The Edge Effect of the Lesser Vegetation of Certain
Adirondack Forest Types with Particular
Reference to Deer and Grouse

By

F. B. BARICK

ROOSEVELT WILDLIFE BULLETIN
Volume 9 August, 1950 Number 1

Roosevelt Wildlife Forest Experiment Station

New York State College of Forestry
at Syracuse University, Syracuse, N. Y.

Joseph S. Illick, Dean

The Edge Effect of the Lesser Vegetation of Certain
Adirondack Forest Types with Particular
Reference to Deer and Grouse

By

F. B. BARICK

ROOSEVELT WILDLIFE BULLETIN

Volume 9 August, 1950 Number 1

VOLUME XXIII NUMBER 4

Published quarterly by the New York State College of Forestry at Syracuse, N. Y.
Entered as second-class matter, October 18, 1927, at the Post Office at
Syracuse, N. Y., under Act of August 24, 1912.

ANNOUNCEMENT

The serial publications of the Roosevelt Wildlife Forest Experi-
ment Station consist of the following :

1. Roosevelt Wildlife Bulletin

2. Service Publication

The Bulletin includes papers of a technical or semi-technical nature
dealing with various phases of forest wildlife, its management and
conservation. The Service Publication is intended to be of general and
popular interest and attempts to interpret forest wildlife research
results and explain the reasons for and methods of their application.

The editions of the Bulletin are limited and do not permit of general
free distribution. Prices of the individual numbers vary as they are
based on actual cost of printing and distribution in accordance with
Chapter 220 of the Laws of 1933. Price lists will be furnished on
request. Exchanges are invited. All communications concerning
publications should be addressed to :

THE DIRECTOR
Roosevelt Wildlife Forest Experiment Station
Syracuse, New York

Notice: This issue is the fifth number of a new series of the Bulletin.
Each issue hereafter will contain only one article and the number of
issues per volume will be determined by the size of the individual
numbers.

TRUSTEES OF THE NEW YORK STATE
COLLEGE OF FORESTRY

EX OFFICIO

Joe R. Hanley, Lieutenant-Governor Albany, N. Y.

W illiam P. Tolley. Chancellor, Syracuse University Syracuse, N. Y.

Lewis A. Wilson, Acting Commissioner of Education.... Albany, N. Y.

Perry B. Duryea, Conservation Commissioner Albany-, N. Y.

APPOIXTED BY THE GOVERNOR

J. Henry Walters, President New York, N. Y.

Francis D. McCotn, Vice-President Syracuse, N. Y.

Charles D. Upson Lockport, N. Y.

George R. Fearon Syracuse, N. Y.

Grant \\'. Ernst Syracuse, N. Y.

Orville H. Greene Syracuse, N. Y.

Frank C. Ash Fulton, N. Y.

James P. Lewis Beaver Falls,

N. Y.

William B. Stark Syracuse, N. Y.

STAFF OF THE ROOSEVELT WILDLIFE FOREST
EXPERIMENT STATION

Joseph S. Illick. D. Sc Dean of the College

Hardy L. Shirley, Ph.D Assistant Dean of the College

Ralph T. King, M.A Director

W ilford a. Dence, B.S Ichthyologist and Assistant Professor

William L. Webb, M.S Assistant Professor

Marianna Neighbour Secretary

[1 ]

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LIST OF ILLUSTRATIONS

PAGE

Fig. 1. The transect employed was a strip of plots extended far
enough in each direction to include typical conditions for each

of the adjacent forest types 31

Fig. 2. A sample plot of young trees, shrubs, and herbaceous vegeta-
tion. The corners of the underbrush plot have been marked
with stakes and the herb plot is marked with sticks laid in-
side the larger plot. On this plot underbrush (shrubs) oc-
curred with a density of Three 31

Fig. 3. Density values. A species of underbrush covering less than
four square feet was ascribed a density of Trace; if it
covered from four square feet to one-third of the plot, it had
a density of One ; if it covered from one-third to two-thirds,
it had a density of Two,- and if it covered from two-thirds to
three-thirds, it had a density of Three 33

Fig. 4. "Reading" the underbrush plot. The density of the shrubs
or herbs could be estimated from one side, but small tree
seedlings required counting 33

Fig. 5. The tree data from each transect were transcribed from the

field sheets to Graphical Analysis Form 1 36

Fig. 6. The herb data from each transect were transcribed from the

field sheets to Graphical Analysis Form 2 36

Fig. 7. Graphical analysis of the distribution of shrubs and herbs

across the edge between Bracken Fern and Aspen 46-47

Fig. 8. Graphical analysis of the distribution of shrubs and herbs

across the edge between Bracken Fern and Second Growth . . 52-53

Fig. 9. Graphical analysis of the distribution of shrubs and herbs
across the edge between Second Growth and Northern Hard-
wood 60-61

Fig. 10. Graphical analysis of the distribution of shrubs and herbs

across the edge between Spruce Flat and Mixed Hardwood. . 68

Fig. 11. Graphical analysis of the distribution of shrubs and herbs

across the edge between Spruce Flat and Northern Hardwood 75-77

Fig. 12. Graphical analysis of the distribution of shrubs and herbs
across the edge between Mixed Hardwood and Northern
Hardwood 85-86

Fig. 13. Graphical analysis of the distribution of shrubs and herbs

across the edge between Spruce Flat and Spruce Swamp.... 93-95

Fig. 14. Graphical analysis of the distribution of shrubs and herbs

across the edge between Spruce Flat and Opening 103-104

Fig. 15. Graphical analysis of the distribution of shrubs and herbs

across the edge between Northern Hardwood and Opening.. 111-113

Fig. 16. Graphical analysis of the distribution of shrubs and herbs

across the edge between Spruce Flat and Open Swamp 115-116

Fig. 17. Comparative value of the vegetation at the various edges

for ruffed grouse, as based on general major requirements.. 124

Fig. 18. Distribution of grouse and deer in relation to the edges be-
tween four of the major types' on the Check Area 133

Fig. 19. Grouse distribution on the Check Area in relation to seasons

and forest types 135

Fig. 20. Deer distribution on the Check Area in relation to seasons

and forest types 136

[3]

LIST OF TABLES

PAGE

Table 1. Forest T ypes on the Check Area of the Huntington Forest.. 23

Table 2. List of transects established Facing 37

Table 3. Derivation of factors for the conversion of graphical data to

a matiiematical basis 39

Table 4. Mathematical analysis of the shrubs and herbs across the

edge between Spruce Flat and Northern Hardwood Facing 41

Table 5. Xunierical equivalents of general abundance groups used in
tables summarizing the principal vegetative features of value
to grouse and deer 43

Table 6. Summary of principal features ad\-antageous to deer and
grouse provided by the lesser vegetation of the edge between
Bracken Fern and .Asi>en 4S

Table 7. Mathematical analysis of the arborescent reproduction across
the edge between Bracken Fern and Aspen expressed in num-
bers per acre 4Q

Table S. Summary of the mathematical analysis of tlie shrubs and

herbs across the edge between Bracken Feni and Aspen 50

Table 9. Summary of the principal features advantageous to deer and
grouse pro\-ided by the lesser vegetation of the edge between
Bracken Fern and Second Growth 54

Table 10. Mathematical analysis of the arborescent reproduction across
the edge between Bracken Fern and Second Growth expressed
in numbers per acre 55-56

Table 11. Summary of the mathematical analysis of the shrubs and
herbs across the edge between Bracken Fern and Second
Growth

Table 12. Summary of principal features advantageous to deer and
grouse provided by the lesser vegetation of the edge between
Second Growth and Northern Hardwood

Table 13. Mathematical analysis of the arborescent reproduction across
the edge between Second Growth and Mature Northern Hard-

wood expressed in numl>ers i<er acre 62-63

Table 14. Summary of the mathematical analysis of the shrubs and
herbs across the edge between Second Growth and Mature
Nonhern Hardwood <>4

Table 15. Summary of princip;il features advantageous to deer and
grouse proxided by the lesser vegetation of the edge between
Spruce Flat and Mixed Hardwood 66

Table 16. Mathematical analysis of the arborescent reproduction across
the edge between Spruce Flat and Mixed Hardwood expressed
in numbers per acre 69-70

Table 17. Summary of the mathematical analysis of the shnibs and
iierbs across tlie edge between Spruce Flat and Mixed Hard-
\\ ood '1

Table IS. Summary of principal features advantageous to deer and
grouse provided by the lesser \egetation of the edge between
Spruce Flat and Northern Hardwood 73

Table 19. Mathematical analysis of the aborescent reproduction across
the edge between Spruce Flat and Northern Hardwood ex-
pressed in numbers per acre 79-80

[-1]

LIST OF TABLES

PAGE

Table 20. Summary of the mathematical analysis of the shrubs and
herbs across the edge between Spruce Flat and Northern
Hardwood 81

