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ABSTRACT

A STUDY OF VARIATION IN THE RED OAK GROUP
IN MICHIGAN AND NEARBY STATES

By William R. Overlease
The limits of variability and the relationship of Hill

oak (guercus ellipsoidalis E.J.Hill) to other species of oaks

 

in the Great Lakes area have been a problem since the species
was first described in 1899 by Rev. E. J. Hill. Questions
have been raised concerning its relationship to scarlet oak
in particular, and also to black oak, red oak, and pin oak.

An attempt to investigate these relationships and to
come to some understanding of Hill oak as an eCological and
taxonomic unit necessitated a study of the related species
mentioned above as well as the Hill oak taxon itself.

A dual approach was used. First, variation in a number
of field morphological characters was studied for a number of
pOpulations along south to north transects. Three transects
approximately 400 miles long were established from Indiana to
northern Michigan. Forty pOpulations in the red oak groupd
(Erythrgbalanus) were located and studied on these transects.
Thirteen characters were studied in the field during the
summer of 1962 and later in the year two additional characters
were studied. This gave a total of fifteen field characters
as a basis for the study of morphological variation.

The second phase was a nursery study of oak seedlings.

This study was designed to help ascertain whether the variation

in field morphological characters was due primarily to

genetic differences or not. Acorns were collected from those

field populations which matured seed and these collections

were augmented by other collections. The acorns were planted
and seedlings grown under common garden conditions in the

Bogue Forest Nursery in E. Lansing, Michigan. Variation was

studied in twenty-five different nursery characters that were

scored in the 2 years that seedlings were observed. Mass
plantings were also made to study variation from single
female parents growing in different portions of the study
region. One mass planting was made of a suspected hybrid to
test for possible segregation among the seedlings.

The results of the study may be summarized as follows:

1. Though field data gave evidence of discontinuities between
scarlet oak (Quercus coccinea Muenchh.) and Hill oak and
within the Hill oak taxon, this did not appear to be sup-
ported by evidence from seedlings grown under common
garden conditions. The evidence from the nursery appeared
to indicate a continuous south to north genetic gradient
with no sharp breaks. For this reason scarlet oak and Hill
oak are regarded as part of a complex, the scarlet-Hill oak
complex.

2. South—north trends were apparent in black oak (Quercus
velutina Lam.) in both field studies and from evidence of
seedlings grown under common garden conditions indicating

south-north genetic trends.

South-north trends were evident in the red oak complex

(Quercus rubra L. and Quercus rubra var. borealis (Michx.f.)

 

Farw.) in both the field and among seedlings grown in the
common garden though not so pronounced as in the scarlet-
Hill oak complex or in black oak.

Black oak, the scarlet—Hill oak complex, and the red oak

complex appear to have small—fruited northern forms.

A STUDY OF VARIATION IN THE RED OAK GROUP IN
MICHIGAN AND NEARBY STATES

By

William Roy Overlease

A THESIS
Submitted to
Michigan State University
in partial fulfillment of the requirements
for the degree of
DOCTOR OF PHILOSOPHY
Department of Botany and Plant Pathology

1964

ACKNOWLEDGEMENTS

The author wishes to express his sincere gratitude to
the following persons for guidance, aid, and inspiration dur-
ing the progress of this study - Drs. John E. Cantlen, Jonathan
W. wright, Philip J. Clark, John H. Beaman, William B. Drew,
Leslie W. Gysel and Charles R. Olien of Michigan State Univer-
sity, and Drs. warren H. wagner, Jr. and Edward G. Voss of the
University of Michigan. He is also grateful to Dr. Terril D.
Stevens, Chairman of the Forestry Department, Michigan State
University, for permission to use the Bogue Forest Nursery
and to Ira W. Bull and his staff at the nursery for their aid
and consideration. The late Dr. Wendell H. Camp of the Uni—
versity of Connecticut was particularly helpful in the early
phases of this study with information based on his experiences
with Hill oak in Michigan. And finally to Edith, my wife,
whose continual aid in the field work and the nursery plantings
was immeasurable.

I also wish to thank the following c00perators who aided
in this study; E. I. Roe and Staff, U. 8. Forest Experiment
Station, Grand Rapids, Minnesota; Robert F. Patton, University
of Wisconsin, Madison, Wisconsin; Grayling Game Club, Grayling,
Michigan; Staff, Raco Guard Station, U. S. Forest Service, Raco,
Michigan; Clayton Wray, The Michigan College of Mining and
Technology, Sault Ste. Marie, Michigan; Maurice Day, Dunbar
Forest Experiment Station, Sault Ste. Marie, Michigan; Staff

ii

Rose Lake Wildlife Experiment Station, Lansing, Michigan;
Frank L. Johnson, Rockford, Illinois; Roland F. Eisenbeis,
Chicago Forest Preserves District, Cook County, Chicago,
Illinois; and the staffs of the following state parks,
McCormick's Creek State Park, Spencer, Indiana; Spring Mill
State Park, Mitchell, Indiana; Brown County State Park,
Nashville, Indiana; Dunes State Park, Chesterton, Indiana;
Pokagon State Park, Angola, Indiana; warren Dunes State Park,
Bridgman, Nichigan; Muskegon State Park, Muskegon, Michigan;
Wilson State Park, Harrison, iichigan; Benzie State Park,
Honor, Michigan; Fort Wilkens State Park, C0pper Harbor,
Michigan; Traverse City State Park, Traverse City, Michigan;
Sleeper State Park, Caseville, Michigan; Burt Lake State Park,

Indian River, Michigan.

iii

Table of Contents
Part I Introduction
Part II Description of Methods
A. General Methods
B. Field Population Methods
C. Nursery Population Methods
D. Computations
Part III Results
A. General Statement
B. Black Oak
C. Scarlet—Hill Oak Complex
D. Red Oak
Part IV Discussion
A. General Statement
B. Black Oak
C. Scarlet—Hill Oak Complex
D. Red Oak
Part V The Red Oak Group in Michigan
Part VI Conclusions

Part VII Bibliography

iv

18

28

32

33

66

92

115

119

129

141

151

165

167

List of Tables

Table Number

I

II

III

IV

VI

VII

VIII

IX

XI

XII

Site descriptions for stands in which
field p0pulations of black oak were
sampled.

Index of similarity for tree vegetation
in 14 stands containing black oak.

Overall differences in 13 morphological
characters between 16 pOpulations of
black oak.

Page

35

37

39

The number of leaves of black oak collected 48

in each of three leaf outline types by pop
ulation.

Results of black oak pOpulations planted
in 1960 under common garden conditions.

Results of black oak populations planted
in 1961 under common garden conditions.

Spring-fall frost damage as observed in
the Bogue Forest Nursery, E. Lansing,
Michigan, 1961.

Site descriptions of stands in which field
pOpulations of the scarlet-Hill oak com-
plex were sampled.

Index of similarity for tree vegetation
in 10 stands containing the scarlet-Hill
oak complex.

Sum of the number of field morphological
characters in which contrasted pairs of

51

55

64

67

68

scarlet-Hill oak populations did not differ

significantly at the 1% level.

The number of leaves of the scarlet—Hill
oak complex collected in each of four
leaf outline types by p0pu1ation.

A summary of the number of trees in each
population of the scarlet-Hill oak com-
plex with winter buds in each of two main
categories, bud pubescence and bud shape.

V

77

List of Tables cont.

Table Number

XIII

XIV

XV

XVI

XVII

XVIII

XIX

Results of Hill oak and small-fruited
black oak populations planted in 1960
under common garden conditions.

Results of scarlet-Hill oak p0pu1ations
planted in 1961 under common garden
conditions.

A summary of the relationships of three
intermediate populations growing under
common garden conditions.

Site descriptions for stands in which
field populations of red oak were
sampled.

Index of similarity for tree vegetation
in 12 stands containing red oak.

Number of characters in which contrasted
pairs of red oak p0pu1ations did not dif-
fer significantly at the 1% level.

Number of trees in each p0pu1ation of
red oak sampled showing presence or
absence of pubescence on the winter buds.

Results of red oak p0pu1ations planted
in 1960 under common garden conditions.

Diagnostic characters of scarlet and
Hill oak.

vi

Page

82

84

89

94

96

97

104

106

134

Figure

10

11

12

13

List of Figures

Overlay map showing approximate loca-
tion of p0pu1ations studied in this
dissertation.

Length of growing season in days in
Michigan and nearby states.

Field p0pu1ation with black oak and the
scarlet-Hill oak complex. Photo.

Outline drawing showing acorn measure—
ments.

Nursery beds at Bogue Forest Nursery.
Photo.

First year seedling in Bogue Forest
Nursery. Black oak. Photo.

First year seedling in Bogue Forest
Nursery. Scarlet oak. Photo.

POpulation means of morphological
characters of 16 black oak populations
under field conditions.

Population means of morphological
characters of 16 black oak populations
under field conditions.

Population means of morphological
characters of 16 black oak populations
under field conditions.

A comparison of variance of black oak
populations.

Basic patterns of leaf outline for 12
field p0pu1ations of black oak from
Michigan and nearby states.

Number of trees in each basic pattern
of leaf outline for mass nursery plant-
ing of black oak seedlings from Dunes
State Park, Indiana.

vii

Page

14

26

26

42

43

44

45

47

Figure

14

15

16

17

18

19

20

21

22

23

24

26

List of Figures cont.

Page
Number of trees in each basic pattern 61
of leaf outline for mass nursery plant-
ing of black oak seedlings from Traverse
City Fair Grounds, Michigan.
Population means of morphological 70
characters of 10 scarlet-Hill oak p0pu-
1ations under field conditions.
Population means of morphological 71
characters of 10 scarlet-Hill oak p0pu-
1ations under field conditions.
Population means of morphological 72
characters of 10 scarlet—Hill oak popu—
lations under field conditions.
A comparison of variance of the scarlet— 75
Hill oak p0pu1ations.
Basic patterns of leaf outline for 9 76

field p0pu1ations of the scarlet-Hill
oak complex from.Michigan and nearby
states.

Number of trees in each basic pattern of 90
leaf outline for mass planting of scarlet-
Hill oak seedlings from Elkhart, Indiana.

Number of trees in each basic pattern of 90
leaf outline for mass planting of scarlet-
Hill oak seedlings from Grayling, Michigan.

Population means of morphological 98
characters of 13 red oak p0pu1ations
under field conditions.

Population means of morphological 99
characters of 13 red oak p0pu1ations
under field conditions.

POpulation means of morphological 100
characters of 13 red oak p0pu1ations
under field conditions.

A comparison of variance of 13 red oak 102
populations.

viii

List of Figures cont.
Figure Page

27 Representative leaves from populations 103
of Quercus rubra.

 

28 Representative leaves from populations 103
of Quercus rubra var. borealis.

 

29 Basic pattern of leaf outline for seed- 112
lings grown from a suspected hybrid tree
from Wilson State Park, Michigan.

30 Acorns from the 1961 nursery planting 122
of black oak arranged on a south to
north basis. Photo.

3! Acorns from two black oak trees near 122
E. Lansing, Michigan, showing the varia-
tion in size of viable acorns.

31 Pathways of gene flow in the red oak . 156
group in Michigan and nearby states.

ix

Appendix

A

List of Appendices

Species basal area in square feet /
acre by stands in which black oak
p0pu1ations were found.

Species basal area in square feet /
acre by stands in which scarlet—Hill
oak p0pu1ations were found.

Species basal area in square feet /
acre by stands in which red oak p0pu-
1ations were found.

A list of the morphological characters
in the red oak group studied during the
summer of 1960 at the University of
Michigan Biological Station at Pellston,
Michigan.

Black oak population mean and variance
for each morphological field character.

Scarlet-Hill oak p0pu1ation mean and
variance for each morphological field
character.

Red oak p0pu1ation mean and variance
for each morphological field character.

Tests by analysis of variance to indi-
cate which groups of black oak field
populations showed a significant dif-
ference at the 5% and 1% level.

Tests by analysis of variance to indi-
cate which groups of the scarlet-Hill
oak complex field populations showed a
significant difference at the 5% and
1% level.

Tests by analysis of variance to indi-
cate which groups of red oak populations
showed a significant difference at the
5% and 1% level.

Page

172

174

175

177

179

180

181

182

186

188

1
INTRODUCTION
In 1899 Reverend E. J. Hill described an oak species

known today as Hill oak,_guercus ellipsoidalis.l Since that

 

time there has been considerable discussion and debate over
the relationships between this taxon and black oak (92ercus

velutina, Lam.), red oak (Quercus rubra, L. and Quercus rubra

 

 

 

var. borealis,(Michx.F.)-Farw}, scarlet oak (Quercus coccinea,

 

 

Muenchh.), and pin oak (Quercus palustris, Mnenchh.). wadmond

 

(1939) made a study of the relationship between Hill oak and
scarlet oak. Ernest J. Palmer (1942) discussed the red oak
complex in Eastern United States with comments on the deep cup
acorn forms of certain species. Curtis (1959) made several
references to the Hill oak problem in Wisconsin with particular
emphasis on the possibility of introgression between it and re—
lated species. Trelease (1919) discussed the problem of Hill
oak and scarlet oak in Illinois. The problem has thus been with
us for over 60 years but to my knowledge a nursery or common
garden approach to the study of the problem has not been attempted.
This dissertation reports a study of some of the field
variation patterns of five related species of oaks, black, red,
Hill, scarlet, and pin, and where acorns were avilable, the
study of seedling variation under common garden conditions.
The results are helpful in understanding some of the questions

concerning the Hill oak complex and its relationships to other

1All species designations in this dissertation follow Gray's
Manual QEDBotony by M; L. Fernald, 1950.

taxa.

One method of approaching the study of Hill oak would be
to study it in the light of some of the already recognized
variation patterns in related oak taxa. Thus the northern

p0pu1ations of the red oak, (Quercus rubra) have long been

 

known to include a small fruited northern race (9. rubra var.

borealis Farwell (1904), Rehder (1940), Palmer (1942), Fernald

 

(1950), Gleason and Cronquist (1963). My own preliminary ob—
servations suggested that if one compared.Michigan and Indiana

p0pu1ations of Q. velutina, it also has small-fruited northern

 

p0pu1ations. With this background one could then advance a

working hypothesis that Q. ellipsoidalis is a similar small-

 

fruited northern race of Q. coccinea, a species which it re—

 

sembles in many morphological features, and the taxon to which
many Hill oaks were originally assigned. The variation patterns
in all three complexes could then be studied both in field popu—
lations and in seedlings grown from these p0pu1ations in a
common garden or nursery.

Three major transects were established along a south to
north axis. These transects were approximately 400 miles long and
crossed one of the major American vegetation—soil boundaries, that
separating the Hemlock-White Pine-Northern Hardwoods from the
more southern Beech-Maple forests, (Braun, 1950). On these three
transects 40 field p0pu1ations were studied intensively. Acorns

were collected from many of these p0pu1ations and germinated

3
under common garden conditions in the Bogue Forest Nursery
in East Lansing, Michigan. Because of erratic acorn produc-
tion in some field sites, other p0pu1ations were also used.
For comparison purposes, acorns from as far away as WiSconsin
were grown. In all, acorns from 216 female sources were
planted in the nursery. These female parents represented 34
different p0pu1ations, 25 of which came from populations
studied in the field.

Specimens from each population.nepresented in this study
are deposited in the Beal-Darlington Herbarium at Michigan
State University, E. Lansing, Michigan. Three specimens from
each p0pu1ation were selected; one to represent the average
in population variation and the other two ia-the variation

extremes.

4

Fig. l. Overlay map (on following page) showing approxi-
mate location of populations studied in this
dissertation. ,

Black Oak Populations

Mc
Ek
Du
Po
Rl
Mg
Pt
W1
81
Et
Ho
Ht
Tp
Tf
Al
Ir
Ro

McCormick's Creek State Park, Ind.
Elkhart, Ind.

Dunes State Park, Ind.

Pokagon State Park, Ind.

Rose Lake Wildlife Station, Mich.
Muskegon, Mich.

Pentwater, Mich.

Wilson State Park, Mich.

Sleeper State Park, Mich.

East Tawas, Mich.

Honor, Mich. (1)

Honor, Mich. (II)

Traverse City State Park, Mich.
Traverse City Fair Grounds, Mich.
Alpena, Mich.

Indian River, Mich.

Rochester, Ind. (Nursery p0pu1ation only)

Scarlet-Hill Oak Populations

Br
E1
Th
Rk
Sk
R1
W1
Fw
wr

Gy
Red Oak

Po
Wd
En
Mp
Bp
Ho
Ap
Bl
D1

Brown County State Park, Ind.
Elkhart, Ind.

Thornton Woods, Chicago, Ill.
Rockford, Ill.

Stockbridge, Mich.

Rose Lake Wildlife Station, Mich.
Wilson State Park, Mich.

Farwell, Mich.

Wisconsin Rapids, Wisc.

Grayling, Mich.

Populations

Pokagon State Park, Ind.
Warren Dunes State Park, Mich.
East Lansing, Mich.

Muskegon State Park, Mich.
Benzie State Park, Mich.
Honor, Mich. (Dunes)

Alpena, Mich.

Burt Lake State Park, Mich.
Douglas Lake, Mich.

+4

Dn — Dunbar Forest Station, Mich.

Ra - Raco, Mich. _

Ch - Copper Harbor, Mich.

Ss - Sault Ste. Marie, Mich. (Nursery p0pu1ation only)

Tension Zone — in Michigan it is an area where northern
and southern flora, fauna, and soil types appear
to intergrade, Veatch (1932), Dice (1931), Potzger
(1948), Braun (1950).

 

Fig. 2. Length of growing season in days in
Michigan and nearby states. Redrawn
from Climate and Man, U.S.D.A~ Year—
book, 1941.

 

 

Part II Description of Methods

A.

General Methods
Field POpulation Methods
Nursery Population Methods

Computations

7
A. GENERAL METHODS

The summer of 1960 was spent perfecting field methods
and studying oak populations in the northern Lower Peninsula
of Michigan. In the fall of 1960, acorns were collected from
103 parental sources representing 21 field p0pu1ations and
planted in the nursery. .A preliminary field study of winter
buds was made in the early spring of 1961 on three transects,
one along the east coast, one along the west coast, and one
through the center of Michigan's Lower Peninsula. In late
spring of the same year a field trip was made to observe frost
damage to oaks in the northern part of the Lower Peninsula.
During the summer of 1961, field methods were further perfected
and in the fall of 1961, acorns from 111 parents representing
14 p0pu1ations Were collected and planted in the nursery. .A
field trip in the spring of 1962 was again made to observe
frost damage to oaks in northern Lower Michigan.

The main field studies of morphological variation were
made during the summer of 1962 on south to north transects
along which 40 p0pu1ations were sampled. The studies were be—
gun in late June and continued into October. A third field
trip to observe spring frost damage in northern Lower Michigan
was made in 1963.

Scoring and maintenance of the nursery p0pu1ations con-
tinued throughout the study. Similarly, the data processing

was more-or-less continuous as a rather large amount of material

8
had to be synthesiszed and reduced to useable form.

Since an understanding of certain of the results requires
detailed knowledge of the methods used, these methods are
described more fully below.

B. FIELD POPULATIONS METHODS

South—north transects were established from south-central

Indiana to northern Michigan and on them populations of the

following taxa of oaks were studied, Quercus velutina, Q.

 

coccinea-Q. ellipsoidalis complex, Q. rubra and Q. rubra var.

 

 

borealis. Natural p0pu1ations were selected on the basis of
the following criteria:

1. Populations of the same species should be located
at approximate 50 mile intervals along the transect.

2. At least 15 individual mature trees of the oak
species should be available for sampling.

3. The p0pu1ations should not be excessively hetero—
geneouSgenetically. In other words an attempt was
made to avoid apparent hybrid swarms.

4. The p0pu1ation sampled should be on a uniform soil
type.

5. The p0pu1ation should be reasonably accessible for
subsequent study, and where possible be located in
a state park or other protected area.

Not every p0pu1ation selected fit all these criteria but

where possible it was adhered to. After a p0pu1ation was

Fig. 30

A woods near Rockford, Illinois,
containing p0pu1ations of black
oak and of the scarlet-Hill oak
complex. A scarlet-Hill oak is
in the foreground and a black oak
in the background.

 

10

selected, a line transect was laid out through it along which
individual trees were sampled. Where p0pu1ation size permitted,
trees for scoring were randomly selected from the first one
hundred mature trees of the species to be sampled encountered
on the transect. A.mature tree is defined as one thought
capable of bearing acorns, but the presence of acorns under a
tree was not used as a criterion. These trees were consecu-
tively numbered and tagged. The distance from the transect
that trees were tagged depended on the stocking of the p0pu1a-
tion. It varied from 50 meters in one population to an average
of 15 meters in well stocked p0pu1ations. One hundred plastic
tabs were given the same numbers as the trees, shaken together
in a container, and selected one at a time until the desired
sample size was reached. The trees were numbered and scored
for designated morphological characters. The size of sample
which varied from 9 to 34 trees was determined by the size and
apparent variability of the population. In uniform populations,
as judged by a preliminary survey of its field characteristics,
approximately 15 individuals were sampled. In variable popula-
tions or suspected variable p0pu1ations about 25 trees were
sampled. If a p0pu1ation proved to be more variable than was
expected, it was a simple matter to increase the sample size by
randomly choosing additional tabs.

Scoring of Morphological Characters

Preliminary field studies in 1960-61 led to a selection of

ll
morphological characters that would be the most useful in
separating species in the red oak group in Michigan. Thirty-
one characters were intensively investigated and six ratios
or other combinations were made with various pairs of these.1
After careful consideration of the resulting data, including
statistical treatment, 13 characters were selected as being
the most likely to prove useful for this study. These 13
characters then became the basis for the field p0pu1ation study
during the summer of 1962.

Of the 40 populations which were studied intensively in
the field, 16 were black oak, 14 were red oak, and 10 were of
the Hill-scarlet oak complex. The following is a description
of the methods used to study each individual character.

53932, In the author's experience, the acorn provides a
series of sensitive and consistent field characters for working
with oak p0pu1ations. In this study the acorn with its cup
was scored for 9 characters. Since considerable emphasis is
placed on acorn characters, the selection of the acorn to be
scored is critical. The following method was used to minimize
bias in this selection. Mature acorns were collected from the
ground beneath that part of the crown area in which there was
little possibility of contamination by acorns from nearby trees.
After the acorns were collected, the maximum sample being 25,

they were laid in a row and numbered. The acorn to be scored

1See appendix B for a list of these characters.

12

was selected from the row by drawing at random, a numbered tag,
from a container. The abundant immature acorns on the ground
under most trees were not considered in selecting mature acorns.

Acorn height with cup_attached (total acorn height).. The
combined length measured in millimeters, from the tip of the
acorn to the t0p of the acorn cup with the cup in place on the
acorn.

Acorn Length. The maximum length of the acorn (minus the

 

cup) measured in millimeters.

Acorn width. The maximum width of the acorn measured in

 

millimeters.

Acorn shgpe. The acorn width divided by the acorn length

 

multiplied by 100. This gave an index of the acorn shape. A
narrow acorn had an index number considerably less than 100,
and an acorn wider than long had an index more than 100. A
perfectly spherical acorn had an index of 100, that is, the
length and the width were the same.

Acorn size. The sum of the length plus the width was used

 

as an approximation of acorn size.

Acorn cup height. The height of the acorn cup measured in

 

millimeters.

Ratio of cgp height to total acorn height. The acorn cup

 

height was divided by the total acorn height and multiplied by
100. This measurement provided an approximation of the degree

to which the acorn cup covered the acorn. For example, an index

13
value of 50 indicated that half of the acorn was covered by
the cup.

Presence of fringe on the acorn cup. An index was es—

 

tablished with values from 1—10. An acorn with very prominent
fringe on the lip of the cup was scored 1. .A cup with no fringe
on the lip was scored 10. Intermediate values were scored with
appropriate numbers between 1 and 10.

Acorn cup scale shgpe. An index from 1 to 10 was established

 

as follows:
2 - Cup scales with rounded tips (typical of red oak).

5 — Cup scales with acuminate tips (typical of the
Hill-scarlet oak complex).

8 - Cup scales with obtuse’tips (typical of black oak).
Intermediate shapes were given values between these established

criteria.

14

Fig. 4. Acorn measurements

A or A’ — Total acorn
height.

 

 

 

 

 

B or Bt - Acorn cup
T height depend-
A A? ing on shape of
l l acorn cup.

C - Acorn cup fringe.

 

 

15

Twig pubescence. An index was established from 1, for

 

very heavily pubescent twigs, to 10 for perfectly glabrous
twigs. Twigs intermediate in their pubescence were scored
with intermediate numbers. Shade branches within 20 feet of
the ground were selected and their twigs observed for the
amount of pubescence. At the beginning of the study a 50 foot
pole cutter was used to collect twigs exposed to bright sunlight,
but this proved to be too time consuming and the long pole was
difficult to handle in the field. A.15 foot pole cutter was
used to obtain all materials in the 1962 field season. Each
tree was scored on the basis of the twig that showed the great-
est degree of pubescence. This procedure was necessary as there
was considerable variation in pubescence on an individual tree
and even on an individual branch. The amount of twig pubescence
of an individual tree appeared to depend on the degree of ex—
posure, twig maturity, and vigor of twig growth. A tree that
showed heavy pubescence on its protected or less vigorous growth
and none on its exposed vigorous twigs would be classified with
a low index number indicating heavy pubescence. This type of
scoring resulted basically in a comparison of the presence or
absence of pubescence on a tree but did not give much informa-
tion on degree of pubescence.

Inner bark color. .A preliminary study indicated that four
basic colors with intergrades would provide an index to inner

bark color. The selected colors according to the Munsell (1929)

16

color coding system were:

Field
Grade Appearance Munsell Code
2 Tan 2.5 Y 9/4
5 Pink 5.0 YR 8/4
8 Yellow 2.5 Y 9/8
11 Orange 2.5 Y 7/8

In the interest of uniformity all bark samples were taken at
4% feet from the ground on the east side of the tree between
bark ridges. The bark color varied slightly depending on
whether the sample was taken between heavy bark ridges or through
them. Height and exposure appeared to have no effect.

