees in our stands with heavy C. fagisuga infestation were probably not yet

infected with Nectria spp. We cannot assume, therefore, that presence of C. fagisuga

64

 

ill-T

equates to beech bark disease. Long-term monitoring will be necessary to evaluate the
radial growth of trees with C. fagisuga as they are progressively impacted by beech bark

disease and dieback occurs.

Leaf biomass

In both 2002 and 2003, the beech component of mean leaf biomass did not differ
among stands with varying levels of C. fagisuga infestation. In other words, C. fagisuga
had not affected foliar productivity of beech and infested and uninfested stands produced
similar amounts of foliage. In a study conducted in New York stands transitioning from
the Killing Front to the Aftermath Stage of beech bark disease, annual litter production
remained constant, beech foliar biomass did not change, and soil chemistry was not
altered by changes in beech leaf leachate, likely because basal area of regenerating beech
compensated for declines in mature overstory mature trees (Forrester et al. 2003).
Although our study was primarily conducted in stands at the Advancing Front, our results
are consistent with conditions reported afier 15 years of cumulative beech bark disease
impact in the New York stand.

Stand-level C. fagisuga classifications were derived from density estimates made
on individual large diameter trees, which are more readily infested by C. fagisuga
(Houston and Valentine 1987), yet foliage was collected at the stand level and included
leaf production from all classes of trees positioned above the traps. Collecting leaf litter
from trees of all classes and conditions likely included productivity from smaller trees not

infected with C. fagisuga in the understory. As trees succumb to beech bark disease and

65

crowns become sparse, suppressed trees of other species in the understory respond to
increased light levels with rapid growth (Canham 1990).

Published equations are useful for estimating foliar biomass of individual trees
based on dbh along with total aboveground productivity for a given stand (Briggs et al.
1989, Forrester et al. 2003, Ter-Mikaelian 1997). In our study, basal area of red maple,
red oak, and other hardwood species were correlated with leaf mass of respective species.
Basal area for beech and sugar maple, however, were not linearly associated with leaf
mass suggesting long-term studies are needed to assess that relationship between foliage
production with basal area.

The complete etiologic complex of beech bark disease was not conclusively
determined in our study and therefore, conclusions are limited to the impacts of C.
fagisuga infestation. Future studies on the impacts of beech bark disease on Michigan
forests should include Nectria sp. present, as tree vigor measurements have been reported
to be more closely correlated with Nectria sp. than C. fagisuga infestation (Jones and
Raynal 1987).

It is plausible that, by the time of this study, beech bark disease had only recently
begun to impact individual trees and stands. One long-term study in New Hampshire
demonstrated that at low levels of infestation, individual tree growth trends varied widely
and occasionally included overall increases (Gavin and Peart 1993). The results
presented here were largely intended to provide a baseline for comparisons of long-term
data and will be useful in elucidating the period over which any changes in radial grth

and biomass production occur.

66

 

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Table 3-3. Spearman's correlation coefficients (rs) and significance value (p)

for the relationship between basal area (mz/ha) of overstory species and
corresponding leaf biomass from 40 stands in 2002 and 54 stands in 2003.
Correlation coefficients witllp<0.05 are significantly different.

2002 2003
American beech
rs 0.588 0.664
p <0.001 <0.001
Sugar maple
rs 0.587 0.638
p <0.001 <0.001
Red maple
rs 0.632 0.679
p <0.001 <0.001
Red oak
r5 0.761 0.797
p <0.001 <0.001
Other hardwoodsa
r: 0.598 0.696
p <0.001 <0.001

aOther hardwood species included ash (F raxinus sp.), aspen (Populus sp.), basswood

 

(T ilia sp.), birch (Betula sp.), and black cherry (Prunus serotina ).

70

Figure legends
Figure 3-1. Intensive research plot design indicating arrangement of subplots 1-5,
overstory composition measurement transects, individual beech trees selected for long-

term monitoring, and leaf litter traps.

Figure 3-2. Age of beech trees (n=94) plotted against the ratio of mean annual increment

for intervals of a) 5-year, b) 10-year, and c) lS-years.