Table 21. Summary of principal features advantageous to deer and
grouse provided by the lesser vegetation of the edge between
Mixed Hardwood and Northern Hardwood 83

Table 22. Mathematical analysis of the arborescent reproduction across
the edge between Mixed Hardwood and Northern Hardwood
expressed in numbers per acre 87-88

Table 23. Summary of the mathematical analysis of the shrubs and
herbs across the edge between Mixed Hardwood and Northern
Hardwood 89

Table 24. Summary of principal features advantageous to deer and
grouse provided by the lesser vegetation of the edge between
Spruce Flat and Spruce Swamp 91

Table 25. Mathematical analysis of the arborescent reproduction across
the edge between Spruce Flat and Spruce Swamp expressed
in numbers per acre 96

Table 26. Summary of the mathematical analysis of the shrubs and

herbs across the edge between Spruce Flat and Spruce Swamp 97

Table 27. Summary of principal features advantageous to deer and
grouse provided by the lesser vegetation of the edge between
Spruce Flat and Opening 99

Table 28. Mathematical analysis of the arborescent reproduction across
the edge between Spruce Flat and Opening expressed in num-
bers per acre 101

Table 29. Summary of the mathematical analysis of the shrubs and

herbs across the edge between Spruce Flat and Opening 102

Table 30. Summary of principal features advantageous to deer and
grouse provided by the lesser vegetation of the edge between
Northern Hardwood and Opening 106

Table 31. Mathematical analysis of the arborescent reproduction across
the edge between Northern Hardwood and C)pening expressed
in numbers per acre 107

Table 32. Summary of the mathematical analysis of the shrubs and
herbs across the edge between Northern Hardwood and Open-
ing 108

Table 33. Summary of principal features advantageous to deer and
grouse provided by the lesser vegetation of the edge between
Spruce Flat and Open Swamp 109

Table 34. Mathematical analysis of the arborescent reproduction across
the edge between Spruce Flat and Open Swamp expressed
in numbers per acre 117

Table 35. Summarj- of the mathematical analysis of the shrubs and

herbs across the edge between Spruce Flat and Open Swamp. . 118

Table 36. Summary of the physical and vegetative characteristics of the

ten edges 120

Table 37. Computation of threshold of edge preference for major types

on the check area 131

[5]

PREFACE

Leopold's statement that wildlife "is a phenomenon of edges"
has become a basic principle of modern wildlife manage.Tient. This
principle implies that a habitat composed of a variety of vegetative
types is more productive of both kinds and total numbers of animal
organisms than are habitats lacking such variety. Most of the wild-
life management being practiced today, however, is in connection
with farm game habitats. Only very recently has there been a trend
toward developing methods of forest habitat improvement. But the
state of our knowledge concerning the habits and requirements of
forest species is decidedly limited, and our knowledge of forest
habitats is even less.

This study, therefore, is an attempt to analyze the wildlife sup-
porting potentialities of forest vegetation. Keeping in mind that
edges have proved to be one of the key factors in the management of
farm game, this study was so organized as to analyze the compara-
tive habitat values of certain forest types and the edges between
them. Although some basic data are presented as well as the methods
of analysis, the most that can be expected from this study is that it
serve as a starting point for further investigation.

ACKNOWLEDGMENTS

The author wishes to express his appreciation to Professor
Ralph T. King, Director of the Roosevelt Wildlife Forest Experi-
ment Station and Head of the Department of Forest Zoology at the
New York State College of Forestry for suggesting and making pos-
sible the development of this study as well as allowing reference to
wildlife census data gathered by the Roosevelt Station staff. The
author also wishes to express his appreciation to Dr. Frank E. Egler,
formerly Assistant Professor, Department of Forest Botany, for
consultation and assistance in securing the proper perspective of the
intricacies of forest ecology. To Professor Wilford A. Dence.
Assistant Professor of Forest Zoology, credit and thanks are given
for checking the manuscript. The author also is indebted to Profes-
sor C. C. Carpenter, of Syracuse University, who was kind enough
to check the statistical analysis of the data.

[6]

The Edge Effect of the Lesser Vegetation of Certain
Adirondack Forest Types with Particular
Reference to Deer and Grouse

By

F. B. Barick
Roosevelt Wildlife Forest Experiment Station
CONTENTS

PAGE

rreface

Acknowledgments 5

I. Introduction g

A. Statement of the Problem 8

B. Description of the Site of the Investigation 10

1. The Huntington Forest — General 10

2. The Check Area 10

3. History , \ 11

4. Geology 12

5. Soils 13

6. Climate 14

II. Review of the Literature 15

A. Concept 15

B. Vegetation, the Viewpoint of the Ecologist 15

1. The Organismal Concept of Vegetation 15

2. The Principle of Vegetation Development 16

3. The Factors Controlling the Distribution of Vege-

tation 16

4. Edge Vegetation 18

C. Forest Types, the Viewpoint of the Forester 20

1. The Basis of Classification 20

2. Forest Types of the Adirondacks 22

3. Forest Types of the Huntington Forest 23

D. Vegetation and Wildlife Distribution, the Viewpoint of

the Wildlife Manager 24

1. The Controlling Influence of Vegetation 24

2. Edge Effect 27

E. Wildlife and Forestry 29

III. Collection of Data 30

A. The Transect 30

B. Arrangement of Plots and Equipment Used 30

C. The Tree Plot 32

D. The Underbrush Plot 32

E. The Herb Plot 34

F. Location of the Transects 35

IV. Analysis of Data — Methods 35

A. Preliminary Tabulation 35

B. Graphical and Mathematical Analysis of Underbrush and

Herbs 38

C. Mathematical Analysis of ./Krborescent Reproduction 40

V. Analysis of Data — Edge Composition with Special Reference to

Grouse and Deer 41

A. The Edge Between Bracken Fern and Aspen 42

1. Vegetative Characteristics 43

2. Wildlife Values 44

B. The Edge Between Bracken Fern and Second Growth

Hardwood 45

1. Vegetative Characteristics 45

2. Wildlife Values 47

[7]

8

Roosevelt Wildlife Bulletin

C. The Edge Between Second Growth Hardwood and Mature

Northern Hardwood 51

1 . Vegetative Characteristics 51

2. Wildlife \'alues 58

D. The Edge Between Spruce Flat and Mixed Hardwood 61

1. Vegetative Characteristics 61

2. Wildlife Values 65

E. The Edge Between Spruce Flat and Northern Hardwood.. 67

1. Vegetative Characteristics 7?

2. Wildlife \'alues \\ 74

F. Tlie Edge Between Mixed Hardwf.od and Northern Hard-

wood 78

1. \'egetative Characteristics 78

2. Wildlife Values 84

G. The Edge Between Spruce Flat and Spruce Swamp 86

1. Vegetative Characteristics 90

2. Wildhfe \ alues 92

H. The Edge Between Si)ruce Flat and Opening 98

1. \'egetative Characteristics 98

2. Wildlife Values 100

I. The Edge Between Northern Hardwood and Opening.... 100

1. Vegetative Characteristics 105

2. Wildlife X'alues 110

J. The Edge Betwen Spruce Flat and Open Swamp 113

1. \'egetative Characteristics 113

2. Wildlife Values 114

VI. Comparative X'alues of the \'arious Edges 119

A. Comparison of the \'egetative Characteristics 119

B. Delimitation of Concepts Relative to Edge Effect 127

C. Computation of Tlirebhold of Edge Preference 130

D. Wildlife Distribution in Relation to Edges 134

VII. Summary 140

VIII. Bibliography 142

IX. Appendix:

List of Plant Species 145

Type Map of the Check Area

I. INTRODUCTION

A. Statement of the Problem.

As pointed out in the Preface, edge vegetation is considered by
the wildlife manager as the most favorable site for wildlife. Leopold
('39) describes game as a phenomenon of edges. He states that
species of low mobility and high type requirements such as quail,
pheasants, grouse, and rabbits occur where they have simultaneous
access to more than one environmental type. It is assumed that an
edge is composed of a greater variety of plant species than would be
found in the center of a single plant association. Theoretically, the
edge furnishes a greater abundance of food and cover than does the
center of a type.

Mttch of the wildlife management now being practiced is based
upon this concept of edge eflfect. Most of it. however, is confined to
farm game and comparatively little is being done, at least in the

Edge Effect of the Lesser Vegetation

9

Northeast, in the way of managing habitats of forest and range game
species. It has been assumed that the theory of edge effect appHes
in the forest environment as weH as in agricuhural lands, the general
supposition being tliat there is a greater abundance of game species
at the edge between forest and open land than in the forest alone or
on the open land alone.

The investigation of forest wildlife habitats to some extent has
been neglected. Unbroken forest areas are often considered as "bio-
logical deserts," consequently the potentialities of forest wildlife
management as a worthwhile effort have been more or less disre-
garded on such areas. As a matter of fact, about the only forest
wildlife management that has been practiced is the opening of dense
stands of timber so tiiat browse would be made available. It has been
generally overlooked that forests are composed of a variety of differ-
ent types and that these types in turn differ in their species compo-
sition, and that with a certain amount of manipulation these might
possibly be greatly improved in wildlife value.