The numerical scoring for inner bark color, as well as
that for acorn cup scale shape, represent an attempt to place
on a linear scale a condition that is fundamentally 3 dimension-
al and the results suffer from this limitation. However, it is
felt that in spite of this, the categories provide useful data.

Tree form. A.tree with extremely fine basic branching

 

was given a value of l. A.tree with very coarse, Open branch—

ing was given a value of 10. In actual practice the trees

varied in scoring from 2 to 9. It was the intention in scoring

this character to estimate the inherent branching character-

istics (basic branching) and not the amount of branching caused

by frost and insect damage or other obvious environmental factors.
Twig_britt1eness. The author is indebted to the late wen—

dell H. Camp for the suggestion that the lower branches of cer-

17
tain species in the red oak group, in particular Hill oak,
exhibit differences in brittleness. A twig that could be
bent double without snapping was given a value of 1, and a
twig that snapped readily on bending was given a value of 10.
Intermediate numbers were given to intermediate conditions.
There was Variation in brittleness on individual trees which
appeared related to shade and sun conditions. Therefore,
when possible, sun branches from the outer crown were used for
scoring. In order to avoid problems in the variability in
hardness of the recent growth, two year old twigs were used
for this test when obtainable. In some of the very tall
southern populations only trunk sprouts were obtainable with
the equipment used in this study.

Leaf outline. A leaf in bright sunlight and a leaf in

 

the shade were selected from each tree where available with
the equipment used. Branches were obtained using a pole cutter
and the first mature undamaged leaf at the end of the branch
was selected. This character was not handled statistically.

Winter buds. Terminal winter buds were collected from

 

individual trees with a pole cutter in the early spring of 1963.
When possible terminal buds on vigorous branches in bright sun—
light were obtained as it was found that the size of the buds
varied considerably in relation to the sun exposure of the

branches. Terminal buds in bright sunlight often were 2 to 3

times the size of terminal buds in the shade. This character

18
was not studied statistically.
C. Nursery Population Methods

The p0pu1ations chosen for comparison under nursery or
common garden conditions were determined by the presence or
absence of reasonable sized crOps of useable acorns. At least
24 viable, unparasitized acorns were necessary from each parent
tree. At the beginning of the study it was intended to collect
acorns from those p0pu1ations in which field morphological
studies had already been completed. It was soon observed from
a study of the remains of past and current acorn cr0ps under
the trees, that many of the field populations were so erratic
in acorn production that they could not be depended upon to
produce enough usable acorns for nursery study in any reasonable
time. It then became necessary to locate fruiting p0pu1ations
that approximated the criteria established for the selection of
the field p0pu1ations (see p. 8). After locating and planting
acorns from these populations in the nursery for two seasons
(1960 and 1961), the areas where these nursery p0pu1ations were
obtained became the basis for the location of the main field
morphological studies conducted during the summer of 1962.

The erratic nature of acorn crops led to the partial waiv-
ing of the original stand selection criteria so that populations
with good acorn cr0ps were chosen even though they might be
only 15 miles apart. The presence of good acorn crops thus be-

came the prime selection criterion.

19

Fig. 5. Nursery beds at Bogue Forest
Nursery, E. Lansing, Mich.
showing one year old seedlings
to the left of the stakes and
two year seedlings to the right.

July 1962

 

20

At the beginning of the study, an attempt was made to
obtain acorns from c00perators in other areas within the range
of Hill oak. From 9 requests mailed out, four sets of acorns
were obtained by mail, one of which did not germinate and
another was erratic in its germination. No further attempt
was made to obtain seed material by mail.

The matter of choosing which of the acorns lying under a
particular tree should go into the nursery was not difficult.
Only in a very few cases was the unparasitized crop large
enough to make selection a problem. In those cases (involving
mostly red oak), acorns were selected in a circle around the
tree or from a particular sector of the circle in which it was
felt there was no contamination of acorns from nearby trees.

Some trees with good crops had to be eliminated as it was
impossible to separate the acorns from those of other nearby
trees with certainty. The acorns of some black oak trees and
:occasionaliy, of whole p0pu1ations of black oak were unusable
due to near 100% parasitism by insects. Infested acorns were
identified by a very minute black spot left on the outside of
the acorn where the insect injected an egg or eggs into the
acorn. There appeared to beP¥LnCLp§l¥tho kinds of infesta-
tion, those producing weevil-like grubs (nut weevils, Curculio
sp.) and those producing groups of hard cysts inside the acorn

(Species of Cynipids). Good acorns could generally be identi-

fied by their color and the presence of stripes. The acorns

21
of black oak, pin, Hill, and scarlet oaks were distinctly
striped with dark lines and those of red oak were occasionally
striped. When the acorns lost their stripes and became yellow-
ish—brown in color, they were generally no longer viable or
were at least a poor risk. Moisture conditions in the forest
floor appeared to be a major factor in influencing the length
of time an acorn remained viable. .Acorns on dry Open sandy
areas appeared to suffer considerable loss of viability due
to their drying out. Korstian (1927) has indicated the impor—
tance of moisture content as a factor in acorn germination.

In 1960 the acorns were planted as soon as possible after
collecting. This varied from the same day to two or three days.
In one or two cases it was as long as a week. The acorns were
placed in paper sacks with a handful of moist Soil from the site
and kept in a refrigerator while awaiting planting. No loss
was detectable from storage treatment for this planting. In
1961 most of the acorns were stored in plastic bags at a tem-
perature just above freezing for a week to as long as 30 days
with no detectable loss of germination.

In 1960 individual acorns were planted by hand and covered
with about 1 inch of soil. In 1961 individual acorns were
planted by driving them into the soft, prepared soil of the
nursery bed with a hammer blow. Though efficient, this latter
method did not result in well covered acorns and nearly 35%

was lost to animal activity, thought to be the abundant pheasants

22
in the area.

After planting, the beds were covered with one inch of
sawdust for protection from frost heaving. The 1961 nursery
bed listed as "Group B" in the results on the following pages
was not mulched. This provided a useful comparison to the
mulched beds as acorns from the same female parent trees were
in both beds.

In order to keep weeding at a minimum, a commercial chemi—
cal weed killer (Amizine by trade name=simazine—amino triazole)
was used with good results. Care was taken not to allow any
of the weed killer to reach the leaves of the young oaks.

The experiment was laid out in a randomized block design,
with four replicates, in seedbeds four feet wide. A plot con-
sisted of one 6—to—8-tree row perpendicular to the long axis
of the seedbed. In the 1960 planting, 6 acorns from one female
parent were equally spaced in each row. In the 1961 planting,
8 acorns were planted in each row. Thus for each female parent
there were 24 seeds in the 1960 planting (6 seeds per row with
4 replicates) and in the 1961 planting, 32 seeds (8 seeds per
row with 4 replicates).

A nursery map was made of all row locations and was the
final authority on plot locations. However, for ready field
reference, a plastic stake bearing the source number was placed

at the end of each row.

Periodic surveys were made of the nursery plantings to

23
detect possible morphological, physiological, and ecological
differences among the seedlings. When a difference was found
that appearedto be related to seed origin, the entire nursery
planting was scored for this character.

Scoring was carried on throughout the year following ger-
mination and for one additional year in the case of the p0pu—
1ations planted in the fall of 1960. The seedling study was
terminated in the early spring of 1963 when the nursery was
moved to a new site.

During the course of the study, 34 p0pu1ations of oaks
from 214 female parents were studied in the nursery as follows,
black oak — 14, red oak — 7, Hill oak - 10, pin oak — 2, and
scarlet oak — l.

The following is a description of characters scored in the
nursery.

Seedling emergence. The number of seedling shoots that

 

had emerged were counted for the 1960 planting on the following
dates: 5/19/61, 5/22/61, 5/26/61, 6/2/61, 6/10/61, 6/18/61,

and for the 1961 acorns on 5/20/62, 5/27/62, 6/2/62, 6/9/62.
The average number of days during which emergence took place
was calculated for each row from the date of first germination
which was 5/19/61 for the 1960 planting and 5/20/62 for the

1961 planting. The resulting figure was known as "the average

24
emergence time".

Frost damage. The number of trees in each row that were

 

damaged by frost on May 26, 1961 was scored on June 2, 1961
and expressed as a percent.

Percent re-sprouting. The number in each row that showed

 

forking of the growing shoot following frost damage was scored
June 10, 1961 and expressed as percent.

Leaf margin. A scale was established as follows:

 

Scale used in 1961. Scale used in 1962

smooth margin
- wavy margin
dentate margin
lobed margin

2 - smooth margin
5 - wavy margin
8 - dentate margin

\D\lJ-\N

Row averages were recorded for leaf margin characteristics on
June 18, 1961 for the 1960 planting and on July 10, 1962 for
the 1961 planting.

Spring leaf color. The average leaf color in each row

 

was scored on June 16, 1961 according to the following scale:
2 - leaves green, 8 - leaves bronze-red.

Leaf angle. The angle formed by the leaf blade with a

 

horizontal plane at the base of the petiole was visually es-
timated in degrees on July 5, 1961, 1961 (June 26, 1962 for
the 1961 planting) and recorded as the row average. Only
positive angles (i.e. leaves tilted upward from the stem)

were scored using the following scale: l-10°, 2-20°,-3—30°,

25
etc. (See Figs. 6 and 7).

Leaf lengthigLeaf width, Leaf shape, Leaf size. The
largest leaf blade from the first flush of leaves (the origin-
al flush is designated as first flush) for each row was se-
lected and measured in millimeters at its longest point for
length, and at its widest point for width. Leaf bristles and
leaf petiole were not included in the measurements. These two
measurements were multiplied to give an approximation of leaf
area. The width was divided by the length to give an approxi-
mation of leaf shape. The 1960 planting was scored for these
characters on July 5, 1961 and October 19, 1962. The 1961
planting was scored on October 20, 1962.

Fall frost damage. The percent of leaves in each row that

 

was damaged by frost on October 1, 1961 was recorded October 5,
1961.

Percent ggeen leaves. The percent of green leaves present

 

for each row was recorded on November 8, 1961 for the 1960
planting and on October 8, 1962 for the 1961 planting.

Percent rabbit damage. The number of stems in each row

 

which had twigs removed by rabbit browsing was counted April
28, 1962. Most of the damage took place during the 1961—62
winter.

Degree of leafingiout. The degree to which each row of the

 

1960 planting was leafed out was scored on May 5, 1962 by the

following scale: 1 - no evidence of expansion of shoots or buds,

26

Figs. 6 and 7. First year seedlings in Bogue
Forest Nursery showing leaf
angle and leaf margin. July 1962.

 

Fig. 6 Black oak

 

Fig. 7 Scarlet oak

27
5 — leaves in extended shoot stage, 10 - leaves expanded.

Percent seedlings in second flush. The number in each
row that had initiated a second burst of new leaves for that
season was observed on July 5, 1961 and June 11, 1962 for the
1960 planting and on June 13, 1962 for the 1961 planting.

Intensity of red color in petioles. The color of leaf
petioles was recorded for each row on June 12, 1962 uSing the
following scale: 1 - none of the trees with red petiole bases,
5 — most of the trees with red petiole bases but not bright
red, 10 - all leaves with bright red petiole bases.

Percent of rows with red petiole bases. The presence
or absence of petioles with red bases for each row was record-
ed on June 9, 1962.

Percent of seedlings with pubescent twigs. The number of
trees in each row with pubescent twigs was observed June 27,
1962.

Presence of pubescence on second flush shoot. Each row
was scored on June 26, 1962 using the following scale: 1 - shoot
heavily pubescent, 5 — shoot with slight pubescence, 10 - shoot
with no pubescence.

Leaf pubescence. The presence of pubescence on the under—
side of expanded second flush leaves was scored July 10, 1962
using the following scale: 1 - heavy pubescence on all leaves,
5 — slight, but evident pubescence in all or most of the trees,

10 - no leaves pubescent.

28

Percent of seedlings with red leaves. The number of
seedlings in each row with red colored leaves was observed
October 4, 1962.

Percent of the leaves lost. An estimate was made on
November 6, 1962 of the percentage of the original numbers of
leaves which had been lost by that date.

Percent of leaves remaining. An estimate was made on
December 3, 1962 of the percent of the original number of
leaves remaining on the seedlings.

Height of trees. The total height of the three tallest

 

trees in each row was measured. On November 20, 1961 the
height was measured in centimeters. In mid November 1962
measurements were made to the nearest inch for the 2 year old
seedlings and to the nearest quarter-inch for 1 year seedlings.
The average was recorded for each row.

Percent of trees emerged on May 20, 1962. The percent of
each row that had emerged by May 20, 1962 (for the 1961 plant-
ing) was calculated from the total emergence for each row.

D. Computations

Field_pgpulations. For each trait studied the population
2)

 

mean (X), and the p0pu1ation variance (8 was calculated. Then
an analysis of variance was performed for which the degrees of
freedom were as follows: between populations -- P-l; within

p0pu1ations -- N-l-(P-l); total -- N-l, where N and P are re—

spectively the total numbers of trees and of populations studied.

29
Sometimes several such analyses were performed for the same
characteristic in order to test homogeneity among populations.
Suppose that the first analysis included the three southermost
p0pu1ations and showed the presence of no significanCebetween—
p0pu1ation differences. A fourth was included and between—
population. differences were significant. It was then concluded
that the first three did not differ but that the fourth one did.
By means of such groups of analyses, a more rational grouping
of p0pu1ations could be effected. The results are essentially
similar to those which would have been obtained by use of the
"studentized range" method.

Chi-square analyses were used to determine the significance
of differences in numbers of black oak trees damaged by frost.

An "index of difference" among populations was calculated
in the following manner. Populations of a species were compared
two at a time for each field character using the analysis of
variance. The sum of the number of characters in which they were
significantly different at the 1% level was shown in tabular form
with 13 being the highest score obtainable.

An "index of variability" was calculated in the following
manner for each population. For each character the p0pu1ation
with the least variance (52) was considered as standard, and the
ratio of the variance in a given population to that standard

variance was computed. If the ratio (F value) was significant

at the 1% level, the population in question was given a score

30
of l for that character; otherwise it was given a score of 0.
The highest possible score was 13; the highest score obtained
was 8.

Nursery pgpulations. Row (= plot) means were used as items
because of the practical impossibility of separating within—plot
variation into environmental and genetic components. Seedlot
means were calculated from the 4 plots (replicates) of each
seedlot; p0pu1ation means were calculated from the_seedlots
collected within each population.

For each characteristic an analysis of variance was per—
formed for which the degrees of freedom were as follows: where
S, P, and R were the numbers of seedlots, p0pu1ations, and
replicates respectively.

Between seedlots

 

‘Within populations (S—l)-(P—l)
Between populations P-l
Seedlot X rgplicate SR-(S-l)-1

Total SR-l

The "Within-p0pu1ation" term.was used as an error term when
testing significance of between—population differences; the
"Seedlot X replicate" interaction was the error term when test-

ing the significance of within—population differences.

Part III Results

A.

General Statement
Black Oak
1. Field Studies
2. Nursery Studies
Scarlet-Hill Oak Complex
1. Field Studies
2. Nursery Studies
Red Oak
1. Field Studies

2. Nursery Studies

31

32

A. General Statement

The large and complex system of populations making up

,Quercus velutina, the Q. rubra-Q,rubra var. borealis complex,

 

 

 

 

and the dercus coccineaaQ. ellipsoidalis complex in the study

region cannot be adequately characterized by the 40 p0pu1ations

examined.

The aim was to examine field populations for trends

in variation and to test whether similar trends, though not

necessarily in the same characters, occurred in seedlings from

these p0pu1ations when grown under common garden conditions.

For each of the three basic species groups in this study,

the following data are provided.

A.

A site description for each locality where a p0pu1a-
tion was examined.

"Indexes of Similarity" showing the relative similar—
ity of the tree vegetation in the stands in each
species group.

The p0pu1ation means for each of 13 field characters
for each group; presented with the p0pu1ations arranged
on a south to north basis.-

A summary of field populations indicated to be hemo—
geneous by the methods used in this study.

An index of variance among the field p0pu1ations.

The p0pu1ation means for each of the seedling characters
for each p0pu1ation studied in the nursery; presented

with the p0pu1ations arranged on a south to north basis

33
by species group.
G. A.summary of the trends or lack of trends in the
nursery data by species group.
B. Black Oak
The data presented below indicate that black oak p0pu1a—
tions are not homogeneous, either when studied as field p0pu—
lations or as prOgeny from these p0pu1ations under common
garden conditions. South to north trends are suggested by
both the field and nursery studies. Some p0pu1ations appear
to be intermediate between species in the red oak group and
progeny from these populations display intermediate features
in the nursery.

1. Field Studies. The stands sampled occupy three

 

transeCts (Fig. 1), one on the west coast of Michigan's Lower
Peninsula and consisting of Dunes State Park, Indiana, (on the
Michigan-Indiana border), Muskegon, Pentwater, Honor I and II,
Traverse City State Park, and Traverse City Fair Grouds; one
on the east coast and consisting of Sleeper State Park, Tawas,
and.Alpena; and one up the center of the peninsula including
Indiana p0pu1ations and consisting of McCormick's Creek State
Park, Indiana, Elkhart, Indiana, Pokagen State Park, Indiana,
Rose Lake, Wilson State Park, and Indiana River.

The vegetations of which the oak populations were species
components are compared using the index of similarity, Curtis

(1959) in Table II. In spite of the obvious difference of the

34
coniferous forest element in the northern stands, there appeared
to be no consistent pattern of difference between the populations
or no evidence of a break in general species relationships at
the "tension zone" (See. Fig. 1). Since only 10 sample points
were used to determine basal area in each stand, these results
must be regarded with caution in comparing stands.

Looking at expressions of heterogeneity in black oak, it
is to be noted that in each of the 13 characters studied,
significant differences (1% level, analysis of variance) were
found among the p0pu1ations. In order to test the degree of
the differences between the p0pu1ations, the variances of each
of the p0pu1ations for the 13 characters were compared and
summarized (Fig.11). It is readily evident that certain p0pu—
1ations (Wilson State Park, East Tawas, Honor II, and Alpena)
were more variable than others. The most variable ones were
from the north.

There were general south—north trends in the following
field characters (See. Figs. 8, 9, 10):

Total acorn height (acorn height with cup attached)
Acorn length

.Acorn width

Acorn size (length plus width)

Acorn shape (width/length)

Acorn cup height

Ratio of acorn cup height to total acorn height
Presence of fringe on acorn cup

Inner bark color

In this dissertation a trend is said to be present when there

is a clear tendency for p0pu1ation means when arranged on a

35
Table l. Site descriptions for stands in which field popu-
lations of black oak were sampled. The p0pu1ations
are listed from south to north. a/ indicates p0pu—
1ations that are represented in the nursery.

Population name Transect
and size. Location length in Soil type
meters.
a/McCormick's Creek NW% NE% Sec. 22 275 Frederick Silt
State Park, Ind. R3W, T 10 N
(22) Owen County, Ind.
a/Elkhart, Ind. SE% NW% Sec. 13 200 Plainfield Loamy
(18) R4E, T 37 N Sand
Elkhart County, Ind.
a/Dunes State Park SW% NE% Sec. 13 200 Bridgeman Fine
Ind. R6W, T 37 N Sand
(25) Porter County, Ind..
Pokagen State NW% SE% Sec. 34 240 Boyer Loamy
Park, Ind. R13E, T 38 N Sand to Boyer
(l6) Steuben County, Ind. Sandy Loam
a/Rose Lake Wild~ Nk SE% Sec. 24 975 Boyer Loamy
life Experiment RlW, T 5 N Sand
Station, Mich. Clinton County, Mich.
(3o)
a/Muskegon, Mich. W% NE% Sec. 35 325 Grayling
(21) R17W, T 12 N (Loamy)
Muskegon County, Mich. Sand
a/Pentwater, Mich. NE% SE% Sec. 2 150 Rubicon Sand
(18) R18W, T 16 N
Oceana County, Mich.
a/Wilson State W% NW% Sec. 21 175 Grayling
Park, Mich. R4W, T 19 N (Loamy)
Clare County, Mich. Sand

Sleeper State
Park, Mich.
(26)

East Tawas, Mich.
(20)

5% NE% Sec. 20 475
RllE, T 18 N
Huron County, Mich.

SW% Sec. 5 300
R9E, T 23 N
Tosco County, Mich.‘

Rubicon Sand

Rubicon Sand

Table I, cont.

POpulation name
and size

a/Honor, Mich.
(29)

36

Site descriptions for stands in which field
populations of black oak were sampled. The
p0pu1ations are listed from south to north.
a/ indicates populations that are represented
in the nursery.

Transect
Location length in Soil type
meters
I NW% NW% Sec. 23 500 Deer Park
R15W, T 27 N Fine Sand

Benzie County, Mich.

Honor, Mich. II NW% NW% Sec. 23 1175 Deer Park

(12)

a/Traverse City

R15W, T 27 N Fine Sand
Benzie County, Mich.

W% E% Sec. 8 175 Rubicon Sand

State Park, Mich. RlOW, T 27 N

(18)

a/Traverse City
Fair Grounds,
Mich.
(19)

Alpena, Mich.
(20)

a/Indian River,
Mich.
(34)

Grand Traverse
County, Mich.

3% NW% Sec. 2 150 Rubicon Sand
RllW, T 27 N

Grand Traverse

County, Mich.

NE% NW% Sec. 11 525 Rubicon Sand
R7E, T 30 N

Alpena County, Mich.

NW% SW% Sec. 27 750 Rubicon Sand
RZW, T 35 N

Cheboygan County,
Mich.

37

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NN. mN. mN. NH. mo. x .xuwm mumum ummmmHm

“H we as am um Hm H3 um w: Hm on an an

.mHmmn cuuoc ou cuDOm n Go Omuonuo GOHuwHSmom .muaHom deEwm OH cuHB
.AwaH .cwsmnammouov Oocuoe coHHHmuuHm mcu waHmS pochuco mmumEHumo mono
mewn Eoum nouSdEoo mum mooHvaH one .xwo HomHn wchHmucoo museum «H GH
coHuwumwo> mouu Mom .AHmmOH .mHuhDOv n x w \ 3N n m kanwHHEHm mo xovcH .ucoo .HH mHan

39

NH

HH
NH
HH

HH

NH

HH

anDQom

OH

HH

OH

OH

H¢

MB

HH

OH

HH

as

NH

OH
NH

OH

NH

um

OH

HH

NH
NH
HH

OH

NH

om

OH

HH

HH

um

OH R x

O OH NH O

Hm H3 on we

AQHV .con .xumm muwum zuHO omum>mufi

O O

Hm om

Harv HH amaze .gon .uoco:
Aomv H mmcsn .son .Hocom

Humv .OUHE .mm3ma .m

AHmv .con .xnmm muwum HodomHm
Hsz .ron .xuwm mumum comHHz
Humv .ron .umuwsucmm

szv .non .cowoxmsz

HHmv .ron .coHumum name
-Hpmaxm mmHHuHHz «awn mmom

Aomv.ocH ..xm .um cowmxom

x Hsnv.ecH ..xm .um mmcsa
« x Axmv .ocH .uumanm
Hozv .eaH ..xm

m n .um xmmho mzonEHoooz
Sm Hm COHuwHDmom

.mHmwn canon ou nuDOm n no mocmnvmm COHu
.AmocmHHw> mo mHmmenwv Hw>mH NH mcu um nocuocw Eoum UmHoHMHO coHuwH
lance oco :oHcs cH mumuomnwco mo unease ocu moumoHch Hogans comm .xwo HomHn
mo mcoHuwHDmom OH cooBumn muwuomumco HmonOHocaHoE mH CH mmocoummeO Hkum>o .HHH oHan

40

HuHV .son .um>Hm cmHecH
m x HH<O .ronz .mcmmH<

m N x AMHV .con .mncsouu
MHMh NuHO mmum>wHH

pH H< we as on or on Hm H3. um m2 Hm om an an cOHumHsaom

.mHmmn sumo: ou cuSom n no mocmsvmm GOHudemom .AoocmHum>

mo mHmNHmcmv Hm>oH NH one um Hmcuocm Eoum OwHoHMHO GOHuwHDQom mco coHca

CH mnouomhmco mo unpaid mcu moumoHch Hogans comm .xwo xoan mo mCOHu
:mHDmom OH coozumn mumuomhmco HwOHwOHocmhoE MH CH mmocmuwHMHO HHmHm>O .ucoo HHH chmH

41
south—north basis, to vary in a given direction as a group
even though adjacent pOpulations or particular p0pu1ations
may not follow this order. NO attempt was made to examine
these trends in a more rigorous fashion.

Variation patterns within the three south—north tran—
sects were also studied. The Pentwater population followed
the south~north trends better when compared only with other
western Michigan populations than with all pOpulations. With
respect to other populations the trends were equally clear
whether considered within a single or all three transects.

The Honor II population was the only deliberate attempt
to study a pOpulation having individuals with intermediate-
appearing characters. The Honor I population was sampled
from Open black oak savanna—like vegetation on a dune area
near Lake Michigan. It was observed that the oak stands on
the better developed sandy soils inland from the dunes were a

form of Q, rubravar. borealis and that in the fringe area be-

 

tween the Open dune soil and the better developed soil, there
were many intermediate trees. The Honor I population was
sampled on the poorly developed soil and the Honor II popula—
tion was sampled about % mile inland near the edge of a red
oak stand. Although the Honor II pOpulation sample was not
as large as that of Honor I, there was a significant differ-

ence in variance at the 1% level (F test) in one character

(acorn cup scale shape) and a significant difference in the

112

Fig. ..