Figure 3-3. Composition of leaf biomass (g/mz) by species from a) 40 stands in of stands

in 2002 and from b) 54 stands in 2003.

71

 

Key to plot diagram:
X Sample tree

El Leaf litter trap

—_ Overstory basal area sampling transect

O Subplot

72

A) 5-year intervals: 1997-2001 and 1992-1996

 

 

 

 

 

 

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14%

74

APPENDICES

75

 

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«H: S: S: «E m m ON H «H H NN.OH HNO NNH
S: Q: S: S: 3 N OH. m NNH NNwH HNO HNH
S: E: 8: 8: 7H m Om H 02 ON.OH HNO 92
S: Q: S: Q: Q: m ow Q v.2 NN.Nm HNO ONH
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«H: S: S: E: m N ON H 6H H Nm.mH ONO mNH
VOH SH mod 8: m N ON 0 62 mva ONO «NH
SH Hod 3H 8: m H Om o 0.2 OH .3. ONO mNH
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«H: E: E: S: 3 N OH H m H EMVN ONO n H H
«E «\c E: 8: 3m H Om H m .m H EOH ONO OH H
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«E «E «E «E m H Om 0 m.mH mem «NO «2
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«E «E «E «E m N on H N. mH «m .«N «NO Nm H
mH.H NmH Nod «E m O OO O mOH H«.H« «NO HmH
mO.H OHH a«.H «E Z H o« D OOH OO.m« «NO OmH
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«E «E «E «E m O o« H m.mH OmHH mNH NOH
O0.0 m«.O O0.0 «NH m N Om U m.mH «m.O« mNH HmH
H«.O m0.0 OmH «E m H Om 0 m.mH Om.m« mNH OOH
«E «E «E «E Z H O« m NH ««.mH mNH OmH
OHH HmH mNH «E m N Om o H.«H «O.m« mNH m«H
«E «E «E «E Z H Om H 2 m«.mH mNH «OH
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«E «E «E «E m O O« m H.«H «HNH mNH me
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«E «E «E «E Z H OO m 0.0 «mm ONO m«H
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«E «E «E «E Z H O« m H .w ««.« ONO m « H
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«E «E «E «E Q N Om H m H m«.ON ONO m«H
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««.O ««.O m«H «E m H O« 0 «.ON «mNm OmO mmN
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«E «E «E «E Z O O« o m.OH O0.0« «mO O«N
«E «E «E «E m O o« 0 H .ON mO. H« «8 OON
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mNH mmH ««H «E Z O O« U m.mH ««.ON N«O OON
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«E «E «E «E >9 O O« U O.mH mO.mN N«O «mN
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«E «E «E «E m H O« Q «.ON mm.N« N«O «mN
«E «E «E «E >9 O O« H m. m H mm .NN N«O mmN
«E «E «E «E >9 O O« H O.mH mm.mN N«O NmN
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110

Appendix 5
Record of Deposition of Voucher Specimens*

The specimens listed on the following sheet(s) have been deposited in the named
museum(s) as samples of those species or other taxa, which were used in this research.
Voucher recognition labels bearing the Voucher No. have been attached or included in
fluid-preserved specimens.

Voucher No.: 2004-02

 

Title of thesis or dissertation (or other research projects):

IMPACTS OF BEECH BARK DISEASE ON TIMBER AND WILDLIFE RESOURCES
IN MICHIGAN.

Museum(s) where deposited and abbreviations for table on following sheets:

Entomology Museum, Michigan State University (MSU)

Investigator’s Name(s) (typed)
Amy M. Kearney

 

 

Date 7/27/04

*Reference: Yoshimoto, C. M. 1978. Voucher Specimens for Entomology in North
America. Bull. Entomol. Soc. Amer. 24: 141-42.

Deposit as follows:
Original: Include as Appendix 5 in ribbon copy of thesis or dissertation.

Copies: Include as Appendix 5 in copies of thesis or dissertation.
Museum(s) files.

Research project files.

This form is available from and the Voucher No. is assigned by the Curator, Michigan
State University Entomology Museum.

111

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112

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