Specifically, the purpose of this study is to determine, in so far
as possible, the vegetative characteristics of a forest edge, and to com-
pare these features as they occur in the various edges and types.
Some of the questions this study will attempt to answer are : What
is the characteristic composition of the edge in respect to specific
amounts of the constituent species? Is there a special edge flora or
is the edge composed merely of the component species of the two
adjacent types? Is there a greater abundance of any particular
species along this line of juncture? How wide is the edge? If the
existing juxtaposition and interspersion do create a favorable ar-
rangement of food and cover, are these so composed and arranged
as to be of value throughout all seasons of the year?

The study presents a graphical and mathematical analysis of the
wildlife habitat potentialities of ten forest types as compared with
the edges between these types. The method of analysis of the vege-
tation in regard to wildlife is presented as being a primary feature of
the publication. The results of the analysis in regard to wildlife
values of the various edges are considered as secondary in importance
and greatly Hmited in dependabihty due to the author's limited
knowledge of the requirements of the species of wildlife considered.
It is suggested that by the method outlined, one may analyze not
only the species composition, the cover value and the general food
value of the various habitats, but also to some extent the seasonal
availability of these constituents. In the light of these determinations,
conclusions may be drawn as to the specific value of these habitats
to the native species of wildhfe.

10

Roosevelt Wildlife Bulletin

The method of mathematical analysis is presented as it may
prove of interest to others initiating this type of investigation.
Though the mass of data may give the method a formidably cum-
bersome appearance, a brief study of its organization will reveal it
to be quite flexible and useful, for it sets upon a mathematically
comparative basis (and thereby simplifies) a type of data which
in the past has been described in only the most general terms.

The ten forest types considered are those most extensively found
on the Check Area of the Huntington Forest and, in general, are
representative of forested areas in the Adirondacks. They are: (1)
spruce flat, (2) spruce swamp, (3) mixed hardwood, (4) northern
hardwood, (5) opening in spruce fiat, (6) opening in northern
hardwood, (7) open swamp, (8) bracken fern, (9) aspen, (10) sec-
ond growth hardwood. The edges, or transitions, considered are as
follows: (1) spruce flat to spruce swamp, (2) spruce flat to mixed
hardwood, (3) spruce flat to northern hardwood, (4) mixed hard-
wood to northern hardwood, (5) spruce flat to opening in spruce
flat, (6) northern hardwood to opening in northern hardwood, (7)
spruce flat to open swamp, (8) bracken fern to aspen, (9) bracken
fern to second growth hardwood, (10) second growth hardwood to
northern hardwood.

B. Description of the Site of the Investigation.

The data for this study were collected on the Check Area of the
Huntington Forest.* Since the problem is ecological in nature, a
fairly complete description of the environment is presented.

1. The Huntington Forest — General. The Huntington Forest is
a 14,984-acre tract, nearly rectangular in outline, situated in the heart
of the Adirondack Mountains just west of the village of Newcomb,
N. Y. The bulk of the area is in Essex County, a small part of the
northwest corner extending into Hamilton County. The main axis
of the Forest lies roughly northwest-southeast ; the 74th meridian
runs just east of the area, while the 44th parallel bisects it.

2. The Check Area. This 4,062-acre block of land has been set
aside to be used for conducting intensive studies of natural phe-
nomena. It is a nearly rectangular block extending two and one-half
miles in the north-south direction and averaging three miles in the
east-west direction, with the north edge one and one-half miles south
of the north boundary of the Forest. A grid system of lines one-
quarter of a mile apart has been established on this area. The trees
along these lines are blazed with rings of orange paint and the inter-

* Hereinafter referred to as the "Forest" or as the '"Huntington."

Edge Effect of the Lesser Vegetation

11

sections of lines are marked with wood signs on metal stakes. (See
Map 1 in the Appendix.)

3. History. There is but little information of a definite nature
available relative to the history of the Huntington Forest previous to
1900. It is known, however, that the Adirondack region in general
was subjected to two series of logging operations before that date.
The first, just after the middle of the last century, removed the ma-
ture white pine which was common along many of the waterways
and throughout the lowlands. The second, toward the close of the
past century, removed a large portion of the spruce. There is evi-
dence that certain parts of the Huntington Forest (including parts
of the Check Area) were subjected to logging during both of these
periods. Great ax-hewn white pine stumps along the lake shores and
streams attest to the fact that a large number of white pine had been
removed prior to the introduction of the crosscut saw ; and rotting
spruce stumps occurring on the flats and slopes indicate another
cutting at a later date.

White pine has not come back to an appreciable extent on the
sites from which it was removed. It is not certain whether this con-
dition is a purely vegetational phenomenon or whether it has been
influenced by some other factor. Maissurow ('35) maintains that, in
Quebec, fire is essential to the re-establishment of white pine in that
it eliminates competition. That white pine is reproducing very spar-
ingly on the Huntington is indicated by the fact that the young
growth is confined principally to the protective cover of the lakeshore
ericads and to a few open old-field sites. Many of the seedlings that
manage to get above the brush cover of the lake shores are promptly
browsed by deer.

Although spruce flat sites normally revert to that type after
logging, it has been suggested that the removal of spruce from the
uplands tends to increase the concentration of hardwoods (Belyea,
'24). This may have greatly influenced the present composition and
extent of hardwood and mixed hardwood types, with possible effects
upon the soil and lesser vegetation.

The area east and south of Deer Lake (as well as a large section
north of the Check Area) was ravaged in 1903 by a post-logging
fire. Most of this area has advanced from the bracken to the aspen
stage or to young growth of other species. Some of it shows a ten-
dency to revert to hardwood while other sections are tending toward
the spruce flat type. Increment borings taken in 1941 from the larger
aspen revealed a maximum age of 30 to 35 years.

There are several localities on the Forest where flooding by

12

Roosevelt Wildlife Bulletin

beaver dams has killed units of several acres of trees and associated
vegetation. The heaver are still resident in some of these places,
while in other places the dams have been deserted but the water level
remains high. These areas have taken on the aspect of open, sedge-
covered muskegs. All of the "open swamp" conditions designated
on the ty])e map of the Check Area were created in this manner.

4. Geology. The .Adirondack Mountains are part of the Lauran-
tian Plateau, which formation extends over the eastern part of Canada
and die northeastern part of the United States. They comprise the old-
est mountain ranges in eastern North America and have undergone a
great variety of diastrophic processing such as igneous activity, fold-
ing, faulting, peneplanation, and glaciation. The characteristic north-
east-southwest valleys of the southeastern and eastern portion of the
Adirondacks are said to be the result of faulting (Bowman, '11).
The results of glaciation are manifested in the varying depths of till
whicli cover both the mountain slopes and the valley floors. In
addition, the valley floors are often covered with stratified drift and
lake deposited material. Many of the lakes, such as Wolf Lake, in
the southeastern portion of the Check Area, have been formed as a
result of morainic deposits in narrow valleys.

The oldest rocks of this region are of the Grenville formation,
and great masses of igneous rocks have been forced upward through
them. Thus there occurs a metamorphosed series of folded sedi-
mentary rocks composed of marbles with interbedded quartzites and
.schists (Miller, T3). "The major portion of the Huntington Forest,
which includes the mountainous parts, is underlain by resistant
syenite and granitic material. Many of the other areas of softer,
more easily eroded marbles have been worn away to form depres-
sions, for example, Newcomb Valley" (Heady, '40).

The topography of the Check Area may be briefly described as
follows: An "L" shaped mountain ridge forms the backbone of the
area; the main axis (Catlin Mountain) extends in a northwesterly
direction and the lesser axis (Panther, Clearing and Observation
mountains) in a southwesterly direction. The large amphitheatre-
like basin in the angle of the two ridges faces Catlin Lake. This
lake has an elevation of 1,597 feet, which makes it slightly less than
1,000 feet below the top of Catlin Mountain. Wolf Lake, which lies
in the eastern corner of the area, is 1,825 feet above sea level while
Deer Lake, on the north edge, is at 1,667 feet. Drainage is north-
westward from Wolf Lake to the outlet of Deer Lake and from here
southwestward to Catlin Lake, which in turn drains through Long
Pond and eventually reaches the Hudson River. The topography of

Edge Effect of the Lesser Vegetation

13

the section north of Wolf Lake, as well as the entire northern third
of the area, is low-lying and not especially rugged. The portion east
of Wolf Lake is tlie northwest slope of Moose Mountain, the hulk
of which lies east of the Forest. Although there are local steep areas
with rock outcrops, the mountain ridges, for the most part, are
characterized hy rounded peaks and gradual slopes.