(‘nn<11llnns.
basis. Thvre 15 a alxnitxcant dlftcrvnro

the 1 % 1vvvl, analysis Of varlnncv. Kvy to

TOTAL ACORN HEIGHT ( acorn wxlh cup attavhvd )

 

 

 

 

 

mm.
19'
S 18-
17
16
15
1
W‘ FkDu Pu n: ‘1. m H 51 H H- m I; If x 1
ACORN WIDTH ( mm, )
16
15
14
13
12
11
MC Ek Du Pu R1 ‘11; Pt ‘n‘l 81H H': H! Tp T! M 1:
ACORN SIZE ( length plus w1dth )
32 _°
30
28
26
MC EkDu P0 R1515: Pt h'l SI 131 HH Ht Tp T! A] Ir
H1: - Wi‘nrmrk’x (‘nwk 39:1!" Park, 1:11
1:1»: - Inimiiurr, Ind.
{m - {hzruw Shim Dark, Ind.
PM - Pwa\wn S‘nrv PJTH, Ind, ,
H1 - stv Ink» h:'ri.:H- S’atxnn, V: n.
V; - Husk“; n, [1.5.
P! - Prai.n'vr, V; J.
hi — h'll\;E]S1:1$o‘~i‘:1:I-l,WHY}.

Emulation moans of morphological characters of 16 hlnc
Names of pupu1ntlun Sit-rs; are IUIthJvllllt‘tl
:lmnmt 3'

k oak pnpulahons under Held
and arrange-d «m a south in north
,u populatums for path chm-rider at
alphrc'wiallnns llfitcd buhw.

ACORN LENGTH ( mm. )

 

 

 

 

 

H I" P in '~!.; 1’! ‘1“ 5'. H Hm HI Tp Ti Al 1r
ACORN SHAPE ( wxdth length x 100 )
102
35
94
SH)
8'1
8‘3
4“
" E'r; 1' I“- H. \' 1" 51* L H Ht Tp 1'1 3". Ir
'1 E\‘\ 1" ' 1 ' v
_I)
‘1
1'5
7
6
:1 .
4
KC 15k D11 Pu R131: P131 Si H, Ho Ht Tp Tf A1 11'
5'. ‘ .\'.-';,n 1‘ 5117-» Park, Huh.
3' - ’rnx Lucy‘s, My"
1! - H mm ‘41»). .; 1 1
H - H 'w:‘ \f;:n_ ( H i ‘
‘I‘r. W, 1122- Stu!» Park, Huh.
5’1 - 13‘f1'\"‘.7'.n" (‘11:; 1.1:? (yr-1,1113%, Huh.
1‘1 _ \. :1 '43, 14.2!“
- , v , , w v. w.

 

 

 

43

PIK'SBL

conditions. Names of

Populatton means of morphological characters of
population sites are abhr
basis. There IS a signtftcant difference among the populationq

16 black oak
eviated

populattons under field

and arranged on a south to north

for each Character at

t ... ". .
he 1 % 1CVL1, analysts of vartance. Key to abbreviattons Itstod below

 

RATIO OF ACORN CUP HEIGHT ( mm. ) / TOTAL ACORN HEIGHT ( mm. ) x 100

ACORN CUP FRINGE PRESENCE ( 1 - frtnge very promtnent, 1

60

56 8
52 7
48 5
44 3
40 4
36 3
32 2
28 1

 

 

 

MC Ek Du P0 R1 Hg Pt WI SI [1 Hm H1 Tp T! Al Ir

ACORN CUP SCALE SHAPE ( 2 - rounded ttps , 5 - arumtnate , 8 thusv )

 

 

 

 

8
7
6
5 5
4 4
3 3
2
1 1
MC Ek Du Po R1 Mg Pt H1 SI Et Ho Ht Tp Tt A1 Ir
MC - McCormick's Creek State Park, Ind.
Ek - Elkhart, Ind.
Du - Dunes State Park, Ind.
M) - Pokagon State Park, Ind.
R1 - Rosv Lake thdltfe Station, MlCh.

Mg - Muskegon, Mich.
Pt - Pentwater, Mich.
W1 - thson State Park, Mich.

0 - no fringe pr

 

 

VJ

bk In. 5». R; x1... m 1.“. 51m Hm m 'rp Tr A1 Ir

TWIG PUBFSCENCE ( 1 - very heavin pubescent , 10 - glabrous )

_,‘-.

 

Hr

Pk

HUHI Tth A1 Ir

Du P» R1 Hg Pt V1 51 it

1% rt *1 at ht no n1 lp It At Ir

Sleeper State Park, M1rh.

hast Tanas, Mivh.

Honor, 311(1). ( 1 )

Honor, Mlvh. ( II )

Trnvvrce City State Park, Mtch.
Traverse Ctty Falr Grounds, Mlch.
Alpena, “It‘h.

Indtan Rtvvr, Htch.

44

conditions.
basis. There is n
the 1 I level,

significant difference among th

)
analysis of variance. Key to ahbreviatiuns

 

5 - pink , 8 - yello' , ll - orange )

Populntton leans of norpholngicnl chararte
rs n! 16 bln k
Names 0! pm ulation sites are abbreviate pd Endogtran p0puu‘;0ns§ under H N

' vdo uth to north ‘

pu ulatiunsn {or each character at

1istud belo

 

 

 

 

 

 

 

 

11
10
9 TREE FORM ( l — extre-ely fine branching, 10 - coarse open branching
a 8
i 7 7
i 5 6
5 5
4 4
3 3
2 2
l 1
Me Ek Du P0 111 Mg Pt v1 31 21 H0 in Tp H Al Ir 1 MC in W PO to w P‘ n S] in Ho m TD T! M Ir
TUIG mtmmss ( 1o - "lg bends double without snapping, WINTER a“) LENGTH ( M- 1
| 0- tl lig snnps reldily on bend 1m,r )
a 8
i 7 7
. 5 6
5 s
4 4
a 3
2 2
i l l
i .
"c 9" m P" m M“ m '1 31 E‘ H” H‘ T" T' A] 1' we Ek m Po RI M; m a". s‘. n in. H1 Tp Ti Al 1r
MC - McCormick's Creek State Park, Ind. 31 - Sizeper-State-Park,M1ch.
E: : 3::22rétatgdgark,n Ho : Honor, High. I
m - mum" S” M Mich. '11; : fishes-39w Slate Park Mic h.
' R1 : H3259:::? Hiidllfe Station, {{ - §r3\:;se City Fair 8Grounds, Mich.
. — .
:1 : 5:1:zfitgfgt: P:1k,M)Ch. Ir - In Stan River, Mich.

 

 

 

45

Fig. ll A comparison of variance of 16 black oak populations.
The height of the bar represents the number of characters
for each population in which the variance is significantly
greater (F test at 1% level) than that of the population
having the least variance for that character. E. g. The
.Alpena p0pu1ation (Al) had significantly more variance
than the p0pu1ation with the least variance for that
character in 8 out of 13 characters.

 

of characters

1’10.

 

 

 

 

 

J

 

Mc Ek

Mc
Ek
Du
Po
Rl
Pt
W1
31
Et
Ho
Ht
Tp
Tf
Al
Ir

 

Du Po Rl 'Ms Pt W1 81 Et Ho Ht Tp Tf Al Ir

‘McCormick's Creek State Park, Ind.
Elkhart, Ind.

Dunes State Park, Ind.

Pokagon State Park, Ind.

Rose Lake Wildlife Experiment Station, Mich.
Pentwater, Mich.

‘Wilson State Park, Mich.

Sleeper State Park, Mich.

E. Tawas, Mich.

Honor, Mich. (dunes) I

Honor, Mich. (dunes) II

Traverse City State Park, Mich.
Traverse City Fair Grounds, Mich.
Alpena , Mich .

Indian River, MHChm

46
means of 4 of 13 field characters at the 1% level (analysis of
variance), acorn fringe presence, acorn cup scale shape, acorn
shape, and twig brittleness.

In addition to the 13 characters studied in the field, two
additional morphological features were examined later but not
handled statistically; these were leaf outline and winter bud
characters. I

The frequency of leaf types illustrated in Fig. 12 was
counted for 12 p0pu1ations. Type A, in which the leaf lobes were
terminally oriented, occured primarily in the north. Type B in
which the lobes of the leaves were asymetrically arranged and
Type C in which the leaves had a regular pattern resembling a
maltese cross occured more or less throngh the study area. See
Table IV.

The results of the study of dried winter buds collected in
early spring 1963 also indicate variability in the black oak p0p-

u1ations. The main points are:

1. There were two basic colors of pubescence on winter buds,
tawny and gray. These colors occured in both northern
and southern p0pu1ations with gray pubescence being more

common in the p0pu1ations in both areas.

2. Terminal buds varied in size on the same tree depending
on whether the bud developed in the shade or bright sun-
light. Sun buds were often 3 or more times larger than

those growing in the shade.

3. There is a trend in bud size with smaller buds more

47

Fig. 12 Basic patterns of leaf outline for
12 field populations of black oak
from Michigan and nearby states.

 

48

Table IV. The percent of leaves of black oak collected in
each of the three leaf outline types by p0pu1a—
tion arranged on a south to north basis.

POpulation Type A. Type B Type C
Percent

McCormick's Creek State Park, Ind. 0 50 50
Elkhart, Inc. 0 75 25
Dunes State Park, Inc. 0 59 41
Rose Lake Wildlife Experiment 0 36 64
Station, Mich.

Muskegon, Mich. O 79 21
Pentwater, Mich. O 80 20
E. Tawas, Mich. 30 40 30
Honor, Mich. (I) 20 35 45
Honor, Mich. (II) 31 38 31
Traverse City State Park, Mich. 19 50 31
Traverse City Fair Grounds, Mich. 12 53 35

Indian River, Mich. 4 58 38

49
cated by the intermediate nature of some of the
populations between black oak and red oak, for
example Honor I and Honor II. See Fig. 10.

4. The five-angled cross—section of black oak terminal
buds is less conspicuous in the northern p0pu1ations,
but is still evident on close examination.

5. The degree of pubescence on the buds is generally
consistent. In p0pu1ations intermediate in their
morphological characters, there were some exceptions,
in particular, in the Alpena and Honor populations-

It was found in the Honor p0pu1ation that certain
trees with red oak characters in their inner bark
color, acorn characteristics, tree form, etc., had
very pubescent buds resembling black oak though they
had the shape of red oak buds.

On field examination, certain of the black oak p0pu1ations
appeared conspicuously intermediate. These were the Wilson
State Park, Honor I, Honor II, and.Alpena p0pu1ations. The
Wilson p0pu1ation appeared to be intermediate between black
land Hill oak and the other three between black and northern red
oak.

Northern p0pu1ations of black oak were observed for frost
damage through three spring seasons. The differences in damage
observed appeared to be related to time of leafing out rather

than to differences in frost resistance.

50

2. Nursery Studies. The progeny tests in the nursery

 

provided a different type and quality of data from that supplied
by the field studies (Tables V and VI). The nursery tests were
replicated so that it was possible to separate variation into
genetic and environmental components. Therefore it was possible
to attach genetic significance to some of the statistics which
were calculated. The trees were growing under more uniform
environmental conditions than was the case in the field. On

the other hand, die nursery studies were necessarily limited

to two years.

The studies of seedlings grown under nursery conditions
confirm the presence of considerable heterOgeneity among black
oak p0pu1ations, the presence of south-north trends in morphology
and ecology, and the presence of populations having morphologi-
cal characteristics appearing intermediate between species.

As indicated in Tables V and VI, 14 out of 48 characters
scored exhibited significant differences at the 1% and 5%
level (analysis of variance). It is of interest to note that
some characters exhibiting significant differences in the lst
year seedlings were not significantly different in the 2nd
year readings, e.g. leaf length, leaf width, and their product
leaf size.

0f the 48 characters examined, 16 exhibited distinct trends
as defined previously. These characters which are interpreted

as eXpressing south-north trends are as follows:

 

1...!Il. IIIIn‘llIt

 

 

 

 

_ Hm>mH NH mnu um uconMHmem u xx
AmocmHum> mo mHmkacmV Hm>mH Nm mam um quoHMchHm n x
ucmoHMchHm uoz n mz

 

 

 

 

 

 

 

 

 

 

xx x m2 m2 xx mz xx mzfl_xx x xx mucoum

ccwauwm.

o.m m.N o.N o.HH o.N m.H o.mH m.q L.almmmmmmw
. I CmGBumnv mSHm> m
ON om m.m 00mm comm mm we «NH mu OHq am m .Lon .um>Hm cprcH
HH om w.m ONmOH mnoN mm Hq HmH we con mm o .soHZ .zuHo mmuo>mue
«N am ooq OOOHH meN Hm oq mmH 0m ooq 0m N .ron .wocom
on me o.m oomHH Hmoq mm mm «ma am one mm a .eoaz .commxmsz
on me c.m omxmfl whom ma om sea ax omm . N .eoaz .mama mmom
0H Nm N.m. mNqu MHoq NOH mm qu ow OHq mm m .ch .uumanm
"m CH sq o.m ommmH mooq No om qu wk com I N .ecH .umummaoom
.ecH ..xm .um
0H Hq o.m oomNH mem mm mm HqH mm 0mm mm N .Ho m.onEMoooz

mpmhw mmmhmmp mummw umumumEHHHHE mnmumEHHHHE HmnEDc
c N H N H N H N H
mw<. mw< m®< mw< mw< mw< mw< mw<
N0\OH\N H©\m\x Ho\wH\o seasons mmmuu
mocmomonsm oncw cHwHwE cumcoH x supH3 nusz fiuwcmH unmade GOHuomHHoo
mmmH mmoH mmoH mmoH mmmH mowH ungom mo uonfifiz mo huHHwooH
.coHuooHHoo

poem oomH .nuuoc ou :uDom Umwmmuuo mmHuHHooOH wDoHum> CH pwuomHHoo pmom Eoum
.con .wchcmH .m or munch N pom czouw mmcHHpoom xmo xomHn mo whouomuoco cu3ouu .> oHan

52

 

Hm>mH NH um ucmonHmmHm u xx
AmoamHum> mo mHmkacmv Hm>oH Nm um ucmoHMchHm x

H

 

 

 

 

ucmonmacmam 602 n m2
m2 m2 m2 m2 .:m2 m2 mpcmum
ncficuwfi.
m.N w.H m.N m.H H.H . «.0 “w..wpcmum
CmmBumnv QDHm> m
0N mN om mm m o.m .soHZ .nm>Hm cmecH
ma om mH 0N oH a.~ .eunz ..xm.xuao mmuw>mua
m mm mH - 0H «H m.m .aoaz .uocom
w mm ON Hm HN m.m .aoaz .commxmpz
o NH NH mm ma m.m .eoaz .oamq «mom
0 mm ma am on w.~ .ucH .uumstm
m km ma em mH o.~ .ecH .memmaoom
.scH ..xm .um
NH wN OH Hm «N o.m .uo m.onEuoooE
uCoohmm mpwmm mpwuw ucoouom ucoouom opmum
No\o\HH No\m\m No\NH\o No\q\OH Ho\w\HH Ho\oH\o
uDo oHOHumm . MOHoo
wchmoH pop mo MOHoo mmmH
umOH powwow monwmp pow amouw mcHhmm GOHuomHHoo
mm>mmH mo>moH mm>mmH mm>mmH mo>on mo>mmH wo zuHHwooH

 

.COHuomHHoo comm OomH
.nuuoa ou nuDOm umwamuhw mmHuHHmUOH mSOHum> SH UwuomHHoo poem Eoum .non
.wchcmH .m um muwmm N Mom caoum mmcHHpomm xwo xoan mo mumuooumso £u30Hu .ucoo .> oHan

53

 

Hm>mH NH um unmoHHchHm

 

 

 

 

 

u xx
AmocwHum> mo mHm%Hmcmv Hm>mH Nm um unmoHMchHm n x
ucmoHMHcmHm uoz u mz

xx xx xx m2 m2 mpcmum

.83, .

N.m o.mH m.N Hf wmnflnfi
Cmm3uonv msHm> m
mm «q oN m Nm .SUHZ «hm>Hm cmecH
me me NH o , on .non ..xm suHo mmum>mue
Nu mm «N OH om .non .uoCom
mm He HH N an .eon .cowmxmpz
m mN NH o Nm .eon .mme mmom
o H OH q we .ecH .uuweme
.NN o OH OH Hm .ocH .Hmummsoom
.ecH ..xm .um
on O OH HH 5N .Mo mconEHoooz

ucmoumm utmoumm mpmuw ucmouom unmoumm
No\HH\m Ho\m\n No\mMNo HomH HHmm No\W\NH
N m®< H mwm pmwmemp

nmus UCOUMm mocmmwnfim unchm wchHwEoM coHuomHHoo
:uHB mowHH uoosm mm>mmH mm>me mo quHwoOH

 

.wchCMH .m up whom» N Mom GBOHw mwcHHpomm xmo xoan mo muouownmro suzouo

.COHuomHHoo comm oomH
.suhoc ou suSom pmwcmuum moHuHHmooH mSOHHm> CH monomHHoo comm Eoum .soHZ

.ucoo .> mHAwH

54

 

AmocwHHw> mo mHmkacmV

ll

Hw>oH NH um uconMHamHm
Hm>mH Nm um unmoHHHcNHN
osmonHCNHm uoz n Nz

xx

II
x

 

 

   

 

 

 

mz Nz museum
. J. .cHeuHsu
N.H H.H ,. .w ,.
cmmzuoHNmDHm> m
«N 0H .son «pm>HN cmHucH
NN HH .NOHz ..xm quo mmum>mua
NN NH .Non .Hocom
NH 0 .son .cowmxmsz
NN NH .aon .mme mmom
NN NH .NcH .uumeHN
Ne NH .NcH .Hmummeuom
.ecH ..Nm .uN
Hq NH .Ho m.oneHoooz
ucmoumm mwmp HonESC
NNNH HomeHN
Eonm
muoosm whom .mEHu
ou owmfimp moCmNMMEm GOHuooHHoo
“Hrnmm mNcHHemmN Ho NHHHmooH

 

.wchcmH .m up whom» N pom czouw mwcHHpmmm zoo HomHn mo muwuomumso su3ouo

.cOHuooHHoo comm cooH
.cuuoc ou runom pmwcmuum mmHuHHmUOH mDoHHm> CH pmuowHHoo comm Eouw .son

.ucoo .> mHan

55

Hm>mH NH ms“ nu ucmoHNHcNHN

 

 

u xx

Awoquhw> mo mHthocmv Hm>mH Nm onu um unonMchHm n x

unmoHNHcNHm uoz a N2
mz m2 m2 m2 m2, m2 x mz «madam g3
N.N N. N.N N.N N.e N.H . .q.ruuaauaN

cmmauwnv mDHm> h
.LUHZ ampsnouo

 

 

 

ooae «N NN NHH oH NHNN NuHo mmum>mua

ooeN NN .NN NHH N .Non .NNNN mumum comHHz

come oooe NN as «N NN NNH .NN N N . .NoNz .NmugSucmN

OONn on Ca NNH m .non .aowwxmsz

_ NHH H .Non_.NchcmH umNN

, .non .coHoNum

ooNN ooem NN NN NN NN NN NoH N N .NNN mNHHNHHz NNNH wmoN

NNNN NN. NN NN N .Non .mNeHunxoouN

NNHN NNNN NN NN NOH HN A N«H . NNH N N .ucH .NNNN mumum amaze

NNNN NN NN HNH N .NaH .NHNNNHN

Amon coucoEHmv

NNH H .uaH .NNNNNHN
N a. N a. N .< N a. N .<
mnmumEHHHHE NumumEHHHHE honest
monochm . mmmhu

nuwcoH x :uUHB cusz SuwawH unmade cOHuumHHoo

NNmH NmmH NmmH uNNHmN No umaasz No NuHHmooH

 

.pmnoHDE uoc

mumB umnu mcuoom I m awoke .umop3om mo noaH H cuHs pmnoHnE mos uwnu pmnpmmm

w cH poumcHEumw paw poucmHm muoz uwsu mahoom :.< @5090 .coHuomHHoo comm HomH

.mHmon nuuoc ou.:u50m o co pmwcmunw mmHuHHmoOH NDOHHm> aH pouomHHoo comm Scum
.son .waHNcwH .m um now» H Mom cBouw mwcHHpoom xwo xoan mo Nnouoonwso zuzouo .H> mHan

56

Hm>mH NH was um quoHNHcNHN
AmocmHum> mo NHNNHmcmv Ho>mH Nm onu um ucmoHMHame

ll

xx
¥
uGNOHMchHm uoz u mz

 

 

 

 

xx mz xx mz xx xx mz m2 94 m2 mpcmum H .dHfiug
H.N N. O.N N. N.HH N.N N.N H.H N.N dwwamueuuw
CmmBuwnv mDHm> h
.son-«mU:Douu
NH NN NN HN O.N NHNN NuHo mmum>mne
NH «N HN ON . N.N .Non .NNNN mumum coNHHz
N NH NN ON NN HN NN NN N.N O.N .NOHz .Nmumsucmm
NH NN ON NN N.N .NOHz .NOmemsz
OH NN NN N.N .Non .NchcNH NNNN
. .NOHz .coHuNuN
N HH NN NN NN ON NN Ne N.N H.N .NNN NNHHNHHN mNmH mmoN
N NN NN ON AN.N «nqu .NNNHNNxoouN
HH OH ON ON NH NN NN NN O.N N.N .NcH .NNNN mumum magma
N ON NN ON N.N .vaH .NNNNNHN
Amme coucoEHmv
N ON ON N.N .NcH .NNNNNHN
N <_ N a. N a. N a. N ,<
UCGOHmm quOMmm mflmhw wmmwmfi mumuw
NN\N\OH NNHNHOH NN\OH\N NNHNNxm NNHOHHN
MOHoo pop ammuw mocoomnsa meam chHmE GOHuomHHoo
mm>wwA mw>wmfl, wwwd Mwmd mwmd MO %HHHMUOA

 

.UonoHSE uoc oum3 umnu mahoom I m amouo
.umsp3wm mo nocH H suH3 pmsoHDE mm3 umsu ponpmom m CH pmumaHEumw cam
.coHuomHHoo comm HNNH .mHmmn Luuoc

pmudea whoa umsu mahoom t_< @5090
ou sumom o co pmwamuum mmHuHHNUOH mSoHHw> cH monomHHoo comm Eonm .non
.mchamH .m an Hawk H Mom c30uw mwaHHpomm xmo xomHA mo mumuomumno nuBouo .ucoo .H> 0HQMH

Hm>mH NH was um ucNoHNHcNHN

II

xx

AmocwHum> mo NHNNHmcmv Hw>oH Nm osu um ucmoHMchHm u x

 

 

 

 

quoHMchHm uoz n mz
N2 N2 N2 N2 mucmum :aHNuHa
N.N N.H N. A..;Nmauum
cmmaumnv mDHm> m
.soHEIémUcDouw
NN N NHNN NuHo mmpm>wua
HN NH .NOHz .NNNN mumum :ONHHN
NN ON O O .NOHz .Nmumzucmm
NN O .NUHz .commxmnz
.son .wchcwH ummm
.NOHE .cOHumuN
NN NN O N .NNN NNHHNHHN NNNH mmoN
NN. O .NOHz .NNNHNNNUONN
NN NN O O .NcH .NNNN mumum mmqsm
, .HEH JNNNNHN
Amme CoucoEHmv
.NcH .NNNNNHN
7
.3 N .< N .4
mmmmmmm mmmmmwm
NNHNH\N NN\O\N
:NDHM mocommhm
pcooom mHOHumm COHuomHHoo

:uHB mmmMH

pow mo>mmH

No NNHHmooN

 

.poLoHSE uoc mHoB uosu mahooo I m mDoHU

.umdpzwm mo socH H cuHB pmsoHnE mm3 uosu Umnpmmm m CH pouwcHEHow mam

poucmHm mum3 umnu mahoow I.< enouu

.coHuomHHoo pomm HNmH .mHmma suhoc

ou nuDON m to nowadays mmHuHHNUOH NDOHHm> SH pmuomHHoo pmmm eonm .fion
mchcmH .m um Ham» H Mom CBOMw mwcHHpmmm xmo HomHn mo mumuomhmno zuaouu .ucoo .H> mHan

58

1960 planting 1961 planting

height, age 1 height, mulched and unmulched
leaf length, age 1 leaf length, unmulched

leaf length, age 2 leaf width, unmulched

leaf width, age 1 leaf margin, mulched and

leaf width, age 2 unmulched

leaf size, age 1 leaf pubescence, mulched

leaf size, age 2 and unmulched

leaf angle

second flush, age 1
seedling emergence
leaf margin
During both the first and second year there were signifi-
cant differences in height of the seedlings of the 1960 plant-
ing. The tallest the first year being from.McCormick's Creek
State Park, Inc. and the shortest from Traverse City, Mich.
By- the second year, however, the tallest were from Honor,
Mich. and the shortest from Traverse City, Mich. These latter
two p0pu1ations were about 25 miles apart and were both from
near Lake Michigan. In contrast to this the 1961 planting in
the unmulched seedbed (Table VI) gave the appearance of a cline
along Lake Michigan with trees from Dunes State Park, Ind. in
the south being the tallest and those from Pentwater, Mich.
in the north being the shortest. This suggestion of a Cline .
was not so distinct in the interior p0pu1ations.
Trees from Indiana and southern Michigan generally had
larger leaves than did those from Muskegon northward. This
was true whether one considered length, width, or the product

of the two. It is interesting to note that the p0pu1ation

with the least growth among the two year old seedlings (Traverse

59

City, Mich.) rwas also the one with the smallest leaves.

Leaf margin and leaf angle appeared to show south—north
trends in the 1960 planting and among the unmulched seedlings
in the 1961 planting.

In the 1960 planting there were significant differences
among progenies from different localities in percentage of
trees with second growth flushes in July 1961 and June 1962,
amount of leaf pubescence, and amount of twig pubescence. In
all cases the percentages or amounts were higher for the more
northerly prOgenies. These trends were not so distinct in the
1961 planting in relation to second flush but were distinct in
leaf pubescence.