5. Soils. The various geological processes have had a marked
effect on the soils of this region. Aside from the basic configuration
of the surface and the elevation, slope, and exposure, the surface soil
perhaps is most markedly affected by glacial action. There is very
little surface that has not been covered with glacial till and v^'ater-
borne silt and sand. "Glacial terraces, alluvial fans, and morainic
dumps are often found in the valleys and are of importance to soil
formation and distribution of vegetation" (Heimburger, '34). The
effect of glacial deposits is reflected in the vegetation not only
through the chemical composition of the surface and subsurface soil
but also in the pattern of deposition. Heimburger (I.e.) points out
that whereas valleys as a general rule have fertile soils, glaciation
has in a large number of instances covered these with coarse till.
This results in the best soil occurring not on the valley floor but
rather on the lower mountain slopes.

The Check Area of the Forest is almost completely covered by
varying depths of glacial till. For this reason the soil, for the most
part, is coarse and sandy and contains an abundance of rocks. This
condition has been modified to some extent by the humifying in-
fluence of the vegetation for, although the Adirondacks lie in the
Podzol Soil Group of Marbut ('35), there exist extensive areas of
mull humus under hardwood forest cover. Mull, however, is not the
general rule, for podzol soils are found under both mixed hardwood
and spruce flat forest types, and these types cover over half the Check
Area.

The site cjuality is modified to a large extent by the type of
hvunus. Heiberg ('41) points out that in the southeastern edge of
the Adirondacks, mull humus type is correlated with much more
rapid growth not only in the hardwoods but also in conifers. He
suggests, also, that the flat and superficial type of rooting in the
conifers may have some influence on the type of humus layer. In
agreement with this theory, Donahue ('40) points out that in the
hardwood mull sites, there is a gradual and uninterrupted decrease
in the number of roots from the surface downward, whereas the
coniferous mor humus types contain a marked absence of roots in
the leached layer. The roots in swamp sites are confined to the
organic layer and thus are very close to the surface.

14

Roosevelt Wildlife Bulletin

6. Climate. While geology and soils determine the local spatial
distribution of plant food species, climate controls the seasonal dis-
tribution of these elements. The climate of the Adirondacks is
characterized by great range in daily temperatures, short mild sum-
mers, and long, cold, snowbound winters. The growing season is
relatively short with a range from 79 to 162 frost-free days. Van
Wagner ('19?), speaking of Essex County, states that the average
dates for the last killing frost in spring and the first in fall are June
10 and September 5, respectively, leaving a growing season of 87
days. These dates are based on weather data gathered over a period
of several years at the Lake Placid Club, about 20 miles northeast
of the Huntington Forest. The elevation of this weather station was
1,864 feet above sea level, which approximates the average elevation
of the Forest.

Weather data collected on the Forest during the past few years
indicate that the frost-free period may be as much as a month longer
than the average indicated above. Maximum temperatures are at-
tained during July and August while the coldest weather occurs in
February. The most rapid warming occurs in April and most of the
deciduous plants begin to leaf about the first of May. The influence
of climate on the persistence of fleshy fruits into the fall cannot be
dependably estimated since there are only a few species that char-
acteristically retain their fruits. These few occur with such a low
frequency and in such small amounts that it is difficult to determine
whether their disappearance is due to climatic influences or to wild-
life feeding.

Precipitation is evenly distributed and averages three to four
inches each month of the year. The prevailing winds are from the
west and most of the precipitation has its origin in the Great Lakes.
A considerable share of the precipitation, however, is carried inland
from the Atlantic on easterly and southeasterly storm routes. The
first snowfall often occurs in September and it frequently remains
until late April. January, February, and March show the greatest
depth of snow, with a maximum depth in February. This is the critical
season of the year for browsing animals such as deer, since the deep
snow not only covers the underbrush browse, but it also hinders the
movement of the animals. As a result many deer die nearly every
winter from malnutrition and exposure.

Edge Effect of the Lesser Vc(/etatio)i

15

II. REVIEW OF THE LITERATURE

A. Concept. Indirectly, wildlife is largely a product of vege-
tation. The composition and arrangement of the vegetation on any
area, to a great extent, control the distribution of the associated
fauna. Since vegetation is the most important single factor in the
local distribution of animals, it is through the manipulation of this
part of the environment that the game manager seeks to control
wildlife populations. These facts constitute the basis of wildlife
management as we know it today. (Adams, '25; Dice, '31; Elton,
'27; Leopold, '39.)

Since familiarity with the vegetation is prerequisite to an inti-
mate knowledge of the local fauna and its distribution, a study of the
vegetation should be one of the first considerations in a management
plan. The study of forest vegetation may be approached from one or
more of at least three different points of view: (1) that of the ecolo-
gist, (2) that of the game manager, (3) that of the forester. The
following review of the literature has been developed from these
three points of view.

B. Vegetation, the Viewpoint of the Ecologist. 1. The Or-

ganismal Concept of Vegetation. From the viewpoint of the ecologist,
"Vegetation is the sum total of plants covering an area. ... (It) is
more than the mere grouping of individual plants. It is the result of
the interactions of numerous factors. The effects of the plants upon
the place in which they live and their influence upon each other are
especially significant. ... A study of vegetation reveals that it is an
organic entity and that, like an organism, each part is interdependent
upon every other part" (Weaver and Clements, '38).

The concept of vegetation as an organism is basic to the study
of plant ecology. Wheeler ('11), Gilchrist ('37), and others have
developed this concept of emergent evolution or holism. Egler ('42)
has brought together the various ramifications, which may be briefly
summarized as follows : Elements of lower categories, such as chem-
ical elements (or individual bees) unite or may be united to form
patterns of higher complexities, as molecules, (or bee colonies) and
the new whole is something more than a mere sum of the parts. The
characteristics of the parts may or may not be present in the whole ;
rather, new phenomena or activities emerge as characteristics of the
whole and these phenomena do no reside in the elements. The whole
is adequately self-sufficient, stable, and generally tends to perpetuate
itself in nature, and therefore may be studied as a unit, without
essential recognition of its parts. Organisms exhibit various degrees
of integration, some being well-developed and others being loosely

16

Roosezelt Wildlife Bulletin

formed: furthermore, organisms may unite to form organisms of a
higher category, as individual organisms group themselves to form
social organisms. These organisms are of varying complexity and
may be considered as part of an ascending sequence of categories as
atoms, molecules, individuals, and bee colonies. Organisms are often
in complex relation with other organisms ; for example, the clover-
bumblebee-mouse-cat-spinster complex cited by Darwin ; and one
element of a complex organism may play a role in more than one
organism, as a human being may function in several distinct social
groups.

2. The Principle of Vegetation Development. It is of the
utmost importance, if one hopes to understand the behavior of organ-
isms, and especially vegetation as an organism, that he think of
them as d3namic, as processes, rather than as structures. The vege-
tation is something happening. The underlying principle of vegeta-
tion development is that of constant change and succession toward a
climax. Often the climax is never actually attained and hence is
purely hypothetical. On any site devoid of vegetation, the tendency
is for an initial association of "pioneer" plants to invade the area.
This group of plants so modifies the site as to make it more favor-
able for another association of plants than for its own continued
development. This is followed by a series of other associations until
eventually one which tends to perpetuate itself is developed. This
final association or climax stage continues indefinitely or until it is
eliminated by some catastrophic influence. It is upon the climax con-
dition that most of the classifications of plant zones and formations
of the continents have been based. (Weaver and Qements, '38;
Lundegardh, '31 ; Tansley and Chipp, 26.)

3. The Factors Controlling the Distribution of Vegetation. The
physiological processes of plants permit them to occur only where
environmental conditions are not inhibitory in any manner. Over-
looking the process of long-time migration, a plant occurs only
where it can fulfill all of its life processes. The distribution of plant
species therefore is controlled mostly by environmental factors:
primary, secondary, and chance and coincidence.

Lundegardh (].c.) lists the four most important primar)- factors
as follows: (1) Light. This is important not only because it is the
primary factor in photosynthesis, but also because of the heat and
other qualities of light such as ultraviolet rays which influence phjto-
physiological processes. Xot only the amount and quality of light but
the distribution throughout the year and throughout each day of the
year is of vital importance in the distribution of plants. (2) Tern-

Edge Effect of the Lesser Vegetation

17

perature. This is described as being the "master factor" in the dis-
tribution of vegetation over the earth, aUhough its action is always
interwoven with those of liglit and water. Raunkiaer's system
(Raunkiaer, '18) of life forms and phytogeography is based upon
this factor, as is also Merriam's life zones (Merriam, '98). Studies
in relation to this factor show that each plant species develops best
within a certain optimum temperature range, and that it can survive
only between certain definite thermal limits, provided other condi-
tions remain the same. (3j Water. The "distribution of soil moisture
and rainfall bring about, more than any other factor, the manifold
types of vegetation." Among these are listed tropical rain forests,
tree steppes and savannahs, deserts, subtropical regions, cold tem-
perate forests, meadowlands, the subpolar coniferous zone, and arctic
vegetation. (4 ) Soil. Soil exerts a powerful influence on the plant
in that the amount of root surface, to a large extent, is controlled by
the texture and structure of the substrate, while the chemical com-
position of the soil is reflected in the physiological development of
the plant.

These four factors (light, temperature, water, and soil) are the
basic environmental factors controlhng the distribution and develop-
ment of plants and plant communities. Although the secondary fac-
tors listed below are of great significance locally, they are not so
fundamental in aspect nor so controlling in the broad sense as are
these.

The secondary factors may be divided into three groups : ( 1 )
site factors, (2) developmental factors, and (3) catastrophic factors.
The first of these includes such influences as geographic location in
relation to the equator, the poles, and bodies of water. Also in this
group are such factors as altitude, exposure and slope. These factors
control the distribution of plants through the influence of the primary
factors. This group of secondary factors is dififerent from the other
two groups in that it directs the development of plants and plant
associations along certain lines.

The next group of secondary factors is one which may operate
on a much larger scale than is generally realized. In this group fall
the influences of pollen-bearing insects and seed-eating birds and
mammals. This conjuncture constitutes a potent influence since it
is only through the action of insects that a great variety of plants are
able to perpetuate themselves, while fruit- and seed-eating animals
assist in the distribution of plants to new areas and to areas that
have been denuded of vegetation. The aeolian element as a factor in
both pollination and seed dispersal must also be included in this
group.

18

Roosevelt Wildlije Bulletin

The third group of secondary environmental factors (catas-
trophic) is controlling in that they usually set back vegetational
development to an earlier stage of succession. These factors operate
locally to determine the serai stage of the plant formation. Among
these factors may be listed: (1) activities of man, such as habitation,
cultivation, logging, stock raising; (2) wildlife activities, such as
browsing by ungulates, flooding by beavers, porcupine damage, de-
struction by carpenter ants; and (3) periodic catastrophes such as
floods, fires, and hurricanes. Sometimes a combination of these
factors operates over a long period and prevents a serai development
from attaining its climax. Such vegetational situations are called
subclimaxes.

To this elaborate set of factors and influences must be added
one more consideration which may best be approached by asking
the questions : What is the nature of the force that operates this
extensive set of controlling factors? Although the activities of man
and some of the activities of certain wild animals may be controlled
to a small extent, what is it that determines when, where, how, and
to what extent the various vegetational influents shall operate? Since
it is not the purpose of this work to delve into a consideration of the
supernatural, it will be necessary to assume that all the interplay of
environmental factors and the resultant effects upon vegetation are
a direct result of chance and coincidence, and that this all-controlling
(or all-noncontrolling) element operates both with and against the
principle of succession toward a climax.

4. Edge Vegetation. Although ecotones are mentioned in the
literature (Weaver and Clements, '38; Braun-Blanquet, '32) they
have been subjected to but little ecological study. Among the few
discussions of edge vegetation in reference to the principle of suc-
cession may be listed that of Gleason ('27). This lack is indeed a
surprising circumstance, especiaUy in view of the w-ide and intense
interest of wildlife managers in this phenomenon. To be sure, nu-
merous recommendations have been made by qualified individuals as
to how field-woodlot edges may be improved by planting food and
cover species of value to certain game animals. Most of this work,
however, has been in reference to farm game habitats and the eco-
logical principles involved have received relatively little attention. A
brief discussion of a few basic features of edge vegetation, therefore,
are appropriate.

The terms "edge," "ecotone," and "transition" (here used inter-
changeably) refer to the zone of juncture between two vegetational
associations. This zone of juncture may be quite narrow or very

Edge Effect of the Lesser Vegetation

19

broad. At the one extreme would be the line between an open field
and an adjacent woodland; at the other would be the very broad
ecotone between the southwestern desert plains and the mixed prairie,
estimated by Clements and Shelford ('39) to be a hundred miles or
more in width through Texas and New Mexico.

In the extremely narrow type of edge, one plant association be-
gins where the other ends and there is no graduation of species from
one association to another. The "transition" type of ecotone on the
other hand, is a zone wherein one association merges gradually into
another and there occurs an intermingling of the characteristic species
of each association. This is the type of transition most widely repre-
sented on the Huntington Forest.

The height, also, is an important consideration of an edge. It
may be very close to the ground ; between an area covered with
bracken fern and an adjacent area dominated by sedges. Or it may
be slightly higher ; between the raspberries of a clearing and the witch
hobble of the forest. Or it may be in the forest canopy, the spruce
and fir of a spruce flat and the beech and yellow birch of a mixed
hardwood type. The edges in each of these strata do not always
occur at exactly the same line, as can be seen by examining the
various graphical analysis sheets presented in the discussion of each
edge.

The edge may be characterized not only by a change in species
composition, but also by a change in the size or amount of a given
species. For example, witch hobble is present in both spruce flat and
northern hardwood but it attains a much better development in the
northern hardwood type. Dwarf dogwood also is present in both of
these types but is much more abundant on the spruce flat side of the
transition.

The type of edge most beneficial to wildlife depends entirely
upon the location, composition and physical characteristics of the
edge and the requirements of the species of wildlife involved. Al-
though we have been accustomed to thinking of edges as being of
value principally to species of comparatively low mobility such as
grouse, quail and rabbits, it should not be overlooked that these same
benefits also may be of value to larger animals, as for example, deer.

The conception that edges are of a more dynamic nature than the
centers of plant associations (Weaver and Clements, '38) should not
be misinterpreted. There is just as much plant activity and competi-
tion in the interior of a plant association as at the edge ; the difference
lies not in the amount of dynamic activity, but rather in the variety of
competing species. The competition in the center of an association is
between individuals of the same species or group of species and is

20

Roosevelt IVildlifc Bulletin

not so strikingly noticeable to the observer. Competition at the edge
between two forest types is much more apparent because here one
can observe the results in the comparative development of the com-
ponents of the adjacent associations. An edge, therefore, should not
be considered especially dynamic except in the sense that it may
represent movement by a vegetative association.

It is only logical to assume that the factors of environment which
determine the location of edges are the same as those determining the
distribution of vegetation. It should be realized, however, that the
factor, or combination of factors, determining the edge on one side
of an association need not necessarily be the same as the factors Hm-
iting the association at the opposite side (Sampson, '39 j. Conversely,
more than one combination of factors may be conducive to the estab-
lishment of the same kind of edge.

Plant associations have been considered as moving organisms
(Glcason. '27) and the movement is most readily discerned at the
periphery of an association. The rate of vegetational movement,
however, varies extremely. Consider, for example, the rapidity with
which an abandoned field is invaded by an adjacent woodland as
compared to the length of time it takes for a spruce flat forest type
to assimilate a bog. The former type of edge advances with relative
rapidity, while the latter may appear comparatively stationary.

The principle of successional development in the study of vege-
tation allows for at least four different kinds of edges : ( 1 ) the edge
of a vegetational invasion into a barren, unvegetated area; (2) the
edge between two serai stages; (3) the edge between the serai stage
and climax ; and (4) the edge between two climax formations. Ex-
amples of each of the last three types will be considered in this study.
These may be compared as to rate of change or movement as follows :
(1) most rapid change — serai stage to serai stage; (2) intermediate
— serai stage to climax: (3) most permanent — climax to climax.

C. Forest Types, the Viewpoint of the Forester. The for-
ester, though his lands are governed by the natural laws as revealed
by the ecologist, is primarily concerned with timber, which is only
one phase of the vegetation. His interest is primarily economic and
therefore he classifies vegetation with an eye to the public market.

1. The Basis of Classification. Forest type classifications and
terminology were controversial issues among foresters for some time,
but indications are that discrepancies have been largely eliminated.
The literature is replete with disagreement as to what constitutes a
type and upon what bases type classification should be placed. The
subject has been made more confusing by the use of ambiguous and

Edge Effect of the Lesser Vegetation

21

special terms. Dana ('13) listed 17 dif¥erent categories of types as
previously recorded in the literature : temporary, transitory, inter-
mediate, derivative, timber, dendrological, cover, permanent, funda-
mental, original, mother natural, physical, natural physical, final,
ultimate, climax, and management. This serves to demonstrate the
extreme variance of thought and terminology, as w^ell as overlapping
concepts prevalent at the time.

Another point at issue has been whether species composition or
site ought to be the basis of type classification. Some individuals
(Zon, '08; Dana, '13) with a more profound appreciation for what
is involved have urged that both be considered. Various opinions
have been expressed relative to the importance of site features such
as soil, moisture, chemistry and climate. Frothingham ('18) main-
tained that height of dominant trees is the key to classification of
site quality, while Bates ('18) declared that "the only final criterion
of the site quality is the current annual cubic foot increment of a
fully stocked stand of the species under consideration." While eco-
logical succession was appreciated by some, the concept of succession
and climax as it is thought of today was not apparent.