For additional characters including these that were not
significantly different among progenies, see Tables V and VI.

Nursery leaves from both the 1960 and 1961 plantings were
mounted on herbarium sheets by p0pu1ation and studied in terms
of leaf outline. Although there was considerable variation in
leaf outline within and between populations, no trends could be
detected by observation nor did any measurements appear promis-
ing.

Winter buds collected in the fall of 1962 from each of the
two—year old black oak seedlings gave the following results:

1. There was no observable regular difference in bud size

or shape between the following populations; McCormick‘s

Creek State Park, Ind., Rochester, Ind., Elkhart, Ind.,

60
Rose Lake, Mich., and Muskegon, Mich.

2. The following p0pu1ations had buds which were dis—
tinctly smaller on observation than the preceeding
populations; Traverse City State Park, Indian River,
and Honor Dunes, all three of which were from
northern Michigan.

3. The seedlings grown from two of the female parents
from the Honor Dunes population had buds of two types.
The buds of one type resembled those of red oak seed-
lings of comparable age in the nursery, while the
buds of the other type resembled those of black oak
seedlings. The ratio of black oak to red oak appear-
ing buds and twigs in both of these female parents
was approximately 2:1.

4. The Indian River population in the nursery appeared
to have three kinds of buds, thOse resembling black
oak, red oak, and Hill oak. The ratios of black:
red: Hill oaks from the progeny of 6 parent sources
were approximately 10:1:1 for black, red and Hill
oak bud and twig characteristics. 4

In addition to winter bud differences observed in the

nursery as mentioned above, the seedlings from the intermediate
field populations (see previous discussion of intermediate
field p0pu1ations) of Honor and Indian River, Mich. were also

distinctly different from other black oak p0pu1ations in leaf

61

Figs. 13—14 Number of trees in each basic pattern
of leaf outline for mass nursery plant—
ing of black oak seedlings from two

   

origins.

Fig. 13 Dunes State Park, 34 32 34 51
Indiana

Fig. 14 Traverse City Fair 3 45 31 76

Grounds, Mich.

62
pubsecence and twig pubsecence (Table V). This would appear
to confirm the hypothesis that these intermediate field pop-
ulations are also genetically different from typical black
oak.

Two mass plantings of black oak were made in the nursery
in the fall of 1961. 151 seedlings from one female parent
from the Dunes p0pu1ation were grown and 155 seedlings from
one female parent from the Traverse City p0pu1ation were grown.
A study of leaf outline was made of leaves harvested on the
same date from these two female parents. It is evident that
there is considerable variation in leaf outline in the nursery
from the same female parent. See Figs. 13, 14. Other results
from this mass planting from two female parents were:

1. The leaves of the Dunes seedlings were generally

very pubescent on their undersurfaces.

a. 6 seedlings had glabrous or nearly glabrous
leaves.

b. 50 seedlings had leaves with intermediate
pubsecence.

c. 95 seedlings had conspiciously heavily
pubescent leaves.

2. In contrast with the Dunes seedlings, the Traverse
City trees were generally glabrous to somewhat pubescent
on their undersurfaces.

a. 73 seedlings had leaves that were glabrous to
nearly glabrous.

b. 63 had leaves that were intermediate in their

63

pubscence.
c. 19 had leaves that were conspiciously heavily
pubescent.

3. The undersurface of the leaves of the seedlings from

the Dunes p0pu1ation female parent were primarily
brown with a slight red color on their edges. 93
were brown and 58 were more red than brown.

4. Again in contrast, the leaves of the seedlings from
the single female parent from the Traverse City pop-
ulation were nearly all red in color on their under—
surfaces, ten being green and 145 being primarily red.

5. In the Dunes seedlings all of the glabrous leaves and

all but two of the leaves that had intermediate
pubescence were in the red color grouping.

Frost damage was observed in the nursery in both the spring
and fall of 1961 (See Table VII). There was no significadidif-
ference among black oak populations for either spring or fall
frost damage. However, if the Indiana populations of McCormick's
Creek State Park, Rochester, and Elkhart are grouped together
(which totaled only 12 trees due to their later germination)
and compared to the Muskegon population (which totaled 68 trees
due to its earlier germination), there was a difference in frost
damage significant at the 5% level (chi-square), with the more
southern p0pu1ation more prone to frost damage. There is also

an indication that the southern populations are more prone to

64

Table VII. Spring—fall frost damage as observed in the

Bogue
1961.

Population

Lansing, Mich.

Indiana2

Muskegon, Mich.

% of trees showing
frost damage on

May 26, 1961

80

75

38

(8/10) I
(9/12)
(26/68)

Forest Nursery, E. Lansing, Michigan.

% of trees forking
from damaged lead-
shoot

50 (4/8)
100 (9/9)
46 (12/26)

Chi-square test of significance of difference between the
Indiana and the MUskegon populations indicated a signifi-
cance at the 5% level in both % trees frost damaged and %

trees forking after spring frost damage.

In the Indiana p0pu1ation, trees from.MCCormick's Creek
State Park, Rochester, and Elkhart, Indiana, were grouped

together and considered as one p0pu1ation.

65
to forking after spring frost damage than the more northern
p0pu1ations (See Table VII).

The seedlings were scored in the spring of 1962 for the
percentage showing cut stems from rabbit browsing.v There was
no significant difference among the populations of black oak.
An obvious trend was evident with the seedlings from the
northern p0pu1ations appearing to have smaller percentages
damaged than those from the southern p0pu1ations (See Table V).
This was probably due more to size differences than to pala—
tability as the smaller northern seedlings were snow covered
or nearly so during much of the winter.

An unplanned experiment resulted when early snow prevented
the usual sawdust mulching on one of the nursery beds in the
fall of 1961. In Table VI, the mulched seedlings are designated
Group A.and the unmulched Group B. The Dunes State Park and
Rose Lake trees in these two beds were from.the same female
parents though not as many of the parents were represented in
the unmulched bed as in the mulched. The Pentwater trees came
fronlthe same population but not the same female parents. The
characters that appeared to be most effected were % green

leaves in fall, % 2nd flush, and leaf angle.

66
C. Scarlet—Hill Oak Complex
As in the case of black oak, the populations of the
scarlet—Hill oak complex were not homogeneous either in the
field or under nursery conditions. Also in a similar way to
black oak, the heteéganeityu was expressed in south to north
.trends in both the field and nursery data and in terms of a
"break" among the populations in the field data. Some p0pu-
1ations were intermediate between Q. coccinea or Q. ellipsoid-
.glig and other species of the red oak group in the field and
in their characters measured in the nursery.

1. Field Studies. Two transects were established for

 

the study of scarlet—Hill oak populations. One from south-
central Indiana to northern Michigan and the other from
northern Illinois to north-central Wisconsin (See Fig. 1).

The vegetations of which these oak p0pu1ations were
species components are compared using the similarity index,
Curtis (1959). The higher numbers in Table IX reflect greater
similarity in tree vegetation between the stands. The Brown
County, Indiana, and Elkhart, Indiana, stands were both dif-
ferent from each other and from all other stands as seen by
the similarity indexes. From the Thornton WOods stand in
northern Illinois northward, there appears to be a clumping of
vegetationally related stands which is accentuated in the four
most northern stands. In general this agrees with the south-

north grouping which is readily evident in the bar graphs of

67

Table VIII. Site descriptions for stands in which field
p0pu1ations of the scarlet—Hill oak complex
were sampled. Populations arranged on a

south to north basis.

Population Transect

name and size , Location length in
meters

Brown County State NE%, NW%, Sec.l7 700

Park, Ind. (23) R 3 E, T 8 N

Brown County, Ind.

a/Elkhart, Ind. 8%, W%, Sec. 9 250
(18) R 5 E, T 38 N
Elkhart County, Ind.

Thornton woods, E%, wa, SE% 400
Chicago Forest Sec. 34

Preserves, Ill. R 14 E, T 36 N

(20) Cook County, Ill.

Rockford, 111. NE%, SE%, Sec. 15 250
(19) R 2 E, T 43 N
Winnebago County, Ill.

a/Stockbridge, Mich. 8%, NE%, Sec. 28 1600
(11) R 3 E, T 1 N
Livingston County, Mich.

a/Rose Lake Wildlife N%, SE%, Sec. 24 975
Experiment Station, R l W, T 5 N

Mich. (18) Clinton County, Mich.
a/Wilson State Park, W%, NWk, Sec. 21 175
Mich. (14) R 4 w, T 19 N

Clare County, Mich.

a/Farwell, Mich. NW%, NW%, Sec. 36 225
(26) R 7 w, T 19 N
Osceola County, Mich.

a/Wisconsin Rapids, N%, NW%, Sec. 16 1275
'Wisconsin (20) R 6 E, T 21 N

Wood County, Wisc.
a/ Grayling, Mich. swx, NEx, Sec. 19 200
(29) R a w, T 26 N

Crawford County, Mich.

a/ Represented in nursery test also.

Soil
type

Muskingum
Stony Silt
Loam

Ottawa Loamy
Fine Sand

Ottawa Loamy
Fine Sand
with moisture
catena

Alvin Sandy
Loam

Boyer Sandy
Loam

Brady Loam

Grayling
(Loamy)
Sand

Omega Loamy
Sand
Grayling
Sand

Rubicon Sand

(toward Kalkaska)

68

I .NN .N ANNNHN mHouso
=.oH5NmoE comm How NoDHm> Hm30H osu mo HHN mo ESN msu NH 3 ram NuHucm uwnuo msu
How NoHSNmmE HHw mo ESN map NH m_.%uHuCo oco How NmHSNNmE HHm mo EDN mfiu NH m_mum£3=IIH

x HNNN .Non .NcHHNNNO
mq. x AHBV .oNHB .NpHmmm CHNCOUNNB
NN. Nm. x Aahv .soHZ .HHNBHmm
NN. NN. NN. x Hsz .Non .NNNN ooooN coNHHz

HHNV .Non .coHomuN

NO. OH. NH. NN. N .NNN oNHHoHH3.onH oNoN
NH. NH. ON. NN. NN. x HNNV .Non .mNoHNNNoouN
NH. NH. NH. NN. HN. HN. N HNNN .HHH .oNoNNooN
AnHv .HHH .No>umNon uwouom

NH. NH. NH. NN. NH. NH. HN. N onoHNO .moooz coocuoca
HH. NO. NO. HH. NO. OH. NO. NH. N HHNV .NcH .oNNNNHN
HNNV .ocH

NH. NH. NH. NH. NH. NH. NH. NH. NO. .NNNN ooNoN Nocooo caoNN

NO N: 3N Hz. HN NN NN NH HN

.NHNmn cuuoc ou sumoN N Go Uqumnum NcOHumHSmom .NucHom NHQENN OH nqu

.ANmmHv awSmnchonu .Uosuoe :oHHHouuHm wsu wcHND pochuno NoumEHuNo Nona HmNmn

Eoum pmquEoo whoa NmoHucH one .onmEoo xwo HHHqumewoN mcu wchHmucoo Npcmum
0H CH GOHuNumwo> mmuu How .HAommHv NHuHSU .n x w \ 3N n m .NuHHmHHEHm mo Nmch .NH oHan

69

N HNOV .Non .NcHHNoNO

o x NHBV .oNHB .NpHdmm cHNcooNHB
OH N N HNNN .Non .HHNNNNN
HH HH N N AHNN .Non .NNNN ooNoN comHHz
AHNV .Non .aoHumuN

N N N N N .NNN oNHHoHHz oNNH oNoN
N N N N N N HNNN .Non .NNoHNNNoooN
N N N N N . N N HNNN .HHH .oNoNNooN
ArHv .HHH .Nm>HmNmum

m N m N N m m N uNmHom ommoHfio .Npooz couCHonH
N O H H N N N N N HHNV .NcH .oNNNNHN
ANNV .ocH

N H H N m m N N N .xnmm oumum Nudfloo ozoum
NO N2 3N H: HN NN NN NH HN coHoNHsNoN

.NHNNQ cuuoa ou :uDoN N Go pqumuh< .AoocmHHN> mo NHNNHNGNV
Ho>oH NH msu um NHucmonHcmHN Hmmme no: uHu NCOHuNHDdoQ xwo HHHquoHHNoN mo
NHHmm UmuNmHucoo noHsa CH Nuwuomumso HNUHNOHOLQHoE UHmHm mo Hmnasc mfiu mo 83m .x NHQNH

70

D
3 " . .
F.,. .5. Ff’tptlicljat‘xon means or" m0rph010K11~31 charact-‘rs n! 10 “(Vivi—Hi“ ‘
It Londltions. Names of pupulatlnn sztus Irv ahnhwhi.. , “‘ 03k PquIatxuns undpr
north hafils‘ Thar" 1‘ 3 Slunxfxvanl dI¢INrNNpp 1L7} tt;d|"” and RFYRUNVQ mm a SHU{
’ '. .. l ‘ V H ‘ H|c1 t‘I" 1‘ U1. i“ O . .‘ _ ‘
It the 1" letcl, “11113515 ”I vanancc‘ Km ta :mh'w.”IAITIIth";I:dL2 Alt” each (hm-miter
_ t v . N A‘-\' _._f U.-

TOTAL ACORN “EIGHT ( “CUT“ ‘11“ CUP attachud ) mm, Afohx LINGTH t7m.)

22'
21-
19
201 .
AH
lgq ._
If 18‘ .
E V
17.
164 . .

15:

 

 

 

 

Br ‘ I u . ., . -
MJX SK R1 n] Ft Br (I. ..; .’. .3; .5g 1.“ 'n. 'r‘- a. m

h

ACORN WIDTH (T7,)

Vt)?“ FHH‘? .;.:'; 1‘ x ,H"

14 .
El(|

mm.

 

 

 

Br £1 Tn RR 5k R; h.

A! URN [71'1“ K? I’l'I'I'T "‘ .I

K)

ACORN SIZE ( lonuth plus width )
LU

34
32

'7"! _

3O
28
26

 

 

 

 

 

8? El Th RK 5k H1 *1 F1 I? 3; Br Ll Th Rk Sk R1 kl Ix hr 0;
R1 - Rose Lake Exldlife Station, chh.
I] - leson State Park, MICh.
[11. PT - Farwell, filth.
Kr - Wisconsln Rapids, its.
- Grayling, Mich.

Br — Brown County State Park, Ind.
E1 - Elkhart, Ind.
Th - Thornton Hoods, Chicago,
Rk - Rockford, 111.
SR - Stockbrldhe. MIch. Cy

 

71

F. .‘ Population means of morphological characters of 10 scarlet-Hill oak populations under
fi" ' field conditions. Names of population Sites are abbreviated and arranged on a south to
north basis. There is a significant difference among the populations tor each character
at the 1% level, analysis of variance. Key to abbrgyiglions listed below-

 

ACORN CUP FRINGE PRESENCE ( 1 - fringe very prominent,
10 - no fringe present )

RATIO or ACORN CUP HEIGHT ( mm. ) / TOTAL ACORN HEIGHT ( mm. ) x 100 10

 

 

 

 

 

 

9
56 8
52 7
48 6
44 5
4O 4
36 3
32 2
28 1

Br E1 Th Rk Sk R1 W1 Fw Wr Cy Br El Th Rk Sk R1 W1 Fw Wr Gy

TWIG PUBESCENCE ( l - very heavily pubescent,
10 - glabrous )

 

 

 

 

 

 

10
- 9
ACORN CUF’SCALE SHAPE ( 2 - rounded tips, 5—acum1nate,
8 - obtuse 8
7
7
6
6
5
5
4
4
3
3
2
2
1
1
' Br El Rk Sk R1 wi Fw Wr Gy
Rk Sk 31 W1 Fr Wr Gy . '
Br El Th t Park Ind. R1 — Rose Lake Wildlife Station, Mich.
Br - 812;“ Couggg Sta 9 ' W1 — Wilson State Park, Mich.
E1 - E ar , .

- , Fw - Farwell, Mich. P
Th ‘ Thozptog W???S, ChicagO, 111' Wr - Wisconsin Rapids. “15.
Rk - Roc or I .

' - ' ‘ Mich.
Sk - Stockbridge, Mich. Gy Grayling,

72

:|‘.|7. Population means of morphological characters of 10 scarlet-Hill oak populations under
field conditions. Names of population sites are abbreviated and arranged on a south to
north baSIS. There is a significant difference among the populations for each character
at the 1% level, analysis of variance. Key to abbreviations listed below.

INNER BARK COLOR ( 2 - tan, 5 - pink
8 - yellow , 11 - orange )

NCO-5005

TWIG BRITTLENESS ( l - twig
10

IbU‘CDKI

OJ

 

 

 

 

Br El Th Rk Sk R1 V1 Fw Hr Gy

 

Br El Rk Sk R1 W1 PW Kr G;

Br
El
Th

Sk

— Brown County State Park, Ind.
- Elkhart, Ind.

— 'ThIIrIItrin MAIININ, (III: aIgI, I ll.
- Rocktnrd, Ill.

- Stockbridge, Mich.

bonds double without snapping,
— twig snaps readily on bending )

'Tllfl .

RI
V1

Fw
Br

(I)

TREE FORM ( 1 — extremely fine branching,
10 - coarse open branching )

 

 

 

Br El Th Rk Sk R1 W1 Fw hr Gy

WINTER BED LINGTH ( mm. )

 

 

In- El Th RR SR R1 III H way

— RHSI'IJIKV hi hilifv StatIInI, Mich.
— kilsun State Park, Mich.

- Faravll, With.

— hiscnnhin Rapids, his.

- Oraxiixg, Vl~a.

73

the field morphological characters, Figs. 15 through 17,
and also in the table of homogeneity, Table X.

The populations were far from homogeneous. In each of
13 morphological characters studied in the field in the scarlet-
Hill oak complex, there was a difference among the populations
significant at the 1% level (analysis of variance).

However, this heterogeneity was not at random. In 9 out
of 13 characters there was a south—north trend as defined on
p.34 , in the population means. This was true on both transects.
The characters that showed trends were; acorn with cup attached
(total acorn height), acorn length, acorn width, acorn size,
acorn shape, acorn cup height, ratio of acorn cup height to
total acorn height, acorn cup scale shape, and twig pubescence,
Figs. 15, l6, 17. It should be noted that 8 out of 9 of these
character trends represent acorn characters.

As reported earlier for black oak, a sharp break in acorn
dimensions occurred between scarlet-Hill oak populations on
different sides of the transition zone. Note specifically the
Rose Lake and Wilson State Park p0pu1ations. (Fig. 15)

Leaves from each tree in the field p0pu1ations were studied
from specimens mounted on herbarium sheets. It was possible
to place the leaf outlines of the trees belonging to the scarlet—

Hill complex into four major categories. Fig. 19. In leaves

of Type A the pattern was regular with lobes balanced in size

and number on either side of the midrib. In Type B the leaf

74
lobes were unbalanced in either their shape or position on
either side of the midrib. In Type C the leaves resembled
those of red oak with rather small and uniform lobes, while
in Type D they resembled those of pin oak in the depth and
type of lobes. See Fig. 19. Table XI summarizes the occurence
of these types by p0pu1ation.

Though only small differences in the pattern of leaf out-
line could be detected, there was a tendency for the leaves of
the northern populations to be smaller than those of the southern
p0pu1ations.

Winter buds were also collected from each of the p0pu1a-
tions in the early spring of 1963. The dried specimens were
later measured and compared in the laboratory. There was a
definite trend from south to north in bud length, the dimension
measured. (Fig. 173),The two southern p0pu1ations were Brown
County, Ind. and Elkhart, Ind. had buds that were substantially
larger than the other populations. The presence or absence of
pubescence on the buds and bud shape were also scored. (See
Table XII).

In general the more northern populations had a higher
prOportion of glabrous buds and also a higher proportion of
buds with pointed apices.

The more pubescent winter buds approached those of black
oak in both the southern and northern p0pu1ations even to the
expression of angularity so characteristic of black oak. Their

size relationship was also similar in that the northern popula-

75

Fig. 18 A comparison of variance of 10 scarlet-Hill oak popu-
lations. The height of the bar represents the number "
of characters for each p0pu1ation in which the variance
is significantly greater (F test at 1% level) than that
of the p0pu1ation having the least variance for that
character. E.G. The Rockford population (Rk) had sig-
nificantly more variance than the p0pu1ation with the
least yariance for that character in 6 out of 13 characters.

 

 

 

wwV—wv-‘u—r.’ 7.,— “w .—-v-—-—

Br El Th Rk Sk Rl 'Wl Fw ‘Wr Gy

no. of characters

Sampled populations ordered south to north basis

Br — Brown County State Park, Ind.

El — Elkhart, Indiana

Th - Thornton Whods, Chicago Forest Preserves, Ill.
Rk - Rockford, Ill.

Sk - Stockbridge, Mich.

R1 - Rose Lake Wildlife Experiment Station, Mich.
W1 - Wilson State Park, Mich.

wr - Wisconsin Rapids, Wisc.

Gy — Grayling, Mich.

76

Fig. 19 Basic patterns of leaf outline for 9
field populations of the scarlet-Hill
oak complex from Michigan and nearby
states.

 

77

Table XI. The percent of leaves of the scarlet-Hill
oak complex collected in each of four leaf
outline types by population arranged on a
south to north basis.

Population Type A. Type B Type C Type D
Brown County State Park, 18 82 0 O
Ind.

Elkhart, Ind. 62 38 0 O
Thornton WOods, Chicago, 56 44 0 6
Ill.

Rockford, Ill. 50 50 O 6
Stockbridge, Mich. 45 52 3 0
Rose Lake Wildlife Ex. 76 24 0 0
Station, Mich.

Farwell, Mich. 59 41 0 0
Wisconsin Rapids, Wisc. 46 54 O 0

Grayling, Mich. 56 3S 9 O

78

tions of the scarlet—Hill oak complex had large buds that were
comparable to the large buds of the more southern black oak and
the northern scarlet-Hill oak populations had small buds that
were comparable to the small buds of the northern populations
of black oak. The glabrous buds approached those of the shade
buds of northern red oak (Q, gubgg var. borealis). They were
also very similar to pin oak buds in size. In the Alpena popu-
lation in which many trees were found that appeared intermediate
between black oak and northern red oak, pubescent buds were
found that closely resembled the pubescent type of Hill oak bud.
The presence of a ring or of several rings of fine pits
around the apex of mature acorns of scarlet oak has been used by
Deam (1953) as a diagnostic character for that species. The
scarlet-Hill oak populations examined in the present study were
scored for pitting with the following results; Brown County,
Indiana, p0pu1ation, 13 trees with pitting, 9 without; Elkhart,
Indiana, 6 with, 15 without. Pitting was absent in the samples
in all other p0pu1ations in this study. However, I have found
individual trees of this scarlet-Hill oak complex in the Rose
Lake, Michigan, area which occupied better drained sites than
the population sampled for this study, which did show distinct
pitting on their acorns. Also out of the 9 female parents

represented in the nursery plantings from Wisconsin Rapids,
Wisconsin, one parent tree had acorns which showed distinct

pitting.

79

Table XII. .A summary of the number of trees in each
population of the scarlet-Hill oak complex
with winter buds in each of two main cate-
gories, bud pubescence and bud shape.

Population Bud Pubescence Bud Apex
Pubescent Glabrous Pointed Round
number of trees number of trees
Brown County, Ind. 11 O 6 5
Elkhart, Ind. 3 0 l l
Thornton Woods, Ill. 3 4 6 0
Rockford,Ill. 5 0 3 2
Stockbridge, Mich. 8 0 7 3
Rose Lake Ex.Sta., Mich. 8 3 10 1
Wilson State Park, Mich. 9 3 ll 1
Farwell, Mich. 5 3 ll 0
Wisconsin Rapids, Wisc. 5 5 10 0

Grayling, Mich. ll 7 15 3

80

The shape of the acorn cup was not particularly dis-
tinctive in Michigan. Nearly all were hemispheric in shape
with only occasional trees showing top-shaped cups. In the
Wisconsin Rapids population nearly one-half of the acorn cups
were top-shaped.

Trees that appeared intermediate between black oak and the
scarlet-Hill oak complex were found in the following stands:
Stockbridge, Mich., Thornton Woods, Ill., Rockford, Ill., and
Wilson State Park, Mich. In the Thornton WOods stand there were
a few trees that appeared intermediate between the scarlet-Hill
oak complex and pin oak. Intermediate appearing trees between
the scarlet-Hill oak complex and red oak (Q. Egbgg var. borealis)
were found in the Cass Lake stand in Minnesota. This is dis-
cussed in detail later under red oak. In the Alpena stand in-
dividuals were found that appeared intermediate between all three
species, black oak, red oak, and the scarlet-Hill oak complex.

Field observations were made of spring frost damage on 3
consecutive years. No differences were noted that did not appear
related to time of leafing out.

2. Nursery Studies. The 1960 planting included 3 p0pu1a-

 

tions of the scarlet-Hill oak complex, from Farwell, Mich., Gray—
ling, Mich. and Wisconsin Rapids, Wisconsin. According to the
field studies, the Wisconsin Rapids, Wisconsin, and Grayling,

Michigan, populations are similar in 9 out of 13 morph010gical

characters. (See Table X).

81

Comparison of the nursery data for these two populations
also indicates a great similarity between them. In 26 or 27
of 29 characters (Table XIII), The Farwell, Mich. p0pu1ation,
represented by a single progeny appeared to differ from the
other two in growth rate and waviness of the leaf margin.

In 1961 an attempt was made to obtain acorns from a larger
number of scarlet—Hill oak p0pu1ations. There was evidence of
differences among the nursery populations. South to north
trends were more evident than in the black oak test. In those
as compared with 4 out of 13 nursery character northern trees,
the southern ones were faster growing, had later autumn color-
ation, had larger leaves with wavier leaf margins and had leaves
attached on more nearly a horizonal plane.