This early period of American forestry was characterized by the
initiation of experimental and scientific analyses of site factors. In-
dividual factors were considered : soil, moisture, sunlight, heat, pH,
and length of growing season. Bates (I.e.) concluded that the role
of light beyond a certain minimum required for photosynthesis is
that of supplying heat so that the thermal threshold of physiological
activities might be reached. Zon ('08) claimed that "no other cli-
matic factor has such direct bearing on forest growth as light." Moore
('17) suggested that osmotic pressure of cell sap might prove to be
an index of forest habitat. This period of research is now taking a
turn toward more complex studies, and toward the realization that
it is not one factor alone but a combination of factors that determines
the vegetation of an area.

The subject of forest type classification on the basis of ground
vegetation has recently received some attention. Most of this work
has emanated from Europe and may be represented by the work of
Cajander ('26). Site classification on the basis of ground vegetation
has been attempted in this country by Ilvessalo ('29) and Heim-
burger ('34). The arguments put forth by the proponents of this
theory are: (1) that minor vegetation reflects the quality of site
better than does the dominant vegetation and (2) that forest sites
are to a high degree independent of the influence of the forest stand
that may occupy the area at the moment. Although Coile ('38)
doubts the validity of these arguments, it has not been conclusively

22

Roosevelt IVildlife Bulletin

shown that this approach is not worthy of further consideration.
Coile admits that there is a correlation between dominant vegetation
and lesser vegetation, but he maintains that the upper story is ex-
tremely influential if not all-controlling in the composition of forest
floor cover. He argues that physical features of topography and soil
are more basic indicators of site than lesser vegetation. Sisam ('38)
correlates herbaceous vegetation with aspen site quality, but Samp-
son ('39) urges caution in the use of indicator species until more is
known about the manner in which they react to the various factors of
the environment.

2. Forest Types of the Adirondacks. The forest types of the
Adirondacks today are considered in essentially the same manner as
Graves ('99) classified them. His classification included : (1) Swamp
land, which consists of low flats with wet, spongy s jil characterized
by red spruce, balsam, black spruce, tamarack, black ash, and cedar,
with frequent "blowdowns" due to shallow rooting. (2) Spruce flats,
or the "level and rolling flats bordering on lakes, streams, and swamp."
The dominants of this site were listed as : spruce, birch, beech, balsam,
and hemlock. (3) Hardwood land, sites located "on elevated benches
and moderate slopes" characterized by a dominance of spruce and
yellow birch if the land is broken, "while hard maple and beech are
confined to the moderate north slopes and bases of the ridges and to
the high benches." These latter sites are listed as being dominated by
sugar maple, beech and yellow birch. (4) Spruce slopes, or steep
slopes with thin stony soil characterized by spruce, yellow birch, hem-
lock and white pine.

The plant formations of the Adirondack Mountains, according to
Harshberger ('05) are somewhat similar, but cover a greater range
of topographic situations: (1) deciduous forest formation, (2) conif-
erous formation, (3) alpine plant formation (on the tops of the highest
mountains), (4) bog formation, (5) hemlock formation, and (6)
rock-gorge formation.

More recent investigations, as those of McCarthy and Belyea
('20) and Donahue ('40) follow this simplified arrange nent of forest
types: (1) spruce swamp, (2) spruce flat, (3) mixed hardwood, (4)
northern hardwood, and (5) spruce slope. The principal difference
between spruce slope and spruce flat is that the latter normally occurs
above 2,200 feet, which is usually above the hardwood type.

Heimburger ('34), seeking to develop a classification based on
lesser vegetation, established a more elaborate set of forest types, first
dividing the entire Adirondack region into : subalpine series, western
series, and eastern series. In each of these series he set up a number

Edge Effect of the Lesser Vegetation

23

of forest sites based on the composition of the herbaceous and shrubby
vegetation as well as the dominant trees. His argument for this
arrangement was that it would simplify the interpretation of site
quality in reference to the dominant trees and it would facilitate man-
agement practices to the extent that the forester could avoid treating
a site for the best development of a species which actually might not
do well on that site. In order to accomplish this, Heimburger redivided
each of the major forest types into a number of forest sites which
can be recognized in the field with but a little extra effort.

3. Forest Types of the Huntington, Forest. Twenty forest types
are represented on the type map of the Check Area (Map 1) but

TABLE 1. FOREST TYPES ON THE CHECK AREA
OF THE HUNTINGTON FOREST

Percent of
Percent of Census Line
Forest Type Area, Acres Total In Type

Northern Hardwood Site :

Northern Hardwood

1486.9

36.60

36.558

N. Hardwood (Second Growth)

47.8

1.17

1.118

3.0

.07

1.9

.05

'.028

Mixed Hardwood Site:

1212.5

29.84

28.252

M. Hardwood (Second Growth)

88.6

2.18

3.287

.9

.02

Spruce Flat Site:

Spruce Flat

777.6

19.14

19.057

Bracken Fern

37.6

.92

1.210

Aspen

57.8

1.42

.896

Spruce Flat (Second Growth) . .

38.5

.95

1.336

16.7

.41

.454

Temporary M. Hardwood

12.7

.31

Forest Opening;

2.3

.06

Lower Spruce Slope Site:

Lower Spruce Slope

144.2

3.55

4.123

Rock Slide

0.7

.02

.103

Spruce Swamp Site:

Spruce Swamp

49.2

1.21

1.189

44.6

1.10

.829

Alder Swamp

24.3

.60

.937

Black Ash

1.0

.02

.055

Pond

Pond

13.9

.34

.570

Totals — All Types

4,062.7

99.98

100.002

24

Roosevelt Wildlife Bulletin

these may be grouped into four climax forms of vegetation: (1)
hardwood, (2) mixed hardwood,* (3) spruce flat, (4) spruce slope.
The hardwood corresponds to \\'eaver and Clements' "Deciduous
Forest" while the spruce types correspond to their "Boreal Forest."
There is no typical "Lake Forest" on the Huntington. The terms
"spruce flat" and "spruce slope" do not of course imply pure conif-
erous composition, for very often such species as yellow birch and
red maple are in the association. The mixed hardwood forest type
possibly ma}' Ije regarded as a broad and fairly stable zone between
the spruce types and the hardwood. Each of these four climax types
is silviculturally mature. From a vegetational standpoint, the other
forest types listed on the type map are merely serai stages and sub-
climaxes in the development toward the climax forms.

The terminology of the forest types on the Check Area (Table
1) is that commonly used by forest operators. The most extensive
single type is northern hardwood, composed predominantly of sugar
maple, beech, and yellow birch, with an admixture of spruce and
hemlock. The mixed hardwood type is next largest in extent and is
composed predominantly of yellow birch, spruce, beech and sugar
maple, with a scattering of hemlock, red maple, and balsam. The
third most extensive forest type is spruce flat and is composed for
the most part of spruce, yellow birch, balsam, hemlock and red maple,
but beech may be scattered locally near the upper edges, while arbor
vitae, white pine and white birch may be found along the lake shores.
These three types comprise approximately 86 percent of the forest
cover of the Check Area. About three percent is in swamps and
clearings and 3.5 percent is in spruce slope, while the remaining eight
percent has been either burned over, or logged or both. Reference
to the type map (Map 1) will give a clear picture of the relative
areas and distribution of the cover types listed.

D. Vegetation and Wildlife Distribution, the Vievirpoint of
the Wildlife Manager. 1. The Controlling Influence of Vegetation.
The wildlife manager's conception of vegetation, as pointed out above,
must, of necessity, be an interpretation of vegetation concerned with
those qualities which make the environment habitable for certain
species of animals. He not only looks for food and cover but for an
appropriate arrangement of these factors, as well as for their presence
at all seasons of the year. His principal interest in vegetation is
tliat it support a certain amount and variety of wildlife.

* The inclusion of mixed liardwood stands as a climax type may be open to
some question since it is not known to what extent early spruce cutting affected
their composition or distribution. Most of the mixed hardwood on the Check
Area is silviculturally mature and contains both hardwood and softwood.

Edge Effect of the Lesser Vegetation 25

The fundamental relationships between vegetation and wildlife
distribution are thoroughly discussed in the writings of Dice ('31)
and Elton ('27). Elton states that there is not a single animal suffi-
ciently elastic in its organization to withstand the wide range of en-
\ ironmental conditions which exist in the world ; for this reason each
different habitat contains a characteristic set of animals. Local vege-
tation is of especially great importance. Its effect, aside from direct
food relations, is to tone down the intensity and modify the extremes
of the outside climate. Some animals are wholly dependent upon one
or two plant species and therefore are found only where these plants
occur. They may be described as being "exclusive" to such areas.
"Characteristic" species, on the other hand, are those with wide
limits, occurring in large numbers in an area but not confined to it or
to any particular single species of plant. Elton (I.e.) further states
that it is probably true that animals living in several zones of vege-
tation show a marked tendency to have their limits of distribution
coinciding with the edges of the plant zones.