Unlike the field data, the nursery data, based on the 1961
planting, do not appear to show any distinct genetic evidence of
two major population groupings (a northern and southern) but in—
dicate a gradual genetic trend from south to north.

Collections of leaves were made from both the 1960 and 1961
scarlet-Hill oak nursery populations. A sample leaf was collect-
ed from seedlings of each individual tree. The p0pu1ation was
studied in terms of leaf outline. No basic differences in leaf
outline could be detected.

Collections of winter buds were made from the 1960 nursery
planting in the winter of 1962. No differences could be detected.

between the three Hill oak populations, Farwell, Mich., Grayling,

82

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87

Mich. and Wisconsin Rapids, Wisc. Also no distinctive separation
was observed betweed the buds of the Hill oak populations and
those of the Traverse City State Park and Indian River black
oak populations. Thelrraverse City population, in particular,
and to a lesser degree the Indian River p0pu1ation had buds re-
sembling those of black oak. There appeared to be no discontin-
uityin size and character of winter bud between these two black)
oak populations and the 3 northern scarlet-Hill oak populations.
In the Honor Dunes population the buds were either similar to
black oak or distinctly intermediate between black oak and red
oak. (See p.60 ) for a more complete discussion of this point.

The exact nature and limits of the taxon Qgercus ellipsoidalis

 

 

and its distinctness from.g. coccinea have been a problem for some
time. The present study began with the assumption that the
northern small—fruited black oaks were really a part of the
scarlet—Hill oak complex. The first year seedlings that develOped
from acorns planted in 1960 before a distinction was made between
these two entities tended to support their relatedness. However,
leaf margin, leaf angle, and height growth did suggest that two
entities were involved. (See Table XIII). The two year old
plants scored the second year from the same plantings indicated
even more pronounced differences. The basic relationship of the
Traverse City State Park, Honor Dunes, and Indian River p0pu1a-
tions to black oak became evident in many character measurements,

for example, leaf size, leaf length, leaf shape, leaf pubescence,

88
presence of red petioles, height growth and shoot pubescence.
The individual character of these three p0pu1ations varied
from the individual character-means-approximaEely those of
black oak to intermediate values between black oak and the
scarlet-Hill oak complex. An intermediate value between these
two species groupings need not necessarily mean that the p0pu—
1ation was genetically intermediate between black and the scar-
let—Hill oak complex as the same intermediate means might also
arise in a population genetically intermediate between black
oak and red oak. See Table XV for a summary of these relation-
ships by character.

Two mass plantings of the scarlet-Hill oak complex were
made in the fall of 1961. 151 seedlings from one female parent
in the Grayling, Mich. population and 132 seedlings from one
female parent in the Elkhart, Indiana, population were grown.

A study of the variation in leaf outline was made from a selec-
tion of a typical mature leaf from each of these seedlings with
the results as shown in Figs. 20, 21.

In the progeny of the Grayling parent a leaf of one in-
dividual was slightly pubescent on its undersurface while none
of the leaves selected from the others were. The leaf colors
were distinctly different between the Grayling and the Elkhart
seedlings. Three leaves were red and 148 green in the Grayling
while in the Elkhart seedlings 105 leaves were red in color and

27 were green. The leaves were collected on the same day in

October.

89

Table XV. A summary of the relationships of three inter-
mediate p0pu1ations growing under nursery con—
ditions.

B - similar to black oak.
H — similar to Hill oak.

' Indian River, Honor, Traverse City
Mich. Mich.(Dunes) St. Pk., Mich.
Leaf size B B H
(61 scoring)
Leaf size H B B
(62 scoring)
Leaf Width B H H
(61 scoring)
Leaf width H B B
(62 scoring)
Leaf shape B B B
(62 scoring)
Leaf margin B H B
Leaf pubescence B B B
Degree red H B B
Petiole
Z Greaileaves H B B
fall
% Red leaves H B B
fall
Height growth B B H
Z Leafing out B B B
spring
Shoot pubescence B B B
Leaf angle B B B
1 Leaves remaining H H B
Leaf length B B B
(62 scoring)
Totals1 Black - 10 black - 13 black - 13
Hill - 6 Hill - 3 Hill — 3

1 In 11 of 27 characters it was not possible to make a black

oak or Hill oak designation due to their similarity.

90

Figs. 20-21 Number of trees in each basic pattern of
leaf outline for mass nursery planting of
scarlet—Hill oak seedlings from two origins.

 

Fig. 20 Elkhart, Indiana ' 17 13 41 61

 

91

It is useful to note that in the 1960 plantings in which
northern p0pu1ations of black oak and Hill oak were grouped
for study purposes, the two taxa were not significantly dif-
ferent from each other at the 5% level (analysis of variance)
in percent of rabbit damage, spring frost damage, and percent
forking after frost damage. There was a significant difference
among the populations at the 5% level in percent fall frost
damage. On the whole, this suggests that in certain basic

reactions to the environment the two taxa were not different.

92
D. Red Oak

There was considerable variability in both the field and
nursery p0pu1ations of red oak. There was evidence of south-
north trends in both the field and in the nursery, but the trends
were not as pronounced as in black oak and the scarlet-Hill oak
complex. Intermediate appearing trees between red and black oak
were evident in several populations and also intermediate appear-
ing trees were found between red oak and the scarlet-Hill oak
complex. A mass planting of one intermediate red oak was made
with observations on possible segregation of the seedlings.

1. Field Studies. Two transects were established for the
study of red oak populations, one along the west coast of Michi-
gan from Warren Dunes State Park to Benzie State Park and one in
the interior of the State from Pokagen State Park in northern
Indiana to Douglas Lake in Michigan. See Fig. 1. Three popula-
tions were studied in the Upper Peninsula of Michigan and one in
Minnesota.

The vegetations of which the oak populations were species
components are compared using the index of similarity, Curtis
(1959) in Table XVII. No pattern was detected in the relative
similarity between stand composition.

Red oak in the study region has long been considered as com-
posed of two races. Thus it comes as no surprise to find that

when all the data on red oak are pooled, each of the 13 characters

show differences among the red oak population which are significant

93
at the 1% level (analysis of variance). The chart on a
comparison of variance (Fig. 26) also presents an interesting
picture of heterogeneity. The p0pu1ations with a score of
two or below whether from the north or not were g. Egbgg and
those with a score above 2 were Q, Egbgg var. borealis.

The variation in field character measurements, as in the
case of the two preceding species, suggests a south to north
trend among the populations. This trend (as defined on p. 39)
appeared in 7 out of 13 field morphological characters, Figs.
22, 23, 24, 25. All 7 of these are acorn characters; acorn
with cup attached (total acorn height), acorn length, acorn
width, acorn size, acorn shape, acorn cup height, and ratio
of acorn cup height to total acorn height. Certain northern
p0pu1ations (Burt Lake State Park, and Dunbar) depart consid-
erably from this average trend. Considerable differences be-
tween southern and certain northern populations appear on the
graphs but due to the variation among the populations, no
definite breaks or break suggest themselves.

Leaves from seedlings of each individual tree sampled
in the population were examined for differences in the pattern
of leaf outline. No basic difference could be detected in
the pattern of leaf outline between the following populations;
Pokagon State Park, Ind., Warren Dunes State Park, Mich.,
Muskegon, Mich., Benzie State Park, Mich., Burt Lake State

Park, Mich., Douglas Lake, Mich., and Dunbar, Race and Copper

94

Table XVI. Site descriptions for stands in which field
populations of red oak were sampled. Popu-
lations are arranged on a south to north

basis.

a/ indicates populations planted in the

nursery

Population name Location Transect Soil

and size length in type
meters

Pokagon State Pk.,
Ind. (18)

Warren Dunes State
Pk., Mich. (21)

East Lansing,

Muskegon State Pk.,
Mich. (20)

a/Benzie State
Park, Mich. (27)

a/Honor, Mich.
(dunes)(9)
Alpena, Mich.
(21)

a/Burt Lake St. Pk.
Mich. (23)

NW%, SE%, Sec. 34 240
R 13 E, T 38 N

Steuben County,

Ind.

SW%, NW%, Sec. 25 500
R 20 W, T 6 S

Berrien County,

Mich.

E%, swa, Sec. 19 350
R 1 w, T 4 N

Ingham County,
Mich.

we, swa, Sec. 17 250
R 17 W, T 10 N

Muskegon County,

Mich.

NW%, NW%, Sec. 27 475
R 15 w, T 27 N
Benzie County, Mich.

NW%, NW%, Sec. 23 1175
R 15 W, T 27 N

Benzie County, Mich.

Wé, SE%, Sec. 27 425
R 7 E, T 31 N

Alpena County, Mich.

NW%, Nwa, Sec. 25 175
R 3 w, T 35 N

Cheboygan County,

Mich.

Boyer Sandy
Loam

Bridgeman
Fine Sand

Hillsdale
Fine Sandy
Loam

Deer Park
Fine Sand

Rubicon Sand

Deer Park
Fine Sand

Rubicon Sand

Rubicon Sand

A

Table XVI, cont.

Population name
and size

a/Douglas Lake
(Univ. of Mich.

Bio. Sta.), Mich.

(26)

a/Dunbar Forest
Station, Mich.
(25)

Raco, Mich. (18)

Copper Harbor,
Mich. (20)

Cass Lake, Minn.
(20)

95

Site descriptions for stands in which
field populations of red oak were
sampled. Papulations are arranged on

a nouth to north basis.

a/ indicates p0pu1ations planted in

the nursery

Location

NW%, SE%, Sec. 33
R 3 W, T 37 N
Cheboygan County,
Mich.

wa, ssa, Sec. 6
R 2 E, T 45 N
Chippewa County,
Mich.

we, NE%, Sec. 13
R 4 W,-T 46 N
Chippewa County,
Mich.

N%, NW%, Sec. 33
R 28 w, T 58 N

Keweenaw County,
Mich.

NW%, Sec. 19
R 30 W, T 145N
Cass County, Minn.

Transect
length in
meters

300

250

800

575

3200

Soil
type

Rubicon Sand

Rousseau
Loamy Fine

'Sand

Rubicon Sand

Waiska
Cobbley
Loam

Cass Lake
Fine Sand

96

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97

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$ ' a Population loans of morphological characters of 13 red oak populations under field

a' ’ Conditions. tunes of population sites are abbreviated and arranged on a south to north
basis. There in a signiiicant difference among the pOpulattnns (or each character at
the 11 level, analysis of variance. Key to abbreviations listed below.

 

TOTAL ACCRN HEIGHT ( acorn with cup attached ) u. ACORN LENGTH ( u. )
39 23
35. 22
2‘4 21
’34 211
22" - 19

3

2H 11;

3 20‘ 17
1% 11f.
111.. 1.3
17. 14
16‘ 13
is.

 

 

 

 

 

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Ph I'dEn apnp H0 11pm n1 Dnlla ChCi

ACGN WIDTH ( II.)

21
20
19
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is
51' 94
16 «m
15 no
14 192
13 m

 

 

 

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0
9
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8
7 7
6 fi
5 5
4 .1 i
l‘
3 3
2 2
l 1
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id - \iarrun Dunvs State Park, Mich. i)‘. - Douglas iriknr, Myth.
lin - i-Zast 121115.111... \iirh. iin — Dunbar Pun-st Station, Mich.
Hp — Wish-tam Stat" Park, Mich. R21 - R‘( 11, H1111,
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Ap - Alpena, ‘iivh.

7 , . —v— v.

99

. Population leans of Iorphological characters of 13 red oak populations under field
Fig. 13. conditions. Names of population sites are abbreviated and arranged on a south to north
basis. There is a significant difference among the p0pu1ations for each character at
the 1‘]. level, analyaia of variance. Key to abbreviations listed below.

 

11m SIZE ( length plus Iidth )

 

 

 

 

 

 

 

 

 

46
M
42
40
38
36 ACORN CUP HEIGHT ( ma. )
34 9
32 8
30 7
28 6
26 g 5
4
Po id En Mp 3p 50 AP 31 01 Du Ra Ch C1 P11 in! 1111 \ip Bp 110 Ap 81 01 Dn Ra Ch Cl
ittiRV ('i'P l‘RlNGi. PRi'SENt‘ivi ( 1 - fringe very prominent,
it) — no iringt- present )
;11
u
1“
RATIO OF ACORN ('1'? ii} lflif’f ( 77.5 11'71! \1 ii-;\ i"- l‘viTI i ' 1 \ ,1111 7
48 n
44 7
40 i
36 It
32 g
28
P" W 11‘- ‘“ 3’“ "1 '1? "- 1' ‘1 "- 11 .11 1h “:9 hp 11 1;» 111 m 1m Ra Ch C1
1’1 A 1‘ :1 1 1.“ 1 1 ‘1 "1'1... “7‘ 1‘ 1’ it 11.1%.; got-gtl‘d’rk, V1111.
:., :1 I ,‘ ‘1‘ LR- “ M 1 :r Pr. ' 51.121111, V1111.
up - ‘1‘ r ‘1- . ‘ ‘1 R1 1 -.
‘7“, D‘r1 1 1 , “‘

100

a I I UlaI i’fin Ill 1 of mo h(l P
F P( I) \ PL “R I I) (If (a “'1 ‘31 rs ’ I [‘9 p
. ‘. (1.111111 I IUHS . Ndlif'S “I. DUDU 1 III I (1)17]. qlli‘t‘;' '11:: h 3;” O l I dd ”3k -"1’Ul.~i'i(ms UndPl field
,3 i » - N 4 ~-' 1 )I (‘V :1 0 and il I‘I‘arli’fv’d ”fl a “Hill I h to I
,. - . nU th

b’ifls I’M‘IP 15 a El " ‘ ‘Q ’I H P I lg H h ' ‘0 at
a N . . k . L,i|1ll( ant dl‘l 'I‘1‘iH‘l" 1 ‘I ' ( a a r
. ‘ ‘A Q 'A th" 0 ,1.) v ‘ l![‘ 1",‘ Q ‘ ‘
e a - , , . ' ‘ M (a 7 7
th 1% ILLLI' analybls 1)! Vdrlank‘c. kt’y IU nt’IlIeVIaIIUIIS llSlL’d btv'l'l“ l I (

 

 

Th 16 R‘IIFSCI'NFF ( 1 — vol-3'. hwavl 1) pu hCSt‘t‘nt I 10 - izlahr()us )

 

 

 

 

 

 

10

ACORN CUP SCALE SHAPE ( 2 - rounded tips, 5 - acuminate, g
6 - nihilist‘ }
8 B
7 7
5 h
5 5
4 4
3 1
2 2
l 1
P0 b‘d En Mp Bp Ho Ap Bl Di Dn Ra (11131 P11 11d E11 ‘vip 8;) HM Ap 111 111 [in Ra Ch Cl

Tkhf EORV ( l - crirvnv}, lint Linn1niug,

 

 

 

 

 

lU -1-u:sv1qnqi:wnn1uhu )
9
INNER BARK COLOR ( 2 - tan, S-pink, B-yvllnw, ll - orange )
H
7 7
6 (i
5 5
4 4
3 3
2 2
1 1
Pg Wd En Mp Bp Ho Ap Bl Di Dn Ra Ch C1 P1 hd En Hp Bp Ho Ap Bl D1 Du Ra Ch Cl—

81 — Burt Lake State Park, Mich.
D1 - D1uglas Lake, Mich.
Dn - Dunbar Forest Station, Mich.

Po - Pokagon State Park, ind. .
Wd - Warren Hines State Park, Mich.
En - East Lansing, With.

Mp - Huskvgun State Park, Mich. Rn — Raco, Hith. ,
Bp - Bcnziv State Park, Mich. Ch - Cuppvr Harbor, Mich.
- Cass Lake, Minnesota

H0 - Honor, Mich ( nines ) Cl
Ap - Alpena. Mich.

101
Harbor in the Upper Peninsula of Michigan. The Copper Harbor
leaves are on the whole more deeply cut than the other popu-
lations but their basic pattern did not appear different. The
following populations were distincly different from those listed
above with leaf outlines that graded toward those of black oak
and scarlet-Hill oak: Alpena, Honor Dunes, Tawas, all from
Michigan and Cass Lake from Minnesota, see Figs. 27, 28. Com-
pare with leaf outlines on pp. 47,76. The Tawas population was
a small sample taken in conjunction with a black oak sample
but not large enough to handle statistically with the rest of
the red oak p0pu1ations.

A study of winter buds showed no pattern of variation in

s... Fig. 12..
bud length. 'Sce-FrgT-ffia The presence or absence of pubescence
on the winter buds is shown in Table XIX.

Trees appearing intermediate between red oak and some other
oaks were conspicuous in the following populations; Honor Dunes,
Alpena, Muskegon, and Cass Lake. In the first three populations
these trees appeared intermediate between red oak and black oak.
In the last population the trees appeared to be intermediate
in their field characters between red oak and the scarlet-Hill
oak complex.

Field observations were made of spring frost damage among
red oak populations in northern Michigan on three consecutive
years. No differences in resistance to frost damage were ob-

served except those related to time of leafing out.

102

Fig. 26 A comparison of variance of 13 red oak populations.
The height of the bar represents the number of
characters for each population in which the variance
is significantly greater (F test at 1% level) than
that of the p0pu1ation having the least variance for
that character. e.g. The Honor p0pu1ation (Ho) had
significantly more variance than the p0pu1ation with
the least variance for that character in 11 out of 13
characters.

of characters

no.

124
'4
10.

1

A

I A A

1. ANI

 

‘yt

L—

 

 

 

 

 

 

 

PoQWd En Mp Bp Ho Ap Bl D1 ‘bn RawCh Cl

Sampled populations ordered south to north basis

Po
Wd
En
Mp
Bp
Ho
AP
Bl
D1
Dn
Ra
Ch
Cl

Pokagon State Park, Ind.
'Warren Dunes State Park, Mich.
East Lansing, Mich. '
Muskegon State Park, Mich.
Benzie State Park, Mich.
Honor, Mich. (dunes)
Alpena,‘Mich.

Burt Lake State Park, Mich.
Douglas Lake, Mich.

Dunbar Forest Station, Mich.
Raco, Mich. '

Copper Harbor, Mich,

Cass Lake, Minnesota

103

Fig. 27. Representative leaves from populations of Qggrcus
rubra showing a typical shade leaf on the left and
two kinds of sun leaves on the right.

 

Fig. 28. Representative leaves from populations of gggrcus
rubra var. borealis showing a typiCal shade leaf
on the left and two kinds of sun leaves on the
right.

 

 

104

Table XIX. Number of trees in each population of red oak
sampled showing presence or absence of pubescence
on the winter buds.

Population Essentially Conspicuously
glabrous buds pubescent buds
Pokagon State Park, Inc. 3 0
Warren Dunes State Park, Mich. 5 0
East Lansing, Mich. 6 2
Muskegon, Mich. 7 3
Benzie State Park, Mich. 8 1
Honor Dunes, Mich. 2 3
Alpena, Mich. 9 5
Burt Lake State Park, Mich. 6 1
Douglas Lake, Mich. 8 3

Dunbar, Mich. ll 0

105

2. Nursery Studies. The number of 99ercus rubra and

 

 

Q. EEQEQ var. borealis progeny established in the nursery was
small because it was not possible to obtain seed material in
the fall of 1960 from the more southern areas due to a poor
crop. As a consequence there were too few p0pu1ations on a
south-north axis to give a very complete picture of trends.
The Holland, Mich. population, the only one that might be con-
sidered southern, consisted of only one female parent tree.
Also the Honor Dunes nursery p0pu1ation came from a field pop-
ulation in which certain individuals appeared intermediate
between red oak and black oak.

In spite of the poor p0pu1ation representation 7 out of
23 character measurements in the nursery showed significant
differences in their means, 3 at the 5% level and 4 at the 1%
level (Table XX). This appeared to show a degree of hetero-
geneity among the red oak nursery populations.

South-north trends were evident in 13 out of 23 character
measurements. These included leaf length, leaf width, leaf
shape, leaf size, leaf margin, leaf angle, degree of red petiole,
% green leaves, % red leaves, % 2nd flush, height growth, %
leafed out, and % leaves lost. Of these 13 character measure-
ments that showed south-north trends, there was a significant
difference among their means in 2 at the 5% level and 4 at the
1% level. 0f the 10 characters that showed no south-north

trends only one character, % second flush (1960 scoring), showed

106

Hm>ma NH we“ um unmUHchwam n a«
Amocwauw> mo mfimhamcmv Ho>oH Nm new on unmoamacwam n a
ucmuamacm.m uoz n mm

 

 

 

 

 

m2 m2 m2 m2 museum Mafia}

5. m.H muu.mficwu%.cww3umnv m5Hw> m

. o .eUHz ..mum ammuom sweeps

oooqa mm A oma ems o .eoaz .mnpmz .mum uflsmm

ooSH mo N3 owe m .532 .oqu mmeSom

oonH as Noa ewe m .eoaz_..xm .um meme swam

ooomH . mo Hma was 0 .euaz .Awwquev nocom

oesea mOH NeH ems , m .30“: ..em mumum maucmm

comma moa moa 5mm H .50“: .uawHHo:
umumumEHHHHE mhmumEHHHHE mumumEHHHHE Hones:
- N awe N owe. N owe N.MMm nook»

suowB x numama :uoaz nuwcma ucmhmm coauooHHoo

Emma new; name unwamm mo umpanz I no suaamooe

 

.cowuomaaoo comm ooaa .mawmn nunoc
ou LuSOm co newcwhhm mmauaawooa mSoHpm> CH omuomHHoo.vmom Eonm .noaz
.wcamamq .m an munch N Mom CBOMm mmcHHoomm xwo ooh mo mhmuowuwso suaouo .Nx wanes

107

Hm>wH NH men um ucmonHamHm as
Amocmflnm> mo mHmhamcwv Hm>oH Nm msu um unonmwcme «
unmonHcmHm uoz n mz

ll

 

 

 

 

 

mz . aa a as «a a museum :aHeuea
H.H ¢.¢m m.m o.q m.m Aquuomwuwcmo3ume~ msam> m
. ma. m.m .soHE «.mum ummhom unease
no mo N NH ‘ «.0 .eon_.mHumz .mum qumm
mm. «a H wH H.o .aon..mawH memsom
mm «N s OH N.o .eon ..Hm .um mHmH sham
mo HN ma MN m.m .nowz..fimocfivv Hocom
ooa - ON «a ma , o.o .Lon ..xm mumum mHNcmm.
OOH mH . mm mH m.s .eon .eamHHom
mowuw ucmwpmm unmonmm mommwwo oomnw
N owe N ems H www H «mm H «mm
No\NH\o . Nm\q\oa Ho\w\HH Ho\ma\n
mHoHuoa mm>me . .. -
ooh can
we omumoo nuHB amouw . mawaw aHmumE SOHuomHHoo
mocha moons mm>woA mmmq mama mo huaawooq

 

.coHuomHHoo omom coma .mHmmn canoe
ou runom Go emmcwnhw mmHuHHwooa mDoHHw> cH omuooHHoo.vomm Eonm .non
.wchcwH .m um mama» N Mom caouw mmcHHommm xmo emu mo mumuomnmno nuaouo .ucoo .xx oHemH

108

Hm>mH NH men um namOHchme

 

 

 

 

n ««
AmocwHum> mo mHmhawcmV Hm>oH Nm mnu um pamOHMHamHm n «
ucmonchHm uoz n m2
m2 m2 s ea mz museum adqnuwa
«.0 m.¢ q.qa N.N : m-. accflumtcom3ume~ osHm> m
z. . t . N.O .eon «.mum ammuom panama
ON OH NN OO N.O .non .oHumz .mum uHsmm
NO N «N mm H.N .SUHz .mme mmesOO
NH a HH mm O.N ‘ .eon_..xm .um mme seem
mm O N. Os H.N .eon .Hmocsev uoaom
mN O N mo O.N .eon ..xm mumum «Huamm
m w m om m.m .non .ocoHHom
unmopmm unwoumm unmoumm unmouom memnw .
H owe N own N own N own H owm
HO\oN N0\m\NH N0\©\HH Nm\m\m. H0\©N\o
UmmmEmU .. . . . ..uDo _MOHoo
umoum wchHmth umoa wafimwma . mcHumm coHuooHHoo
mm>mm4 mm>mmq mm>moq mm>wmA mm>mmA mo huHHmooq

 

.coHuooHHoo comm coma .mewn nuhoc
ou susom co owwGMHum moHuHHwooa msownw> GH UmuooHHoo comm scum .soaz
wcwmcwu .m um when» N Mom cBOMw mwcHHommm xmo no» mo mhmuomhmno :u3onw .ucoo .xx oHan

109

Ho>oH

Amocwanw> mo mamhawamv Hm>oH

NH men um uanHchmHm u a«
NO men on pcOOHchme n *
unwowmacmHm uoz n mz

 

 

 

m2 m2 s m2 m2 .Il museum nafifiwmw
N.N H.N n.m bmaflmwmmmudmwwaumnv mDHm> m
0.0 OO ON OO .:OHz ..uum ammuom Hansen
O.N mm mm «m mm .non..oHuwzv.oum uHDmm
0.0 OO OO NH OO .aOHz_.mqu muHOOOO
0.0 mm OO NH ON .SUHz...am .um mme unsm
0.0 ON on «O OO .nOHz_.Hmmg:OV uocom
0.0 . NN OO OH OO .eon ..xm mumum «Hucmm
O.N . OO HO NO OOH .30 z .OanHom
ammo unmouom unmonmm unmouom ucndnom
H www N «mm H mwd N own A «we
owmewc
umoum HOOH
HONOHNO NONHHNO HONONN maHnam wcHuam
Eonw.m%mo Sanam., _ nouns vommawo
.oEHu mocmw uaooom suH3 wcaxhom ,umohm coHuooHHoo
uhmaw mcHHoowm moons momma mm>wmA mo muHHwoou

 

.GOHuooHHoo comm coma .mHmmn nuhoc

ou suSOO co oomcwhuw moHUHHwOOH OSOHnm> SH nouooaaoo.ooom Eoum .non

.wchGmA .m us when» N How nsouw mmaHHooom xmo ooh mo whouomhwso nu3onc

.ucoo .xx mHema

110

Hm>mH NH men um OchHHchHO
Amocmanm> mo mHmkchmv Hm>oH Nm map on unmonHcmHm u s
ucmuHHchHm uoz u Oz
Oz mecmum naHnuee
h; wvmwum.mmm3umn~ msam> m

H
“X
-3<

 

 

.eon «.mum ummsom Humane

 

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OO .Oon .mamH mmHmsOO
OH .eOHz ..xm .OO mme Haze
me .soHE .Ammcnvv hoaom
om .soHZ ..xm mumum oHNCmm
OO , .eOHz .OamHHom
UCmUMmm.
H own
Noma
muoosm
ou mwmawo coHuooHHoo
“Hague Ho NuHHmUOH

 

.coHuooHHoo comm coma .mHmon sunoc
ou nuDOO co Ummcwpuw moHuHHwooHn mDOHuw> CH UmuooHHoo comm Eoum .non

.maHmch .m um munch N Mom c3onm mccHHomom xmo ooh mo whouowhmso nuBOHO .ucoo .xx manme

111
a significant difference among the populations at the 5%
level.

Careful study failed to reveal any basic differences
among the nursery p0pu1ations in,pattern of leaf outline.

. Winter buds were collected from the 1900 planting in
the winter of 1902 representing the following populations
from Michigan; Holland, Benzie State Park, Honor Dunes, Burt
Lake State Park, Douglas Lake, and Sault Ste. Marie. No dif-
ferences in winter buds, in size, shape, or character could
be detected by observation.

One mass planting of acorns from a suspected hybrid tree
from Wilson State Park, Michigan, was made in the fall of 1961.
The parent tree appeared intermediate between red oak and
either Hill oak or black oak. Leaves from 175 seedlings re—
sulting from the planting were collected and classified ac-
cording to leaf outline. They could be assigned to three major
types. See Fig. 29. A comparison of these leaves with other
leaves in the nursery of comparable age indicates that 47 of
the leaves resembled Hill oak leaves and 79 red oak leaves.
45 leaves were of an intermediate type.

Frost damage studies were made on the red oak nursery
p0pu1ations in the spring and fall of 1961. There were no
significant differences among the p0pu1ations in spring or

fall frost damage, (Table XX) or in percent forking of the

leading shoot after spring frost damage. There is an indica-

112

Fig. 29 Basic pattern of leaf outline for seedlings
grown from a suspected hybrid tree from Wilson
State Park, Mich. The largest terminal leaf
was collected from each seedling for this
study. Number of seedlings in each pattern,
47 on left, 45 in middle, 79 on right.

 

113

tion in Table XX that southern red oak populations may be
less susceptible to frost damage, but it was not significant
at the 5% level.

There were no significant differences among the p0pu1a—
tions in percent rabbit damage during the winter of 1961-02.
The rabbits appeared to work in certain areas of the nursery
planting rather than on species groups. This seemed to be

related to amount of nearby cover and snow depth.

Part IV Discussion
General Statement

Black Oak

Scarlet-Hill Oak Complex

Red Oak

114

115
A. General Statement

Complexes of natural interbreeding populations exhibiting
variation both between individuals and between local p0pu1a-
tions occupying different environments have been described by
a number of workers. In many cases part of the field observed
variation can be attributed to genetic differences such that
even when grown in a common garden or under controlled conditions,
individuals from these p0pu1ations remain different, Turesson
(1922), Clausen g; a1. (1940, 1945, 1958), McMillan (1959),
Goodwin (1941, 1944), Mooney and Billings (1961), wright (1959),
01sen.§£ g1. (1959), Kriebel (1957), Vaartaja (1959). Genetic
variation across environmental gradients has been specifically
examined by several of the preceding with the work of MCMillan
being possibly the most nearly concerned with the relationship
of the genetic differences to the environmental gradient.

Stebbins (1950) in commenting on the variation patterns
in the genus 92ercus notes that Ascherson and Graebner (1913)
did not find well—defined geographical races in the European
white oaks although the species are quite variable and apparent
hybridas frequent. With regard to American oaks Stebbins makes
the following points:

1." "...that a much larger number of species (than at the

time of writing) will be found to possess geographic
variation when their intraspecific variation patterns

are studied more carefully."

2. "...hybrid swarms are often found in regions where the
ranges of markedly different allopatric species overlap."

Hill,

116

"Species may occur together in many regions without
showing any signs of intergradation, but in other
areas these same species may be connéctédl by both
first-generation hybrids and other intergrading types
which are the result of segregation or backcrossing."

"The evidence available suggests that the number of
genes by which species of oaks differ from each other
is considerably smaller than it is in the cases of
most other plant groups."

Referring to Palmer (1948); "the small percentage of
oak hybrids in nature in comparison to the total p0p-
u1ation is not due to the difficulty of producing
interspecific hybrids, but to the small chance which
the hybrids have of becoming established."

"In 92ercus therefore, we have a rather large number
of species, many of them polytypic which are often
capable of exchanging genes with other, morphologi—
cally distinct populations occurring in the same
region and in similar habitats. These sympatric
p0pu1ations have always been kept apart as distinct
species, but some of them, such as g. bicolor and g.
macrocarpa, behave genetically more like subspecies."

 

In initiating the present study on Quercus ellipsoidalis

three more general questions were in mind.

a.

In the highly variable red oak group does the observed
field variation show any tendency to reflect environ-
mental differences?

In common garden studies of seedlings grown from
acorns collected in the various p0pu1ations, do
variation patterns show any relationship to environ-
mental patterns?

Is interspecific hybridization suggested either by

the field or the common garden variation and if so

does this hybridization appear to be related to en-

117

vironmental patterns?
Each of these questions involves consideration of the environ-
mental patterns in the area. The most singular environmental
feature in the region is the vegetationai, climate and soil
boundary approximately corresponding with the separation of
the Hemlock - White Pine, NOrthern Hardwood Climax Forest
region from the Beech Maple Forest region (Braun 1950). Such
a change in vegetation, climate and soil should result in dif-
ferent selection pressures and thus give some evidence of
genetic differences on a south-north basis. This is suggested
in all three species complexes studied.

Since different characters are used in characterizing the
field and the nursery p0pu1ations the trends of the field can-
not be compared directly with trends among the nursery p0pu1a-
tions. e.g. nursery seedlings were too young to obtain acorn
characters and even the foliage tended to be juvenile in first
year seedlings. Thus the observation that acorns are smaller
in northern p0pu1ations cannot be assigned to genetic differ-
ences although it is reasonable to expect that the highly sig-
nificant S-N differences among the various seedling populations
in the nursery are of genetic origin.

In those characters observed in the field for which there
is a comparable character observation from the nursery, for
example leaf size, twig pubescence, and leaf pubescence in

black oak the trends were similar. Though leaf pubescence

118
and degree of twig pubescence were not scored in the field
p0pu1ations (presence or absence of twig pubescence not de-
gree of pubescence was scored, see methods), experience
with these p0pu1ations left no doubt as to a reduction in
the degree of expression of pubescence in the northern popu—

lations in these two characters in black oak.

119
B. Black Oak

Black oak, according to Fernald (1950), is found in dry
woods from north Florida to east Texas, north to south Maine,
center New Hampshire, Vermont, New York, south Ontario, Michigan,
Illinois, south Minnesota, and southeast Nebraska. The region
in this study would include the northern limits of black oak
in Michigan and some of its variation in Indiana. All of the
p0pu1ations (except McCormick's Creek State Park) were found on
a type of sandy well drained soil of Wisconsin or later age.

See Table I for specific soil types.

Black oak is common in the study region up to its northern
limits in the Lower Peninsula of Michigan where it is replaced
by northern red oak (9. 32253 var. borealis).

The classification of black oak has been studied by Trelease
(1912, 1924) and most recently by Camas (1952). Fernald (1950)

lists four forms named by Trelease (forma missouriensis, f.

 

macrophylla, f. dilaniata, f. pagodaeformis). Rehder (1940)

 

 

 

lists only the species and one variety Q. velutina var.

missouriensis Sarg. Gleason (1952) and Gleason and Cronquist

 

(1963) list no varieties or forms. It thus appears that the
present taxonomic concept of black oak includes few named forms
and varieties and that the tendency is to reduce the number of
these.

That black oak in the field in the study region presents

a heterogeneous population is readily evident from the field

120
data in which all 13 field characters showed a significant
difference among the p0pu1ations at the 1% level (analysis
of variance). Differences wére also found in the leaf outline
studies and in the winter buds studies though these were not
handled statistically.

Acorn size showed a distinct reduction northward, those
populations below the tension zone (Fig. 1) averaged (length
plus width in mm.) from 28.5 to 32.0, and those from north of
the tension zone 25.8 to 28.4. See Appendix C.

These significant differences in acorn size warrent at-
tention in their own right. They could be due to genetic dif-
ferences or to environmental differences or both. That en-
vironmental conditions alone are unlikely is suggested by two
lines of evidence. Individual black oak trees on different
soil types for which I followed the acorn crOps over a period
of years tended to retain their differences in fruit size

(Fig. 31).

 

Also Gysel (1957) reported acorn weights of g, velutina for
medium and poor oak sites, per 100 feet of crown area for a
four year period. The one stand (medium site on Fox sandy
loam) was from the Lansing area (Rose Lake Wildlife Experiment
Station) and the other (poor site on Grayling sand) was from
about 40 miles north of Lansing (Flat River State Game Area).
Calculations made from his results indicate that acorns from

the medium black oak site weighed .00306 lb. per acorn and

121
those from the poor black oak site weighed .00318 lb. per
acorn. These 4-year records indicate that black oaks on
more droughty,, sandy sites do not necessarily have smaller
acorns than those on less droughty site and suggest that the
gradient in size indicated in Fig. 30 is genetic rather than
environmental in nature.

Thus, small—fruited individuals are occasional in the
Lansing area, more abundant northward and not observed south- '
ward, although occasional individuals with abnormally large
acorns do occur southward. This is suggestive of selection
Operating in favor of individuals with small acorns northward
or with large acorns southward, and leads to the question of
whether acorn size is associated with any other observed
characteristics.

Acorn size has long been known to be associated with seed-
ling vigor, Kfirstian (1927). Cieslar (1923) presents evidence
of its influence on height growth in seedlings in the white oak
group up to seven years and Johnson (1951) up to 18 years.
Johnson also presents evidence of the influence of acorn size
on the percentage of germination in the white oak group. The
evidence in this dissertation appears to suggest a strong in-
fluence of acorn size on height growth the first year, but little
the second. The influence on leaf dimensions appears to be
present in both years. See Table V.

Small acorns might be associated with characters which '

Fig. 30.

Fig. 31.

122

Acorns from the 1961 seed crop used in nursery
planting of black oak and arranged from left to
right on a south to north basis as follows:

Dunes State Park, Ind., Elkhart, Ind., Rose Lake
Wildlife Experiment Station (Lansing, Mich.) Mich.,
Muskegon, Mich., Pentwater, Mich., Wilson State
Park, Mich., Traverse City Fair Grounds, Mich.

 

Acorns from two black oak trees near E. Lansing,
Mich. showing the variation in size of viable
acorns. The two trees were about 60 feet apart
on a similar soil type. The two trees were ob-
served for three fruiting seasons and the size
relationship was always the same.

 

123
have selective advantage northward. An observation that
points in this direction is a consistently higher number of
acorns produced by the smaller fruited black oaks observed in
the Lansing area as compared with the associated large-fruited
ones.

From the limited data now in hand, it is not possible to
determine the significance of the pronounced shift to small
acorns in black oak in the northern part of the species range
in this study. It is clear that acorn size is variable and
must be used with care in diagnosing species. There is some
suggestion of a break between the northern and southern field
populations based primarily on data from acorns. See Figs. 8,
9, 10. Also the occurrence of occasional small-fruited indi-
viduals south of the tension zone in the Lansing, Michigan area
suggests this northern expreSsion can behave as a reasonably
discrete entity. Further work may suggest that rec0gnition of
a northern race is warranted comparable to the northern race
in red oak, Q. gghgg var.borealis.

In reference to variation in black oak resulting from the
study of seedlings in the nursery, Schreiner and Santamour (1961)
came to the following conclusion based on their work with 17 black
oak female parent sources from 9 eastern states;

"...no pronounced tendencies were observed that would in-

dicate racial variation except fall coloration in progenies

from localities with longer growing season tended to be re-

tarded. There was wide individual variation in the factors
effecting growth."

124
This conclusion was based on the reported scoring of three
characters, percent germination, average height at 1 year and
3 years, and autumn color rating. The progenies were replicated
twice.

I found, on the other hand, a difference in height growth
significant at the 1% level for the 1960 planting the first
year and at the 5% level after two years of growth (analysis of
variance). However, the growth rates the first and second year
were not related. Certain populations with the slowest growth
rate the first year (Indian River, Honor Dunes) were among the
tallest the second year. Conversely, the tallest p0pu1ation
the first year (McCormick's Creek State Park, Ind.) was among
the slower growing the second year. Table V. In the 1961
planting there was no significant difference in height growth
in the seedbed that had been mulched in the fall with sawdust.
There was ac significant difference at the 5% level in the un-
mulched bed. See Table VI.

Thus one may interpret the different findings on height
growth between Schreiner and Santamour and the present study in
at least two ways. First, that the Michigan populations of
black oak exhibit greater genetic differences related to growth
than the populations studied in eastern United States. Or,
secondly, as Cieslar (1923) and Johnson (1951) have shown, seed-

ling growth is strongly influenced by the size of the acorn.

These differences might be essentially lost by the 3rd year and

125
the differences in the present study (which were less signi-
ficant in the 2nd than the lst year) would be due primarily to
differences in seed size.

In fall leaf color in the 1960 planting, I found no signi-
ficant difference in percent green leaves in the lst year read-
ing nor in percent red leaves in the 2nd year reading. However,
there was evidence of a trend in percent green leaves. There
was no significant difference in percent green leaves or percent
red leaves in the mulched seedbed though again there was a dis-
tinct south-north trend. In the unmulched seedbed there was a
significant difference atrth'el percent level for both percent
green leaves and percent red leaves. Table VI. These results
of fall coloration appear to confirm the work of Schreiner and
Santamour.

In addition to the latitudinal variation pattern illustrated
by the acorn size, small scale variation is also present. A
comparison of the nursery populations from Benzie Dunes and
Traverse City State Park indicate that it is possible to have
genetic distinctness among black oak populations in an area not
much larger than a county. See Table V. These two p0pu1ations
are approximately 25 miles apart.

Some aspects of the variation pattern in black oak, both
from.nursery and field data, strongly suggest introgression be-
tween it and other species. Such a view gains support from the

work of Stebbins gt El. (1947), Tucker (1951, 1961), Tucker and

126
Muller (1957), Cooperrider (1957), Sillman and Leisner (1958),
and Benson (1963). The two taxa most suspect in the northern
part of the study area are 9. Egbgg var. borealis and 9.,
ellipsoidalis. '

Intermediate trees between black oak and northern red oak
were not uncommon in the northern stands. For example in the
Muskegon State Park, Alpena, Honor I and Honor II populations
they were conspicious. In the only one of these in which nursery
data were available (Honor Dunes p0pu1ation) there was some evi-
dence of segregation of the buds and twigs in the seedlings of
certain parents into two groups, those resembling black oak and
those resembling red oak. (See p. 60).

Intermediates between black oak and Hill oak are suggested
by fall leaf color, inner bark color, and winter buds. Typical
fall leaf color in black oak p0pu1ations is yellow while typical
color of Hill oak is red. In the Grayling area fall color varies
from yellow to red-brown with many intermediates suggesting that
there is an intermixture of black oak and Hill oak. This inter-
mixture is also suggested by the intermediate nature of the inner
bark color in the Indian River population. It is a lighter color
than any of the other black oak populations except Alpena (See
Fig. 10), which is itself an intermediate population but between
black oak and northern red oak. The intermediate nature of the
Indian River population is also quite evident in the nursery par-

ticularly in leaf and twig pubescence. See Tables V and XIII.

127
Also nursery prOgeny of the Indian River p0pu1ation have winter
buds that appear to segregate into three groups, those resembling
black oak, Hill oak, and red oak.

Fig. 11 on page 45 depicting a comparison of variance in
the field characters is particularly useful in this discussion
of intermediate populations. It is assumed that p0pu1ations
with intermediate trees should show a considerable increase in
variance over normal populations. That this is the case in black
oak is indicated by Fig. 11. The p0pu1ations with the highest
index of variance are Alpena, Wilson, E. Tawas, Honor I and II,
and Traverse City Fair Grounds. This corresponds almost ex-
actly with field experience with these populations. The inter—
mediate nature of Wilson State Park stand is discussed under
red oak in this dissertation. The E. Tawas stand was the only
one in which the intermediate nature of certain red oak appear-
ing trees was not readily evident in the field. Later study of
the leaf collections in the laboratory suggested certain of
these trees may be intermediate between red oak and black oak.

Twoacharacters that appear to be useful in identifying
individuals intermediate between black oak and red or Hill oak
are the shape of the apex of the acorn cup scale and the shape
of the acorn. Typical black oak has obtuse scale apices and
round to oblong acorns; typical red oak has rounded shaped
scale apices and round to oblong acorns; while typical Hill oak

has acute to acpminate scale apices and ellipsoidal acorns.

128
Whenever a black oak or a red oak is found with either acute
or acuminate acorn cup scale apices or particularly narrow

shaped acorns, it suggests a past gene exchange between any

two of the three species above.

129
C. Scarlet-Hill Oak Complex

A major cause of confusion in studying Quercus coccinea

 

and g. ellipsoidalis is the general lack of a clear understand-

 

ing of the limits of the latter, Hill oak. In looking over
labeled herbarium sheets in many midweastern herbaria, I have
come to the conclusion that frequently any specimen from the

red oak group in the Great Lakes area that was of a questionable
nature was assigned to the Hill oak taxon. This includes speci-
mens intermediate between any two or more of the following
species, black oak, red oak, scarlet oak, and pin oak.

The species description by Rev. E. J. Hill (1899) assigns
very broad limits to many of the diagnostic characters and this
may be a major reason for its becoming the catch-all among the
red oak group in the midwest. The following statements quoted
from Hill's description of the taxon appear to be the most use-
ful in trying to define it:

1. "...the spray fine and repeatedly dividing, the limbs
often descending low down on the trunk and the lower
most drOOping."

2. "It (innerbark) is coarsely cellular, dull red within,
with a thin band of yellow or yellowish bark 1-2 mm.
thick next the wood."

3. "The winter buds are 4-8 mm. long, ovate, obtuse or
acutish, sometimes slightly angled. The scales are
ovate to oval, ciliolate, the outer brown to reddish-
brown, with a rusty or grayish pubescence, the apex
blunt or rounded."

4. "They (leaves) are from 6-15 cm. long, 5-12 cm. wide,

oval-orbicular or somewhat obovate-orbicular in out-
line, the broadest part usually just above the middle,

130

and are deeply cut into 5—7 lobes by broad sinuses
rounded at the base which extend halfway or nearly
to the midrib. The lobes are generally oblong, the
terminal broader and somewhat quadrangular, their
sides nearly parallel or a little diverging above
the middle. They end in three to five triangular
teeth with slender bristles. The base of the leaf
is bluntly cuneate to truncate, the parenchyma on
one side usually a little lower. The petioles are
rather slender, 2.5-5 cm. long (generally about

4 cm.), usually tinged with red on the upper side."

5. "The color of the autumn leaves varies, but in most
is yellowish to pale brown, more or less blotched
or tinged with red or purple. Some change to a
vinous or crimson purple, giving the tree a dark
reddish appearance. In winter they are pale brown."

6. Acorns.

a. "...the cup turbinate or cup-shaped, thinnish,
covering one third to one half or more of the
nut and commonly tapering into a peduncle 8-15
mm. long."

b. "In some forms the cup is thickened near the
margin, forming a kind of shoulder."

c. "The scales are narrow-ovate, obtuse or trun-
cate, brownish, pubescent, closely appressed,
sometimes a little loosened near the rim on
drying."

d. "The nut is chestnut-brown, often striped with
darker lines, puberulent, 12-20 mm. long, 10-15
mm. wide, ellipsoidal, varying from a cylindri-
cal to a shorter somewhat globular form."

A type specimen of dercus ellipsoidalis was not designated

 

by Hill in the original description. There are five herbarium
sheets of Qgercus ellipsoidalis from Rev. Hill's herbarium in
the Chicago Museum of Natural History herbarium with the nota-
tion "sp. nov." on the label. It is assumed that these were

used as the basis for the description and drawings. The speci-

131
mens at present carry the Field Columbian Museum numbers
(now the Chicago Natural History Museum) but no specific speci-
mens are listed or referred to in the original species descrip-
tion. The numbers and their labels are as follows:

50193, 50195 "Woods along Calumet River near Halstead
Street, Chicago, 111., Sept. 17, 1896."

50194 "Dry clayey ground, Glenwood, 111., Sept. 24, 1896."

50196 no locality, Sept. 23, 1895.

50199 no locality for the herbarium sheet lable but sever-
al specimens are mounted on the same sheet, some of
which are labeled "Glenwood, Ill."

A sixth sheet with the herbarium label of Agnes Chase with

a notation in red ink in large letters "Part of Type" is at the

present in the type folder for Quercus ellipsoidalis at the

 

museum. Its number is T1547 and it has the following label

"Along Thorn Creek, Thornton, Ill. Sept. 12, 1899". A specimen

of flowers on the same sheet was dated May 3, 1899. The leaf
specimen is very similar to the drawing, plate 111, in the species
description. However, the specimen was collected in Sept. 1899

and the species description in The Botanical Gazette is datedIWank

 

1899.

It thus appears that the herbarium specimen of Agnes Chase
(T1547) was collected after the species description was published
and designated at the time of collection or at some later date as
the "type specimen". However, this designation of an Agnes Chase
specimen as the type when material collected by Rev. Hill is

available is not in keeping with the International Code gf Botani-

132
cal Nomenclature, 1960. The code states that as long as any
"element used by the author ... is extant, it automatically
fixes the application of the name concerned". Thus one of Hill's
five sheets should be designated as a lectotype.

Later authors have generally interpreted Quercus ellipsoidalis

 

in a narrower sense than Rev. Hill's description would warrent.
For example, these are major points in the keys of the following
authors; acorns narrowed at both ends, with top-shaped cups,
winter buds pointed, Palmer (1942); winter buds small, 1-3 mm.
long, Wadmond (1933); winter buds small, pointed, Camas (1952).
The populations designated as Hill oak in this dissertation are
also interpreted more narrowly than the species descriptions of
Rev. Hill. The one exception occurs in the handling of the 1960
nursery data. Here northern populations of trees that resembled
black oak except for very small acorns were lumped with trees of
Hill oak (in the narrow sense). These northern black oak—like
p0pu1ations have often been considered a form of Hill oak by
field workers in Michigan and fit reasonably confortably into
Rev. Hill's description of his new species. It was possible after
compiling data from the nursery for two growing seasons to sep—
arate these taxa with reasonable confidence. See p. 87.

As I have interpreted Hill oak, it does not have yellow in-
ner bark in spite of the species description published by Rev.
Hill. Trees with yellow inner bark from the Michigan area, I
have referred to black oak or its variants. In all 40 of the

p0pu1ations studied, including red oak, northern red oak, black

133
oak, Hill oak, scarlet oak, and pin oak, I did not find a
single tree with a yellow band of inner bark between the wood
and an outer reddish band of inner bark as Rev. Hill describes
his species. (See point 2 of the species description previously
mentioned). The populations I examined included a sample of 88
trees from Thornton Woods, Illinois, which is or is very near
the original type locality as interpreted in this dissertation.
This sample included black, pin, Hill, and red oak trees.

A major problem that has concerned workers on Hill oak has
been its relationship to scarlet oak. Trelease (1919), Wadmond
(1933), Palmer (1942), and Camas (1952), have maintained these
two taxa as separate species. Table XXI lists the points con-
sidered to be the most diagnostic in separating these two species.

Rev. Hill even after his published description of Hill oak,
continued to identify a separate entity as scarlet oak at Thornton,
Ill. A sheet from his herbarium dated October 9, 1899 from

Thornton, Illinois was named Quercus coccinea. In the species

 

description for Hill oak, Rev. Hill includes the following
description of the relationship of the two species.

"When first observed in the fall of 1891, with some mis-
givings it was called Q. coccinea, partly on account of
acorns, which in general appearance are more like some
forms of fruit in this species, and partly on account of

a tendency to red, of a purple or crimson cast, however,
seen in some of the autumn leaves. Its leaves also divide
more like those of g. coccinea and it has the reddish tinge
of the inner bark, but the leaves do not turn scarlet in
the autumn, the outer bark is darker colored and much less
rough and broken than on boles of Q. coccinea of similar
age or size. It is also far less lichen-covered than

134

Table XXI. Diagnostic characters of scarlet and Hill oak.

Scarlet Oak
Buds rather large, 3-5 mm.

Acorn shape, oval or oblong

ovoid. Acorns dull colored.

(Wadmond)

Leaves oblong-obovate or
elliptic in outline,
abruptly contracted or
truncate at base.

Winter buds conic-ovoid,
obtuse or rounded at apex.

Upper winter bud scale con-

spicuously pubescent on
margin. (Palmer)

Buds rather large, conical-
ovoid, obtuse or rounded
at apex. (Camas)

Hill Oak
Buds 1-3 mm. long

Acorn ellipsoid to semi-
globose but always taper-
ing to a point. Acorns
shining as though freshly
varnished. (Wadmond)

Leaves oblong-obovate in
outline, abruptly narrowed
or rarely truncate at base.
Winter buds pointed. Upper
bid scales inconspicuously
cilate-pubescent or nearly
glabrous. Acorns narrowed
at both ends with top-
shaped cups rarely over
1.5 mm. wide. (Palmer)

Buds small, pointed. (Camas)

135

trunks of Q. coccinea (generally it is comparatively

free from 11cEEEEIT‘EOr the scarlet oak seems pre-

eminently chosen among the black oaks by these plants,

which sometimes are so close and large as nearly to

conceal the color of the bark and at once reveal its

specific character."