Dice (I.e.) states that plants probably are the most important
feature in the environment of mammals, as well as of all other
terrestrial animals and that all mammals are dependent directly or
indirectly upon plants for food. Furthermore, plants also furnish
many mammals with shelter from the weather, sites for homes, nest-
ing material and protection from enemies. He states that mammals
usually are dependent upon types of vegetation rather than upon par-
ticular species of plants for food, homes and protection ; that although
the forest in major degree makes the habitats for many of the in-
habitants, and while the height and density of the forest cover are
important environmental factors for the mammals, the precise tree
species involved probably are of little consequence, so long as the
quality of food and shelter is the same.

Admitting that mammals generally are very adaptable, and that
there are many forms which range over several vegetation types,
Dice (I.e.) notes that there is. nevertheless, a considerable amount of
correlation between mammalian distribution and types of vegetation.
In the case of the "Characteristic" species mentioned above, the
presence of certain plants might be considered to indicate conditions
suitable for the presence of particular mammal communities. Dice,
however, cautions against the use of plant species as indicators in
this sense because the habits of mammals apparently differ to some
extent in different regions, and so also does the growth of plants.

Another point brought out by Dice is the relation of vegetational
succession to animal distribution. A number of ecologic plant com-
munities are found in every region which are successional stages

26

Roosevelt Wildlife Bulletin

leading toward the climax. Some of the successional stages in any
given region may be very different in vegetation type from the climax
and, therefore, will constitute a very different environment for the
animals. Some of these successional communities, however, are of
quite constant occurrence and some cover extensive areas ; therefore,
from the standpoint of mammalian distribution and successional com-
munities they may be almost as important as the climax community.
Although only of presumably brief duration on any spot, certain of
these communities may be present at all times in some part of the
province and, therefore, constitute an important factor in the long-
range evolution and adaptation of faunal societies.

As was mentioned above, many species of animals range over
several vegetation types, and individual animals may move freely
within several plant communities. Therefore it may seem useless to
try to apply the very minor ecologic subdivisions to these animals.
Even mice, shrews, and other small mammals which have relatively
short cruising radii, show little evidence of limitation to minor plant
communities. Dice indicates that in view of these facts one should
consider animal distribution to be more strongly correlated with
major vegetation types.

The fundamental relations presented by Elton and Dice are
further demonstrated by other authors. Beecher ('42) states that
"it is not difficult to demonstrate that, within any selected area, the
vegetation is the prime limiting factor in the distribution of birds."
Van Deventer ('39), in discussing the influence of secondary plant
communities on bird populations in an area of cut-over woodlot at
Kankakee, Illinois, set forth a classification of communities based on
permanence. He found each of three communities characterized bv
a different group of birds. Dambach and Good ('40) also found a
definite correlation between agricultural land use practices and bird
populations.

Harper and Harper ('29), in a study of animal habitats in cer-
tain portions of the Adirondacks, came to the conclusion that "Gen-
eral conditions for animal life appear to be slightly more favorable in
predominantly deciduous woods rather than in predominantly conif-
erous woods."

Bennett, English, and McCain ('40) offer a method of deter-
mining the difference in deer habitat value of various forest types,
on the assumption that these animals have definite habitat preferences.

Davidson ('32) made a study of invertebrate forms in a series of
successional stages of vegetation and found several significant bio-
ecological relations: (a) The climax maintains a cooler, moister and
less variable microclimate than that of subclimax stages, (b) The

Edge Effect of the Lesser Vegetation 27

populations increase from the pioneer stage of vegetation to the
cHniax stage, (c) The number of seasonal and incidental species
decreases as the succession proceeds and is least in the climax, (d)
The number of perennial predominants increases toward the climax,
(e) The preclimax stage is characterized by several groups of species
of relatively large numbers. Current predominance of any of these
groups varies with environmental conditions exclusive of those
correlated with the vegetation.

2. Edge Effect. Most of those investigators who have studied
animal distribution in relation to vegetation have come to the con-
clusion that edges are of greater value than the centers of plant
associations. For example, King ('38) stresses the importance of
interspersion and juxtaposition of vegetation, while Lay ('38) found
that the margin between forest and clearing had 95 percent more
birds representing 41 percent more species than did the interiors of
corresponding woodland. Leopold ('39) states that wildlife is a
"phenomenon of edges."

Beecher ('42) found a remarkable correlation between edges and
bird distribution in bird communities of northern Illinois. He says
"the population density of most nesting birds, and doubtless many
other animals as well, varies as a direct function of the amount of
edge per unit area." Beecher broadened the concept of edge effect
by suggesting that edges are efifective as such only when they are
above a minimum "threshold" acreage (ratio of edge to acreage).
This is illustrated by a plant community so small or narrow, as for
example a hedgerow, that a nesting pair on or near one edge is no
appreciable distance from a pair nesting on the opposite edge — in
fact sufficiently narrow so that it is impossible to tell which edge the
pairs react to. If the area of the community is expanded, the nest
distribution, following the edges outward, will leave a center rela-
tively free of nesting. When the area reaches sufficient size for even
a slight edge preference to be evident, the corresponding sparsity of
nesting in the center will cause a lowering of the density per unit
area of the whole. This point is described as the "threshold acreage"
of edge preference.

While Beecher's concept of "threshold acreage" is of value in
certain phases of wildlife work, the manner in which the concept is
developed presupposes a habitat so diverse in composition and form
as to be highly attractive to many species of animals, whereas a
threshold condition usually is associated with a minimum population.
The concept of "threshold acreage" of edge preference should there-
fore not be confused with a straightforward definition of the threshold

28

Roosevelt W'Udlijc Bulletin

of edge preference. The threshold of edge preference is attained when
the edge between two plant associations becomes sufficiently varied
in form and composition so that it holds a stronger attraction for
certain species of wildlife than do the centers of the adjacent asso-
ciations. This of course assumes that there are edges which do not
exert any edge effect, and starting with this condition, there are
edges which exert varying degrees of attraction for certain species
of wildlife. The extent to which an edge affects wildlife varies
according to the requirements of wildlife species and the extent to
which the edge vegetation fulfills these requirements. An edge may
attract certain species of wildlife and at the same time hold no
attraction for others. In fact, as observed by Beecher (I.e.), some
animals are actual!}- repelled by certain types of edges.

Through its influence upon the distribution of herbivorous ani-
mal species, edge effect may be found to function in the activity,
habits, and distribution of carnivorous species. For example, Aldous
and Manweiler ('42), in a study of the short-tailed weasel, found
that "Trapping operations were most successful . . . within two
hundred \ards of the swamp periphery and along drainage ditch
grades running through the region that produces the same general
flora."

Davidson's {'22) observation that spiders occurred with greatest
percent of frequency in the forest edge presocies is further evidence
of this effect.

Among other authors attesting the wildlife value of edges may
be listed Deem ('38) who says 'Tn any manipulation of forest cover
seeking to improve environment for wildlife this edge effect must
be recognized. This means that overly-large contiguous areas are
to be avoided." He points out that conifer types in younger stages
provide good cover sites, especially for grouse. He advises selection
cutting in coniferous types as well as clear-cutting of small areas,
and points out that sprout hardwood provides good deer browse.

Edminster ('39) says that "To preserve edge effects, no point
within a woodland should be more than three hundred feet from a
margin. \'ariety in vegetation should be sought and succession
allowed for. while devoting field borders to shrubbery promotes the
edge effect and increases food and cover." Bump ('35). too, stresses
the value of openings and forest margins. He states that seven times
as much game was found in natural mixed cover as in areas refor-
ested w'ith conifers only. The need for maintaining openings for the
good of wildlife is emphasized in a report of the George ^^'ashington
National Forest (\\'ayland, '38). It was found that the grouse and
turkey populations increased threefold to eightfold by clearing areas

Edge Effect of the Lesser Vegetation

29

and improving them for wildlife. Calculations are presented to the
effect that expenditures for wildlife improvements are justified even
on the best timber areas and that making clearings which result in
more wildlife can be done at a profit.

King ('37) points out that it is very important to have a favor-
able distribution of various cover types in the management of rufifed
grouse. Edminster ('35) in analyzing a reforestation project found
that it is only the marginal portion of the reforested area that is used
by grouse, rabbits and deer, and that the interior is barren of wildlife.
It is therefore recommended that coniferous plantations be not more
than six hundred (preferably four hundred) feet in diameter and
that they be separated by plantings of hardwoods and fruit-bearing
shrubs and interspersed by open lands.