Later Trelease (1919) and subsequent botanists in that
area eliminate scarlet oak as a local species and named all of
the scarlet oak-like trees as variants of Hill oak. Deam (1953)
lists scarlet oak for northern Indiana, and it is interesting
that the range of scarlet oak as described by recent authors
(Palmer, 1942) tends to follow the Indiana-Illinois state line.
The late Dr. wendell Camp in personal conversation with the
author strongly expressed the View that Q. coccinea or trees.
closely related to it did occur in northern Illinois and southern
Wisconsin.

From extensive traveling in Michigan, Indiana, and northern
Illinois, from a survey through Wisconsin and Minnesota and from
the study of herbarium specimens at many of the major herbaria in
the midwest, I now question the separation of Opercus ellipsoidalis
as a separate species. As a working hypothesis, I have consid-
ered Q. coccinea and g. ellipsoidalis (in the strict sense) to
form a large complex without sharp discontinuities. The field
and nursery studies were designed to test this hypothesis.

From a careful study of the graphs of the morphological
characters of the field populations, the tables of winter bud

characters, and other diagnostic characters, there appears to be

three major groups of populations represented in the study area:

136
l. A southern, large fruited group with large more rounded
winter buds and frequent pitting at the apex of the acorn.
2. A middle region large-fruited group with small more pointed
winter buds, with no pitting at the apex of the acorn, and oc-
curing in northern Indiana, northern Illinois, and southern
Michigan south of the tension zone. 3. A northern, small-
fruited group with small, generally pointed winter buds, no
pitting on the apex of the acorn, occuring north of the tension
zone in lower Michigan and probably in Wisconsin. The first
group would fall within the present interpretation of Q. coccinea,
the latter two within the range of variation recognized for g.

ellipsoidalis.

 

The-problem is three-fold:

a. Is the variation observed in the field due primarily
to genetic differences between localities or is it a
plastic morphological response of a more-or-less homo-
geneous genetic system to a pronounced environmental
gradient?

b. If the variation in the complex is genetic, is it
essentially continuous or does it have clear dis-
continuities?

c. Do the discontinuities, if present, represent sufficient
impediment to gene flow to constitute a break of specific
rank or is it more nearly sub-specific in nature.

It would be presumptious to attempt to answer such difficult

137

questions in any final way with the limited data now in hand.
However, the field data are not incompatible with suggesting
one discontinuity at the approximate location of the transition
zone (Fig. 15). The nursery data on the other hand fail to in-
dicate any such clear discontinuity and for 9 out of 13 characters
suggest nearly continuous variation from south to north (See
Table XIII). A study of the pattern of leaf outlines from the
nursery also showed no basic difference from south to north
except for a tendency toward smaller leaves in the more northern
populations. Since the field data includes the taxonomically
more useful acorn characters not available from the nursery,
the possibility of a genetic discontinuity is not ruled out.

Another of the problems associated with Hill oak has been
the occurence of populations or individual trees that have been

intermediate between Hill oak and black oak, red oak (guercus

 

rubra), northern red oak (Quercus rubra var. borealis), and pin

oak (guercus palustris). It is these variants or "odd—balls"

 

that are found deposited in herbaria which have caused consid-
erable confusion as to just what constitutes Hill oak. Curtis
(1959) reports that intermediates between Hill oak and red oak
and black oak have constituted a problem in identification in
ecological field studies in Wisconsin.

Two intermediate p0pu1ations were studied in the nursery,
Honor Dunes and Indian River. The Honor p0pu1ation was inter-

mediate between a northern expression of black oak and northern

138
red oak. The Indian River population appeared to be inter-
mediate between Hill oak and black oak. Due to its yellow
inner bark, heavily pubescent winter buds, and more or less
pubescent twigs, it was considered a variant of black oak.
The nursery studies appear to bear out this assignment of the
population. As far as I can tell, this intermediate p0pu1ation
occurs widely in and near southern Cheboygan County, Michigan,
in suitable habitats indicating that intermediate populations
may cover considerable area.

Trelease (1919) in his article on Hill oak in Illinois in-
cludes a photograph showing variation in acorn forms. The small
globose forms pictured are almost exact replicates of the small-
fruited black oak forms such as found in the Honor Dunes popula-
tion. It also includes types very much like those in the Indian
River intermediate p0pu1ation. This confusion of a small-fruited
black oak type and Hill oak in the strict sense, has been a major
stumbling block in understanding the Hill oak problem. See Figs.
_30, 31 for some examples of small globose black oak acorn forms.

The only published study known to the author involving
regional progeny testing in scarlet oak provides further in-
sight on the scarlet-Hill complex. Schreiner and Santamour (1961)
report on a progeny test of scarlet oak from four areas, Virginia,
Tennessee, Illinois, and Alabama, involving 7 parent sources.
Their conclusions based on three kinds of measurements, percent

germination, average height at 1 year and 3 years, and autumn

139
color rating were as follows;

"The few provenance of these species included in the

study showed no evidence of racial or ecotypic varia-

tion. In 9. coccinea there was a highly significant

difference in fall coloration between the Alabama

progeny and the average of all seedlots, but the data

are not sufficient to conclude that there is a north—

south trend in this character."

The evidence in this dissertation indicates a south-north
trend in fall leaf color. Table XIV. However, Schreiner and
Santamour's lack of evidence for racial or ecotypic variation
or south-north trend is in distinct contrast with the south-
north trends that appear in the Michigan data.

It must be emphasized, however, that the seedlings I studied
were grown from acorns which differ in size, the acorns of the
southern populations being at least twice and in some cases,
three times the size of those from the north. Since Johnson
(1951) reports an influence of acorn size on percentage germina-
tion and height growth for the white oak group in EurOpe, some
of the seedling differences reported in the two scarlet oak
studies may also be related to this source of variation.

It would appear from these two very inadequate studies that
Q. coccinea is relatively homogeneous over much of its range ex-
cept in the Great Lakes Region where a small-fruited form is
found which is related to climatic and site conditions. This
evidence is not interpreted to mean that there are no differences

between these p0pu1ations. They are different, but the differ-

ences between them are not as great as their similarities. The

140
south-north variation pattern in the study area based on field
and nursery data, appears to indicate a complex of closely re-
lated infraspecific entities rather than two or three species.

Occasional reports of Quercus ellipsoidalis from the east

 

coast have crept into the literature, Grier (1924), but have
been disregarded due to the fact that they would be completely
outside of the designated range. It would appear normal in
terms of the concept of the scarlet—Hill oak complex prOposed

in this study for individual trees and possibly even popula—
tions of Hill oak—like trees to be found where ever scarlet

oak is found. If the range as listed in the manuals were not
considered definitive by field taxonomists, there is no doubt

in my mind that a cr0p of Hill oak identifications would sudden-

ly occur throughout much of the range of scarlet oak.

141

D. Red Oak

The red oak populations in this study can be divided into

 

two groups, those resembling Quercus rubra and those resembling

Q. rubra var. borealis based on the following criteria:

Q. rubra

Little or no pubescence on twigs.

Shade twigs brittle

Branching of trees is Open
form

Apex of acorn cup scale tend
to be rounded

Acorn cup lip shows very little
or no evidence of fringe.

Acorn is large except in the
most northern populations,
generally more than 21 mm.
in length and 18 mm. in width

Acorns are nearly round with an
index (width/length x 100) of
85 or more.

Q. rubra var. borealis

Some evidence of slight
pubescence on twigs

Shade twigs have a tendency
to bend before snapping

Branching of trees tends to
be twiggy.

Apex of acorn cup scale tends
to be acute or more acute.

Acorn cup lip generally shows
some evidence of fringe.

Acorn is generally less than
19 mm. in length and 16 mm.
in width.

Acorns are more narrow with
an index less than 85.

Using these criteria the populations in the present study

can be grouped as follows:
Q. rubra

Pokagon State Park, Ind.
warren Dunes State Park, Mich.
E.Lansing, Mich.

Dunbar Forest Station, Mich.
Raco, Mich.

Copper Harbor, Mich.

Intermediate

Muskegon State Park, Mich.
Benzie State Park, Mich.
Burt Lake State Park, Mich.

142

Q. {3253 var. borealis

Honor Dunes, Mich.
Alpena, Mich.
Douglas Lake, Mich.

Cass Lake, Minnesota

This grouping by the above listed criteria also appears
to be supported by evidence of grouping in the winter bud studies
and the leaf outline studies.

Among the red oak populations are those listed as inter-
mediate between the two groups though generally closer to Q.
Egbgg in most field morphological characters such as the follow-
ing; large acorn dimensions, less tendency for pubescence on the
winter buds and twigs, more round shape to the acorns, less ten—
dency for the presence of fringe on the acorn cup, and a more
round shape to the apex of the acorn cup scale. Also these
populations have a tendency for a more Open branching in tree
form and for more brittle shade twigs. There appears to be a
continuum of populations from those that are distinctly different
from.Q, Egbgg to those that are very similar.

Of the Q, EEQEQ var. borealis field populations studied,
the following contained a conspicuous number of trees intermediate
between red oak and black oak; Honor Dunes, Alpena, and Muskegon,
all from Michigan. The Cass Lake, Minnesota, pOpulation appeared
intermediate in its field characters between Hill oak and red oak.
See p.148 for a further discussion of the Cass Lake pOpulation.

In eastern Upper Peninsula of Michigan and at COpper Harbor,

at the tip of the Keweenaw Peninsula where neither Hill or black

143
oak occur or probably have occured in the recent past, the red
oak, from my observations is Q, EEQEQ (using the criteria pre-
viously established). They suggest Q. gghgg var. borealis only
in acorn size and such a reduction in size in the northern part
of the range of Q. 52233 is to be expected from the trends of
acorn size which were found. See Fig. 23. In the other char-
acters listed for separating the two groups such as twig
pubescence, shape of apex of acorn cup scale, twig brittleness,
acorn cup fringe etc. they are Quercus rubra.

Field experience with these two red oak population types
and their intermediates appears to indicate that Q. 52233 is
found on better soil types and Q. Egbgg var. borealis on poorer
soil types. This is indicated graphically by the soil types on
whch each of the above pOpulation groups is found.

Q. rubra
Pokagon State Park - Boyer Sandy Loam

warren Dunes State Park - Bridgeman Fine Sand (very calcareous)

East Lansing - Hillsdale Fine Sandy Loam

Dunbar Forest Station - Rousseau Laomy Fine Sand
Raco ~ Rubicon Sand

Copper Harbor - Waiska Cobbley Loam

Intermediate

Muskegon State Park - Deer Park Fine Sand
Benzie State Park - Rubicon Sand
Burt Lake State Park - Rubicon Sand

Q. rubra var. borealis

Honor Dunes - Deer Park Fine Sand
Alpena - Rubicon Sand

Douglas Lake - Rubicon Sand

Cass Lake - Cass Lake Fine Sand

144

In the “soils of Michigan: E. P. Whiteside e; 31.. (1959),
the Rubicon Sand is described as having low natural fertility
and low moisture-holding capacity. From this "poor" soil the
soils in this study grade into those that are intermediate in
these qualities such as the fine sands to those of higher fer-
tility and moisture holding capacity, the loams.

The occurence of intermediate trees in many typical Q.
Egbgg var. borealis populations, the apparant relationship of
Q. ggfigg and Q. ggpgg var. borealis to site soil conditions, and
the occurence of Q. ggbgg and not Q. ggbgg var. borealis in the
eastern Upper Peninsula of Michigan and at Copper Harbor, suggest
a possible explanation as to the origin of the differences be-
tween the two groupings.

_Quercus rubra var. borealis appears to be the result of

 

genetic introgression of red oak (Q. Egbgg) by either black oak
or Hill oak and in some cases possibly both. Farther south scar-
let oak replaces Hill oak in this relationship. The occurence of
Q. Egpgg var. borealis on poor sites is a result of the former
or present occurence of either black oak or Hill oak which are
the genetic sources for the variability of these trees which per-
mits them to compete on this type of site.

This explanation appears to fit the picture as it appears in
Michigan. Also it appears to coincide with E. J. Palmer's (1942)
observation of the Q. EQQEQ - Q. Egbgg var. borealis relationship.

He says as follows:

145

"There is evidence to indicate that the two forms do

not 'come true? from seeds. In a row of street trees

planted near the Arnold Arboretum a complete series

of variation in the size of the acorn and shape of the

cups can be found; and several trees grown in the Ar-

boretum supposedly from the same seed, show similar
fruit variations." "...the facts seem to indicate that
one is only an unstable form of the other."

If the hypothesis of the origin of the borealis variety
is correct, their present occurence is a possible indication
of the former presence of either black oak or the scarlet-Hill
complex in the area. For example, in Benzie State Park a
borealis population occurs on the better developed soil some
distance from Lake Michigan (approximately 2 miles) and black
oak is presently restricted to the more recent, more poorly
developed soil nearer the Lake. In the case of the Douglas
Lake population, the nearest black oak is about 9 miles south
but its former presence in the Douglas Lake stand is strongly
suggested by black oak tendencies in certain individuals in the
populations particularly in a gray pubescent acorn cup scale,
the presence of fringe on the acorn cup, and unusually deeply

cut leaves.

It would also be possible for a Qgercus rubra var. borealis

 

type that was competitively superior to black oak on intermediate
soils but inferior to Q. ggbgg on good soils to advance and spread
beyond the present range of Q. Egygg, black oak, and the scarlet-
Hill Oak complex. Though this is a possibility, I feel this would

be more applicable for short distances, unless there was a large

continuous receptive habitat available. However, these conditions

146
could have been present in the several oscillations in vegeta-
tion following the onset and retreat of the Wisconsin glaciation.

I have noticed in the Upper Peninsula of Michigan beyond the
range of the other oaks that red oak (Q. {2253) occurs at least
occasionally on Jack Pine sites (see the Raco pOpulation basal
area table, Appendix A). Southward in the Lower Peninsula of
Michigan, on this type site, black oak or Hill oak would be the
important oaks with possibly an occasional Q. Egggg var. borealis.
This suggests that the absence of red oak (Q. ggbgg) on some of
the poor sites in the Lower Peninsula is not so much an inability
to grow on these poor soils. Perhaps competition from species
better adapted to these sites such as black oak, Hill oak, and
Q. ggpgg var. borealis may keep this species out.

The results from the nursery planting ole. ggpgg and Q. £2233
var. borealis strongly support the presence of south-north trends
in red oak. In 13 out of 23 character measurements (Table XX)
such a trend is suggested. However, contrary to the field data,
nursery results give no support for recognition of a poor site
genetic entity. Part of this may be due to the fact that the only
poor site population in the nursery was from Honor Dunes. The
poor site (Q. Egygg var. borealis) areas that I have observed for
three years have occasionally produced some acorns but not enough
for replication in the nursery. A half a dozen usable acorns would
generally be a good crap from one of the poor site trees. The

red oak trees from.Honor Dunes that produced crops good enough

12.7
for use in the nursery were on the fringes of a poorly de-
veloped dunes soil area and adjoining a good acorn producing
red oak site. The good site areas under observation in the
north during the time of this study and those intermediate
between good and poor sites such as Burt Lake State Park and
Benzie State Park were heavy producers of acorns.

The nursery data present no evidence of a genetic dif—
ference other than that expected in the normal south-north
trends between Q. gyhgg, good site, and intermediate site
Lower Peninsula populations of red oak and populations from the
Upper Peninsula of Michigan (for example compare Burt Lake State
Park and Sault Ste. Marie or Dunbar populations, Table XX).

Due to their erratic or generally poor acorn production,
it may be difficult to obtain good representation of poor site

(Qgercus rubra var. borealis) populations for comparison for

 

future‘studies of the problem in the nursery.

The best diagnostic characters Of the borealis variety are
acorn or mature tree characters (branching, shade twig brittle-
ness) and only small differences occur in other aspects of their
field.morphology. In the nursery I have been unable to detect
any seedling differences between the Q. 52253 and Q. rubra var.
borealis types. It may be necessary to raise the trees to the
point of acorn production to study this phase of the problem.

Schreiner and Santamour. (1961) conducted a progeny test on

red oak from 15 areas in eastern United States including the

148
following states; New Hampshire, New Jersey, Pennsylvania,
North Carolina, Kansas, ILlinois, Michigan, Massachusetts,
and west Virginia. This test represented 23 parent sources
and was scored for germination percent, average height at 1
year and 3 years, and autumn color rating. Their conclusions
were as follows:

"...no pronounced tendencies were observed that would

indicate racial variation, except that fall coloration

in progenies from localities with longer growing seasons

tended to be retarded. There was wide individual varia—

tion in the factors effecting growth."

The species which appeared to be represented in their sow-
ing was Q. {2233 with no mention Of Q. Egbgg var. borealis.

The results of their study in conjunction with the evidence
reported in this dissertation suggest that red oak is rela-
tively homogeneous genetically over much of its range. 0b-
servations further suggest that a small acorn form occurs on
poor sites especially in the north. This small acorn form may
be genetically different (though the differences may be small)
from Q. gghgg. However, I obtained no evidence from the nursery
to support this.

The relationship of the Cass Lake, Minnesota, population
has presented a considerable problem. Acorns from this site
were sent to the author by mail but none germinated. This may
have been due to adverse conditions in mailing between the time

of collection and planting. On the first visit to this site in

the summer of 1960, the population was thought to be a variation

149
of Hill oak. Later on revisiting the area for field study
in the summer of 1962, it was felt that due to the acorn size
and the general appearance of the trees, that it was an inter-
mediate population between red oak and Hill oak but more strongly
resembling red oak. It was placed with the red oak populations
on this basis for study and presentation. After compiling the
field data and comparing the results of this population with
that of other red oak p0pu1ations in all characters, I feel
that it should now be placed with Hill oak and that it is in-
termediate between Hill oak and red oak but primarily leans
toward Hill oak. I have no nursery material from this pOpula-
tion. The evidence from a tree at Wilson State Park, Mich.
intermediate between red oak and Hill oak on pp.lll,lBZalso
indicates the possibility of such an intermediate condition.
The seedlings grown from acorns of this tree showed evidence
of segregation. The problem of the amount of segregation
evident when there is intrOgression, or apparent introgression
in the oaks is not clear. The Wilson State Park suspected
hybrid reported in this dissertation appeared to be an F—l or

close to it.

Part V The Red Oak Group in Michigan

150

151
The Red Oak Group in Michigan1

On the basis of the studies presented, certain features
of the red oak group in the study region are suggested. There
appear to be south-north trends in field measurements, especially
in acorn characters. In the nursery studies south-north trends
also occur indicating genetic differences. A break occurs in
the field morphological characters in both black oak and the
scarlet-Hill oak complex at approximately the transition zone.
Fig. 1. However, if it occurs in red oak, it is masked by two
edaphically different types of populations in the north. The
nursery data do not support the occurence of a genetic break
at the transition zone in any of the three species studied, but
rather suggest a continuous south-horth trend. This suggests
that differences in soil, climate and other environmental con-
ditions on the two sides of the transition zone mold morpholo-
gical characters, especially acorn dimensions differently or
that the seeds represent a broader range of genetic types than
can survive to mature acorn producing individuals.

It is interesting at this point to note that Gillis (1962)

reported a break in the morphology of Rhus Toxidodendron (poison

 

Ivy) in approximately the same area in Michigan as reported here
for black oak and the scarlet-Hill oak complex. He lists two
taxa, a southern and a northern one as varieties of the species.

1The red oak group in Michigan (Erythrobalanus) includes one
other species not discussed in this dissertation, shingle Oak,
(Q. imbricaria. Blackjak oak, Q. marilandica, does not occur
as a native tree in Michigan. The old reports of its occurence
appear erroneous identifications of black oak sprout growth.

 

152
No data on genetic differences was presented.

One of the major problems encountered in this study was
that of intermediate trees. This has been frequently reported 1
in the red oak group, Palmer (1948), Stebbins (1950), Zamierowski
(1960). It has been speculated that there are no effective
genetic barriers between closely related species in the red oak
group, Palmer (1948), Stebbins (1950), Mu11er (1952). In the
study region I have found trees, and in some cases even popula-
tions, that appeared intermediate in their field morphological
characters between the following; red oak - northern red oak,
red oak - black oak, northern red oak - black oak, red oak -
Hill oak, northern red oak — Hill oak, red oak - scarlet oak,
red oak - pin oak, Hill oak - pin Oak, pin oak - black Oak,
Hill oak - black oak, black oak - scarlet oak.

A preliminary study of the pigments of the inner bark in
the red oak group indicates a similar relationship. Brother
Edward Zamierowski (unpublished M.S. thesis, Ohio State Univer-
sity, 1960) gave the following data from a paper chromatography
study of the pigments in a population in Ohio containing black
oak, scarlet oak, and red oak:

Field identification, ’ Identification by paper
number of trees by species chromotographic pattern,
number of trees by species

black oak 26 16
scarlet oak l3 3
red oak 6 15

'2 2 3

153
Ten of the individuals showed intermediate patterns between
species inferred by Zamierowski to be indicative of back-

crossing or hybridization.

Black oak - scarlet oak 5
Black oak - red oak 2
Red oak - scarlet oak 3

Palmer (1948) lists recognized hybrids for the following:

Q. velutina X ellipsoidalis (X Q. palaeolithicola), Q. velutina

 

 

X borealis (X Q. hawkinsii), Q. velutina X palustris (X Q. vaga),

 

 

Q. borealis X coccinea (X Q. Benderi). In this last case Palmer
does not try to differentiate between Q. 53933 and Q. {2233 var.
borealis. (Q. borealis = Q. ggpgg var. borealis in the strict
sense).

In his article on "Hybrid Oaks Of North America" (1948),
Palmer makes the following points:

"It has been stated that most or all of the species within
the two main sections of white oaks and black oaks are
interfertile and that in some cases at least they produce
viable fruit in abundance. There is also evidence that
some of them reproduce themselves with little change,
retaining characters intermediate between the parent
species. Since several species are often found growing
together in the deciduous forests throughout North America,
it may be asked, as Englemann asked, why are hybrids not
more common and why do they not tend to spread in their
native woods and to become dominant in some areas?

"Perhaps there are some physiological reasons that tend
to reduce fertility of some oak hybrids, but evidence is
lacking on this. And without resorting to such an ex—
planation probably a sufficient answer can be found in
the keen competition for survival and reproduction among
plants in nature. In a well occupied forest there is
little chance for expansion and reproduction is held to
a minimum except where the pressure is relieved by some

155

extraordinary chance or vicissitude, such as artificial

clearing or thinning or a similar result brought about

by such natural causes as fire, hurricane, climatic ex-

tremes, or disease among some of the competing species."

It would appear that during the various advances and re
treats of the Wisconsin glacier and on into post Wisconsin time
to the post logging period Michigan and other nearby states be;
came natural laboratories for experimentation among the oaks.
Newly available habitats, site variability, climatic variability,
cutting and fire were all at work sorting out of this grand ex-
periment those forms that were best suited for the changing
habitats. It is from such a background that the present species
variants, forms, and intermediate pOpulations in the Michigan
area were formed.

These inter—actions in the Michigan area as seen today can
perhaps be summarized by a diagram such as Fig. 32. With an
understanding of the possibilities of intermediate trees and
pOpulations in the directions as shown on the diagram, and the
effect this may have on field morphological characters, and also
a recognition of the sorting effect of the environment (including
competition of other species) in selecting the better suited
genotypes, I feel that the variation patterns of the red oak
group as it is encountered in Michigan and nearby states can be
logically understood.

The evidence appears to be mounting for this type of pattern
not only in the red oak group, but also in the white oak group.
There are reports Of intermediate populations between Q. elhg

and Q. montana, Allard (1949), Sillman (1958), and frequent re-

156

Fig. 32 Pathways of suspected gene flow in the red oak group
in Michigan and nearby states. Major pathways are
designated by broad connections.

 

 

 

157

poets of intermediate pOpulations between Q. bicolor and Q.

macorcarpa, Stebbins (1950), Bray (1960). I have noted near

 

Minneapolis, Minnesota, what appear to be intermediate trees

between Q. macrocarpa and Q. alba.

 

Stebbins (1950) describes the genetic structure in Qgercus
in these words:

"...we have a rather large number of species, many of them

polytypic, which are often capable of exchanging genes with

other, morphologically distinct pOpulations occuring in

the same region and in similar habitats. These sympatric

populations have always been kept apart as distinct species,

but some of them, such as Q. bicolor and Q. macrocarpg,
behave genetically more like subspecies."

 

The apparent panmictic character of the red oak group under
appropriate conditions is most clearly evident in populations
that contain trees intermediate between two closely related
species. For example, when the patterns of leaf outline of
all oak samples in the Honor Dunes stand are compared, there
is an obvious morphological continuum between red and black oak
even in the small sample obtained in this study. This was also
found to be true in comparing leaf outlines in other stands
that contained intermediate trees, i.e. Alpena, E. Tawas,
Stockbridge, Wilson State Park, Thornton WOOds, and Rockford.

If the leaves collected from these particular stands were sorted
as a group, they would form an apparent continuum in leaf out-
line between northern red oak and black oak, black oak and Hill
oak, and pin oak and Hill oak. It was possible to detect dif-

ferences between certain populations ole. rubra var. borealis

158

' and Q. Egyge in leaf outline but with connecting forms. How-
ever, in other Q. Egbgg var. borealis populations there were
no basic differences between the species and the variety in
leaf outline.

The areas where morphologically intermediate populations
occur appear to be sites or regions in which environmental
gradients are conspicuous or in which disturbance has upset
normal relationships. For example, several of the stands in-
volved (Honor Dunes, Alpena, Muskegon State Park, Thornton Woods)
occupy an area across which soil and/or drainage undergoes or
has undergone point to point variation. Also, the broad
climatic gradient across Michigan appears to reach critical
conditions for a number of species between Lansing and the
Straits of Makinac. An example is the black oak found in the
northern end of this zone. Here at its range limit, a fringe
of intermediate trees between black oak and red oak (Q. £2253
var. borealis) is found that, as far as I have been able to de-
termine, extends across the northern part of the Lower Peninsula
of Michigan where the two species are found together. Popula-
tions sampled from this fringe include Alpena, Traverse City
State Park, Traverse City Fair Grounds, and Benzie Dunes.

This together with the large pOpulation at Indian River
appearing intermediate between black oak and Hill oak suggest
that even when some environmental conditions on the range limits

prevents the persistence of the full gene complement of the local

159
pOpulation, its niche may not be fully pre—empted by the
"next hardiest" species. As a consequence, hybrids which
embody enough of the traits this "next hardiest" species to
mature, but also incorporate enough of the genes of the "range
limits" species to be more or at least as effective on this
particular type of site as the pure "next hardiest" species
occur in significant numbers on these sites. It might be
thought of as a special case of the "hybridization of habitat"
of Anderson (1948).

The suggested relationships between the members of the red

 

oak group is best illustrated ble. ellipsoidalis. In the broad
sense, it has forms which tend toward scarlet oak, jpin oak,

red oak, northern red oak, and black Oak. It could be pictured
like a wheel with the spokes representing the different species
and Hill oak representing the hub. Hill Oak as a taxon in the
broad sense, in accord with the species description by Rev.

Hill actually symbolizes the nature of the inter-gradation in
the red oak group. Even in the strict sense, as it has been
used in this dissertation, the more narrow acorns of the
northern populations and their less acuminate scales (See figs.
15, 16) suggest introgression with black oak and also probably
red oak. This is most dramatically illustrated in the two basic
kind of buds of the northern populations, those which are heavily
pubescent and those which are essentially glabrous.

To paraphrase Wittaker's (1956) use of Sewell wright’s

...
concept of the "adaptive peak", it appears as if the red oak
group represents a series of peaks in a genetic landscape in
which the peaks represent favorable combindations of genes
with valleys representing less favorable combinations. This
relationship would be dynamic and would change with time
and with changes and trends in environmental conditions. Thus
black oak, red oak, pin.oak and scarlet oak would represent
major peaks with northern red oak (Q. Egbgg var. borealis)
and Hill oak representing either minor peaks on the slopes
between the peaks, portions of the slapes between the peaks,
or broad bajades of locally successful coalesced contributions
from 2 or more of the peaks.

One area that may seem to have been neglected in this
dissertation is the position of pin oak in this whole problem.
.After observing pOpulations of pin oak in Indiana, Illinois,
and Michigan, it appears as if pin oak is the most stable of
all the species of the red oak group under study. It appears
to contribute to the genetic variability of the other species,
especially in the Thornton WOOds population, but tends to occur
in those stands as a relatively stable entity itself. Of two
sources of pin oak I grew in the nursery, one collection from
south-central Indiana and one from an area about 40 miles north
of Lansing, Michigan, significant variability was lacking even
in time of germination. This sample was very small, only two

female parent trees, and can serve only as a tentative point

161
reference. This does not mean pin oak is unimportant in
understanding the red oak group in theMichigan area, and
in particular the Hill oak problem. I strongly suspect that
it is one of the key species in the origin of the scarlet-Hill
oak complex.

Based on the apparent interrelationships between Hill oak,
pin oak, and black oak in the Thornton WOOds population, on
similar relationships in the Elkhart scarlet Oak population,
and also on my observations in northwestern Ohio, I would like
to speculate On a possible origin of the pOpulation first given
the Hill oak taxon.

In all of these areas a pin oak population occupies the
poorly-drained soils of a moisture gradient with black oak
nearby on well-drained soils, and the bulk of the Hill Oak found
in the intermediate part of the gradient. If we speculate that
these populations of Hill oak represent the result of gene ex-
change between black and pin oak, individuals could also occur
at both extremes as a result of the backcrossing. This would
also indicate that individuals of Hill oak could be found through-
out the moisture gradient from dry to moist, depending on the
competition in the stand in which it is found and the particular
genetic complement it contains.

In the Lansing area and in southern Michigan, I have found

that Hill oak in the strict sense is most commonly found on im-

perfectly- to well-drained soil near the edge of swamps or lakes,

162
often forming a narrow band along the zone between poorly-
drained and the well-drained soil. In some of these areas
pin oak is no longer present. However, if this speculation
were correct, the presence of Hill oak in the narrow sense,
would be suggestive either of the former presence of pin oak
or of migration of the intermediate into new areas. How far
this segregate might spread from its point of origin if suit-
able habitat were available is difficult to say. Immigration
of hybrids into old, well established communities might be
highly unlikely, but in the young glacial terraine of the study
region change is characteristic.

Not all of the resultant combinations from gene interchange
have survival value. Only those combinations that are able to
compete with the existing vegetation survive. However, many
of those that do survive appear able to maintain themselves
for generations. This may be due to a linkage pattern in which
certain combinations of physiological relationships and capacities
have survival value while other combination along a genetic
continuum are incapable of growth, are sterile, or unable to
meet the competition with the more successful combinations.
This is in line with the adaptive peaks concept presented
earlier.

The graphs of variance presented with each of the species
discussed in this dissertation appear to support the conjecture

as to the origin of Hill oak . In the study region the scarlet-

163
Hill oak complex as a group definitely has more characters
with a significantly high variance than either black or red
oak, particularly when obvious intermediate populations of
black and red oak are not included. This higher variance
would be expected from a group which had its origin through

hybridization and backcrossing.

Part VI Conclusions

164

165

Conclusions
There is a south-north trend in certain field morpho-
logical characters in the red oak group in the Michigan
area, especially in acorn dimensions.
There is a south-north trend in the red oak group as is
indicated by nursery studies under common garden conditions.
There appears to be a break in the south-north trends in
the field morphological characters near the transition
zone in Michigan in black oak and the scarlet-Hill oak
complex. It was not, however, readily evident in either
red oak (Q. EQQEQ) or northern red oak (Q. Egbgg var.

borealis).

 

There is no evidence of a genetic break among the nursery
populations of black oak or among the nursery p0pu1ations
in the scarlet—Hill oak complex in the Michigan area but
of south-north genetic trends.

Northern small acorn pOpulations of the red oak group in
Michigan with yellow inner bark are basically more closely
related to black oak than Hill oak though they may be in-
termediate in nature between the two or between black oak

and northern red oak (Q. rubra var. borealis).

 

There is evidence that it may be possible to recognize a
northern small-fruited race of black oak comparable to the

northern race in red oak (Q. rubra var. borealis).

10.

11.

166
Acorn size alone cannot be used as the basis for separating
species in the red oak group in the Michigan area but may
be useful in separating races.
Hill oak and scarlet oak are considered part of a complex
and not as separate species in the Michigan area. The

taxon Quercus ellipsoidalis being regarded as a small-

 

fruited northern race of Quercus coccinea.

 

Northern red oak (Q. rubra var. borealis) in Michigan is

 

thought to be the result of introgression of red oak (Q.
ggbgg) with either black oak or Hill oak or both and as
a result is restricted to disturbed or poor sites where
it can compete with them and with red oak.

The field and nursery evidence concerning the red oak
group in Michigan supports the view that the group con-
sists of a series of species which may freely intergrade
in all directions among themselves when favorable site
conditions are available for their survival. However,
there appear to be major and minor pathways for this
intergradation or gene exchange.

A conjecture is made that the type locality Hill oak
p0pu1ation and other populations under similar conditions
of a changing watertable and soil conditions may have

originated from a black oak - pin oak gene interchange.

167
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Stebbins, G.L., Jr., E.G. Matzke, and C. Epling. 1947.
Hybridization in population of 99ercus marilandica
and Quercus ilicifolia. Evolution. 1: 79-88.

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171
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172

Appendix A

Species basal area in square feet / acre by stands
in which b1ack_oak p0pu1ations were found.
area obtained by Bitterlich method, Grosenbaugh
(1952) with 10 sample points.

Species

Quercus velutina
Quercus alba
Fagus grandifolia
Liriodendron tulipifera
Acer saccharum
Quercus rubra

(and var. borealis)
Carya ovata
Prunus scrutina.
Carya glabra
Cornus florida
Sassafras albidum
Tilia americana
Fraxinus americana
Populus grandidentata
Quercus bicolor
Ostrya Virginiana
.Acer rubrum
Quercus ellipsoidalis
Ulmus americana
Anelanchier sp.
Pinus strobus
Pinus banksiana
Pinus resinosa

‘Population basal area

is ..
an: x
HH H
0 Cd
0. 0 Q4
01.3 'U
~ga c! a)
.x H u
09-: ‘ :3
a. u m
MU $4
om cd to
ou .1: a)
.gcn £3 a
a m a
58 32 145
30 , 33 2
15 1
15
10 1
6
2
13 2
7
9
1 1
137 88 158

Ind.

Pokagon State Park,
Ind.

NW
ub-

'ha

P‘H‘ to
c:

DDCDNHl-‘Hw

L0

125

Rose Lake Wildlife

b .
H

25
17

89

Exp. Sta., Mich.

Basal

‘5“,
figs
44 d)
23.3
«344-3
Com
04-)
DOCUCO
.3588
COGS-£1
53:2
42 24 39*
9 5 5
39*

9 5

57

1
60 92 83

‘*This Hill oak-black oak p0pu1ation was grouped together for
gathering data in the field and the total divided in half

for each species.

er State Park,

R.

Slee
Mic

w
H0

15

73

173
Appendix A

Species basal area in square feet / acre by stands
in which black oak populations were found. Basal
area obtained by Bitterlich method, Grosenbaugh
(1952) with 10 sample points.

E. Tawas, Mich.

Traverse City

State Park, Mich.
Grounds, Mich.
Indian River, Mich.

Honor, Mich.
- Traverse City Fair

Species

«b
(A)
N
\O

Quercus velutina 27

Quercus alba .

Fagus grandiofolia

Liriodendron tulipifera

Acer saccharum

Quercus rubra ' .5
(and var. borealis)

Carya ovata

Prunus serotina

Carya.glabra

Cornus florida

Sassafras albidum

Tilia.americana

Fraxinus americana

POpulus grandidentata

Quercus bicolor

Ostrya.virginiana

Acer rubrum

Quercus ellipsoidalis

Ulmus americana

Amelanchier sp.

Pinus strobuS' '1 1 29 25

Pinus banksiana 42 15 2 - 5

Pinus resinosa 1 10 10

w

1“
MH
I—‘
b
N
0

Population basal area 47 28 82 98 34

Appendix A

174

Species basal area in square feet / acre by stands

in which scarlet-Hill oak populations were found.
Basal area obtained by Bitterlich method, Grosen-

baugh (1952) with 10 sample points.

Species

Quercus velutina
Quercus alba

Prunus serotina
Carya glabra
Sassafras albidum
Tilia americana
Fraxinus americana
Populus grandidentata
Quercus bicolor

Acer rubrum

Quercus ellipsoidalis
Ulmus americana
Amelanchier sp.
Quercus prinus
Quercus coccinea
Nyssa sylvativa
Quercus palustris
Quercus macrocarpa
Populus deltoides
Populus tremuloides
Fraxinus pennsylvania
Pinus strobus

Pinus banksiana

Pinus resinosa

Population basal area

Brown County St.
Pk., Ind.

box-

[—1
OLA)

38
20

89

34

13

Elkhart, Ind.

28

22
25

Thornton WOods, Ill.
. Rockford, 111.

O\
\OU'IO\

(AM

PM“

Stockbridge, Mich.

35

16

33

Rose Lake Wildlife
Ex. Sta., Mich.
Farwell, Mich.

.b-
...:

H

25 -
17 62

. d
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5 8
39* 47 41
1 1
34 46

94 100 117 110 89 65 83 82 96

*This Hill oak-black oak p0pu1ation was grouped together for
gathering data in the field and the total divided in half for

each species.

175

Appendix A

Species basal area in square feet / acre by stands
in which red oak populations were found.
obtained by Bitterlich method, Grosenbaugh (1952)

with 10 sample points.

. Pokagon St. Pk., Ind.

Species

Quercus velutina 34
Quercus a1ba 23
Fagus gradifolia
Liriodendren tulipifera 1
Acer saccharum
Quercus rubra 20
(and var. borealis)
Prunus ”retina,
Carya ovata
Carya glabra
Sassafras albidmm
Tilia americana
Fr axinus americana
Populus grandidentata
Ostrya virginiana
»-Nyssa:sy1vatica
Ulmus americana
Carya cordiformis
Cornus alternifolia
Hamamelis virginiana
Ulmus fulva
Quercus muehlenbergii
Celtis occidentalis
Juglans cinerea
Quercus macrocarpa
Populus tremuloides
Cornus florida 1
Pinus strobus
Pinus banksiana

PM“
oabnbcnrorduahl

Warren Dunes State
Park, Mich.

11
35

bOON-bl-‘H

10
10

E. Lansing, Mich.

Muskegon St. Pk., Mich.

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7 .5
35 .5
2
6

Basal area

Burt Lake St. Pk., Mich.

+4
u:

60

20

32 20 l 10

3 15

3

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23 1
33
2 15
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36

Cass Lake, Minn.

16

21

176
Appendix A
Species basal area in square feet / acre by stands

in which red oak populations were found. Basal area
obtained by Bitterlich method, Grosenbaugh (1952)

with 10 sample points. . .d
, .5 a .3
'o 44 .é . g; :3
2101: 0A .
H8 ..déjr—Sq; 45,
238,8125599 ...
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Q4 2 Phi/NO.“ “a
.35.;Q43agm a“
swamp-g “345-02
£352 *1 C U) .3 Fl l2 :4 g 6‘
C we vmm 22:11.3
0C1~C100<D r-I‘UIH “3
00 w.¥ to m ~4 L: r4 m . p ,4
m u a ,1 a: N o 44 cm L: o: a)
£03m0w58.’58“%&$
. e. :3 k1 :§ :2 z: a: c: :3 a: o w
Spec1es c5 c3
Pinus resinosa 11 6 15 1 4 31
Tsuga canadensis 1 20
Betula papyrifera 3
Abies balsamea 13 28
Betula Lutea 2
Thuja occidentalis 11
10

Picea glabra
Population basal area 125 108 133 98 84 28 122 64 89 51 101 114

177

Appendix B

Alist of the morphological characters in the red oak group

studied during the summer of 1960 at the University of Michigan

Biological Station at Pellston, Michigan.

waH
O O O

\DOO\IO\U1
00.00

10.
ll.
12.
l3.
14.
15.
16.
17.
18.
19.
20.
21.
22.
23.
24.
25.
26.
27.
28.
29.
30.
31.

Leaf b1 ade length

Leaf petiole length

Leaf blade width

Average leaf body width (the width of the unlobed portiOn
of the leaf)

Distance of the widest part of the leaf from the leaf tip

Leaf pubescence - upper surface

Leaf pubescence - lower surface

Leaf surface (whether shiny or dull) - upper surface

Leaf surface - lower surface

Leaf color - upper surface

Leaf color - lower surface

Leaf vein axil pubescence - lower surface

Leaf shape (whether obovate, etc.)

Twig pubescence

Length of acorn with cup attached

Acorn cup height

Acorn cup width

Acorn length

Acorn width

Acorn cup shape

Shape of acorn cup scale

Shape of tip of acorn cup scale

Presence of humping on acorn cup scale

Acorn cup scale color

Presence of fringe on lip of acorn cup

Tree self-pruning

Twig brittleness

Tree diameter at 4% feet from ground
Inner bark color

Bark texture at 2 feet from ground

Bark.texture at 7 feet from ground

The following characters were calculated from the above measure-

ments:

32.
33.
34.

35.

Leaf shape (width / length)

Leaf area (length x width)

Ratio of blade length to petiole length
Acorn shape (width / length)

178
Appendix B
36. Acorn size (length plus width)
37. Ratio of cup height to height of acorn with cup
attached
Notes were also made on the following population characteristics;
presence of pitting at the apex of the acorn, leaf petiole color,

presence of striping on acorns, basal area of stand, soil type,

and legal location.

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182

Appendix D

Tests by analysis of variance to indicate which groups of black
oak field populations showed a significant difference at the 5%
and 1% level. These tests were used to indicate homogeneity

among populations that did not show this significant difference

and as the basis
These tests were

character.

Field Character

Total Acorn
Length with
Cup Attached

.Acorn Length

.Acorn Width
Acorn Shape

.Acorn Size

.Acorn Cup
Height

IRatio of Total
.Acorn to Cup
Height

Acorn Cup Fringe

Presence

.Acorn Cup Scale
Shape

'Pwig Pubescence

for tests between p0pu1ations two at a time.
carried out for each morphological field

a? .a
'U'U .CIU
- Cc: OH
H HHO) ~42:
U3 U4 2
«mg-lo o a
o cor-[£030.00 0
$4 ..L’F‘fi'UUonxfi-S
o ammu-I-r-Io-H.xrloo.£:o
“U HZHED-t HU-r-l
‘0 C m - E: 2: 0 2: -H 2:
"' HUU a 0".“ z
r3 (Scam-«puma a
U «u .xmcmm man—1
22mgsasacmgHH
h-cfimbo mBOQmHH
O'UfimmmvxummE-«oo
UC.‘ GACVJNUJGu—lm CC!
UHHgoohJDQJHr—l-OO
'21 m mm zmzmmmn:
X XXXXX XXX
X XX
X XXXXX XXX
X XX
X XXX XX
X XX
X XX XXX X
XXX
X XXXXX XXX
X XXX XXX
X XX
X XXXXX
X XXX
X XXXXX
XXXXXX
X XXXX
XXXX
X XX XX
X XXXX X X
X XXXX XX X X
XXX X
X X X
XXXXX X
XXXXX
X X

183

.30“: .HH pocom

.soHZ .H nocom

.SUHE .mmame .m

.nofiz ..xm .um pmammfim
.nofiz ..xm .um cowHHB
.50“: .uwuwaucmm

.non .commxmsz

.50“: ..mum .xm
mMHHnHfiz mxmg mmom

.ncH ..xm .um aommxom
.eaH ..xm mumum magma
.naH .uumnxflm

.vcH
..xm .um .uo m_xoflenouoz

Field
Character

X

Inner bark

color

X

Tree Form

Twig Brittle-

HESS

.5
>50
“ 32:“
8
Field $.85
Character g .
w.u
L! U)
5*
Total Acorn
Length with x
Cup Att.
Acorn Length
x

Acorn Width
Acorn Shape x

Acorn Size

Acorn Cup
Height

Ratio of Total
Acorn to Cup
Height

Acorn Cup Fringe
Presence

Acorn Cup Scale X
Shape x

Twig Pubesc. x
x

Inner Bark Color

xxNXNN

Traverse City

Fair Grounds, Mich.

N

184

Alpena, Mich.

N

N

N

Indian River, Mich.

X

x

XXXXN

F value, analysis of

level
** = Significant at 1%

= Significant at 5%

variance

7':

1.83
1.02

2.29*
1.48
1.97
2.69*
0.91
0.98
7.36**
2.23*

no sig.

2.32
2.46
2.01
1.32
4.09**
12.83**
2.94
2.34
1.42
24.84**
0.06
24.91**
1.26
4.38**
1.15
2.32
0.06
21.92**
1.07
1.43
1.81

level

dif.

185

Hm>oH
NH um quUHMHcmHm u ¥«
Hm>mH
Nm um ucmMHchmHm n s
. mocmwum>
mo mamhamcm .mDHm> m

.noaz .um>Hm amaecH

.suazv.wcmaw<

wean: .meqsonu yawn
huau omhm>mue

.Lofiz...xm .um
huwo mwhm>muH

Field
Character

1.20

Tree Form

5.23**
0.22
1.78
2.37*
1.25

Twig Brittleness

186

Appendix D

Tests by analysis of variance to indicate which groups of the
scarlet-Hill oak complex field populations showed a significant
difference at the 5% and 1% level.
indicate homogeneity among populations that did not show this
significant difference and as the basis for tests between pop-

ulations two at a time.

morphological field character.

.d

U)

>.

4.)

c: .
Field 8'2
Character 59**

El»:

84m

m
Total Acorn, x

length with
cup attached
Acorn Length x

Acorn Width x

Acorn Shape x
x
Acorn Size x

Acorn Cup Height

Ratio of Total
Acorn to Cup
Height

Acorn Cup Frings
Presence

Acorn Cup Scale
Shape

X

>4

X

Elkhart, Ind.
Thornton woods,

>4

Chicago, Ill.

Rockford, Ill.

>4

Stockbridge, Mich.

These tests were used to

Rose Lake Wildlife
Ex. Sta., Mich.

>4

Wilson State Park, Mich.

>4

Farwell, Mich.

Wisconsin Rapids, Wisc.

>4

Graylino, Mich.

F value, analysis of

variance

These tests were carried out for each

level
** = Significant at 1%

= Significant at 5%

‘k

2.19

level

not sig.

1.05
2.13
5.12**
1.71
5.08**
4.84**
3.43**
2.08
1.60
4.01**
2.08

10.79**

8.46**
0.18
2.83
1.48
3.5788

2.61*
1.74
2.29
5.92**
1.21
2.81**

187

Hm>mH

NH um unmeamacmam n 88
Hm>mH

gm um unmoHMHcmHm u «

mocmauw>
mo mammfimcw .msam> m

.SOHE .wEHH%MHU
.omflz..wpadmm Camcoomfiz

.5032 .Hamzumm
.son ..xm mumum comaflz

.sofiz ..mum .xm
mmafieaaz_mxmg mmom

.8032 .mmaapnxooum
.HHH .8uomxoom

.HHH .OwaHSU
.mpooz couahonfi

.vGH .uumnxam

.vcH ..xm
.um hucfioo fiBoym

r
e
t
C
Mm
e a
ih
FC

x 3.01

Twig. pubsc .

9.17**
0.43
1.90

X

Inner Bark

Color

2.92**
0.37
2.09

x 28.41**

X

X

Tree Form

X

3.96

X
X

X

Twig Brittle-

HESS

2.69

188
Appendix D

Tests by analysis of variance to indicate which groups of
red oak field p0pu1ations showed a significant difference
at the 5% and 1% level. These tests were used to indicate
homogeneity among p0pu1ations that did not show this sig-
nificant difference and as the basis for tests between pop-
ulations two at a time. These tests were carried out for

each morphological field character. . .é
.c o
o O U ‘H
'U H .1': A '21
C“ E U CD a
H H (D e .C
o a 2 fl 0 U
r .5 f3 .8. . a a 8
.54 U) w-l 0.. . v
p. z ,3 . . ..
m - D4 0 I: «U CD
0 Q) 0 A U ..C U U) x
5‘ as no U) . o :1 CU
Field Character (3.3 E. g 30’ E A g ’4
a 2: m o . m m
a) c v c: co m . m t: w
00 m,« :6 a) *1 p 5 r4
:8 t . h: z: N c: 0 4J no
.24 .24 “1 G :1 Ch 3 :3
O (GD-a 0‘ :3 (D O r-l 0.-
9-! 3 m E m 3 <21 an Q
Total Acorn, x x x x
length with cup x x x x
attached x x x x x
Acorn Length . x x x x
x x x x
Acorn Width x x x
x x x
x x x
Acorn Shape x x x x x
x
x x x x
x x x x x x
x x x x x x
x x x
Acorn Size x x x x x
x x x
x x
x x x x
x x x
Acorn Cup x x x x x
Height x x x x x x x
x
Ratio of Total x x x x x x x x
Acorn to Cup x x x x x x x x

Height

Dunbar Forest Sta., Mich.

Raco, Mich.

>4

>4>4>4

189

.gofiz .oumm

.nUHz ..mum ammuoa panama

.soazm.mxmu mmflwsom
.53: .53 .um. 33 “:3
.nofi2_.mcomw¢

Ammcnmv .non .uocom
.son ..xm.um mnncmm
.non ..xm .um commxmpz
.5032 .mcfimcmg .m

.soaz ..81
.um mogsa aohumz

.ucH ..xm .um ammmxom

Field
Character

X X

X X
X X
XXXX
X X
X

X X

X XXX

X XXX

X XXX

X XXX

8

g e

n .1

.1 a

r C

F S

pep

ucu

CnC

mm... m

rema

OHOh

C CS

A A

Twig. Pubsc.

X

Inner Bark Color x

XXX

Tree Form

X
X

X

Twig Brittleness x

XXX
XXX

190

Field Character

Total Acorn
length with cup
attached

Acorn Length

Acorn Width

Acorn Shape

Acorn Size

Acorn Cup
Height

Ratio of Total
Acorn to Cup
Height

Acorn Cup Fringe
Presence

Acorn Cup Scale
Shape

Twig Pubescence

Inner Bark Color

>4>4>4

>4

COpper Harbor, Mich.

Cass Lake, Minn.

F value, analysis of

variance

level
** = Significant at 1%

= Significant at 5%

7‘:

4.35**
1.13
2.92*
2.80
1.08

,3.08

3.21*
3.51**
1.46
0.16
0.43
1.86
2.24*
0.94

14.95**

5.45*
0.53
9.16**
4.25*
0.39
2.88**
no.-sig.
1.74
2.87**
7.97*
2.75*
2.92*
0.51
2.43*
2.69*
2.55
1.51

14.38**

5.1288
1.69
5.4488

level

191

Hm>mH
NH um ucmoamacwam n «4
Hm>mH
gm um uchHMHcmHm n 8
mocmfium>
mo mfimzamcm .mDHw> m

.CCHE .mxmq mmwo

.soHE .nonumm ummdoo

Field Character

Tree Form

* * **
*** ***
62671588
14167846
00......
63512340
X

XX

S

S

e

n

e

1

t

t

.1

r

B

00

.1

W

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