E. Wildlife and Forestry. The point of view taken in this
study is a combination of all of the above-mentioned three. Any
sound investigation of vegetation must be based upon fundamental
ecological principles. On the other hand, it must be kept in mind
that many areas in the region represented by this study are main-
tained primarily for their timber value and that any application of
wildlife management must be organized on the basis of vegetational
types as established and employed by the forester. There has been
considerable difference of opinion as to the compatibility of forest
and wildlife management on the same area. It is believed, however,
that the present trend is toward the acceptance of forest wildlife as
a resource worthy of consideration.

Although McAtee ('36) states that wildlife management and
forestry are almost entirely opposed there are several other prominent
authorities who see the way clear to a compromise favorable to both
interests. Hosley ('37) maintains that there is no antagonism be-
tween the production of trees and of game; that in fact "There
seems to be no silvicultural operation which cannot influence wildlife
to advantage, if properly used." As an example, we may cite Hormay
('40) who, in discussing the effect of logging on forage, reveals that
although grazing capacity is reduced about twenty percent for a year
or two following logging, it rapidly builds up and surpasses the
capacity of the former timber-dominated forage.

Leopold ('30) stresses the importance of coordinated effort be-
tween forestry and game management. He points out that the com-
bined effort of foresters and game men would yield two crops where
only one was produced before. Going one step further, Morton ('36)
claims that foresters fail "in their duty to the people and to the pro-
fession when they do not give proper recognition to wildlife in the

30

Roosevelt Wildlije Bulletin

plans of management . . . Participation in the development of the
forest for the greatest good to all is not only a forester's privilege,
it is his duty." Clepper ('36) offers a compromise of mutual advan-
tage to wildlife and to forestry when he states that orderly improve-
ment cutting will increase carrying capacity of forests and improve
the condition of the deer. He further indicates that such work
actually has been initiated on Pennsylvania state forests.

III. COLLECTION OF DATA

A. The Transect. The transect used in this work was a
modification of the belt transect described by Weaver and Clements
('38) in that a series of plots instead of a continuous strip was em-
ployed. It may be noted, in passing, that the widths used were con-
siderably wider than recommended above in order to insure an
adequate sample. The chief advantage of the plot method is that data
gathered in this manner lend themselves more readily to both graph-
ical and mathematical analyses. In order to sample the three layers
of vegetation, i.e., tree canopy, underbrush and herbs, three sizes of
pk)ts were used. The trees were recorded on a series of contiguous
circular plots 74.46 feet in diameter and covering one-tenth of an
acre. The shrubs, together with the arborescent reproduction (col-
lectively referred to as underbrush), were recorded on a series of
quadrats 6.6 feet square covering an area of .001 acre, i.e., one-
hundredth the area of the tree plot. The herbaceous vegetation was
recorded on 3.3-foot quadrats located in the proximal lefthand
quarter of the underbrush plot. These were .00025 acre in size.

The advantages of this arrangement of plots may be listed as :
( 1 ) the various plots are large enough to embrace adequate samples
of each of the three life forms: (2) data from a series of plots may
be gathered more rapidly and are more easily analyzed mathema-
tically than those from a continuous belt; (3) the fact that the tree
plots cover one-tenth acre and are one hundred times the size of
the underbrush plots, and these latter four times the size of the herb
plots, makes it convenient to convert all the data to an equal areal
basis, or to a per acre basis.

B. Arrangement of Plots and Equipment Used. The under-
brush-herb plots were spaced at 24.8-foot intervals along the transect.
Three times this figure is 74.4, the diameter of the tree plot. In other
words, there were three underbrush-herb plots in every tree plot, and
every third underbrush-herb plot marked the center of a tree plot.
Reference to the accompanying diagram (Fig. 1) will assist the reader
in understanding this arrangement.

Edge Effect of tlie Lesser Vegetation

31

Fig. 2. A sample plot of young trees, shrubs, and herbaceous vegetation.
The corners of the underbrush plot have been marked with stakes and the
herb plot is marked with sticks laid inside the larger plot. On this plot
underbrush (shrubs) occurred with a density of Three.

32

Roosevelt IVildlijc Bulletin

Equipment consisted of a two-chain steel tape, a hand compass
and a pair of hinged sticks 6.6 feet long. The hinged sticks were used
to delimit two sides of the underhrush plot; the proximal half of each
arm was unpainted while the distal half was painted red. Thus the
colored end next to the tape established one side of the herb plot while
the adjacent side of the herb plot was bounded by the chain.

Three sides of the underbrush plot were delimited, respectively,
by the chain and the two arms of the hinged sticks. Points were
marked in orange paint along the length of the chain 24.8 feet apart.
Thus every paint mark established the location of an underbrush-herb
quadrat, and every third point was used as the center of a tree plot.
For the sake of randomization all the quadrats were laid to the right
of the direction of travel so tliat eacli paint mark on the chain set the
lower left-hand corner of the quadrat. One arm of the hinged stick
was laid perpendicular to the tape opposite the paint mark and the
other arm parallel with the tape. (See Fig. 1.)

C. The Tree Plot. As mentioned abo^•e, these were a series
of contiguous circular plots 37.2 feet in radius or one-tenth acre in
area. The center was marked with paint on the chain while at least
two radii could be determined at 56 links from the center. With
these two sides to check on, the other radii were determined by pac-
ing and ocular estimate.

The trees were tallied by species and size. The diameter size
classes w^ere rather broad, ranging thus: l"-3", 3"-6". 6"-12". 12"-18",
18"-24", 24"-30" and over 30". Trees under one inch in diameter
were classed with the underbrush and were tallied as reproduction
on the underbrush plot. These broad size classes facilitated estimat-
ing and at the same time this segregation gave a good indication of
the relative numbers of dominants, subdominants, and suppressed
individuals of each species.

D. The Underbrush Plot. As stated above, these were quad-
rats 6.6 feet square, covering .001 acre. Two groups of data were
recorded from these plots : ( 1 ) young trees under one inch in dia-
meter, and (2) shrub species.

The young trees were tallied in a manner similar to that em-
ployed for the mature stand, except that the size classes were based
on height rather than diameter. These size classes were : 0"-6" high,
6"-24" high, and 24" high to 1" diameter. This segregation set ofT
arborescent winter browse (24" high to 1" diameter) from that
available at other seasons. It also served as an index of the age at
which the excess dominant vegetation dies, and as an indication of

Edge Effect uj the Lesser Vecjctatioii

33

rj. DENSlTli:S

Fig. 3. Density values. A species of underbrush covering less than four
square feet was ascribed a density of Trace; if it covered from four square
feet to one-third of the plot, it had a density of One; if it covered from
one-third to two-thirds, it had a density of Tivo; and if it covered from
two-thirds to three-thirds, it had a density of Three.

Fig. 4. "Readiii- Iib-I. The density of the shrubs or herbs

could be estimated fidin one .-ide, but small tree seedlings required careful

counting.

34

Roosevelt IVildlifc Bitlletin

v/hat species were coming into the plant association, and was thus a
measure of the stability of the existing forest type. (See Fig. 2.)

The shrub species tallied from the underbrush plots included all
woody plants with more than a single stem, not attaining twenty feet
in height, and growing in a clump rather than as individual plants.
Among these were such species* as witch hobble**, alder, honey-
suckle, and raspberry.

Since it would have been impractical to use the tally method on
the shrubs, the method adopted was based on relative density and
frequency. An arbitrary range of four degrees of density was es-
tablished, much as was done by Webb ('42). A density of "trace"
denoted less than four square feet of cover while a density of "one"
indicated that the species in question covered from four square feet
up to one-third the entire area of the quadrat. A density of "two"
was ascribed to a sjjecies if it covered between one-third and two-
thirds of the total area ; and a density of "three" indicated that the
species covered from two-thirds to all of the area of the quadrat.
(See Figs. 3 and 4.)

Total density of the shrubs for herbs) refers to the combined
coverage of the shrub (or herb) species on the quadrat. The total
density, however, is not necessarily the sum of the densities of the
individual species. Furthermore, the total density does not always
exceed the largest individual density. The reason for this can be
seen in the width of the density classes ; for example, a quadrat
might have three species of shrubs, each covering just over four
square feet, and therefore each having a density of one, but the total
area that they cover might be less than one-third of the area of the
entire quadrat ; hence, the total density also would be only one.

E. The Herb Plot. As stated above, the herb plot occupied
the proximal left-hand corner of the underbrush quadrat. It was 3.3
feet square, or a quarter mil-acre in area. The herbaceous species
were recorded on it in much the same manner as was done in the
case of the shrubs. The areas of coverage necessary for densities
of one, two, and three were proportionally less but were still limited
by the same fractions. One square foot co\er was the upper limit
of the trace density.

Although over 600 of the 761 vascular species on the Hunting-
ton Forest are herbs, relatively few of these were included in this

* This term is used ratlier broadly in a few instances ; for example, all grasses
have been himped together as one "species," as have all sedges and violets. Other
aberrations are covered in the list** mentioned blow.

** The scientific names of all plant species referred to in the analysis are re-
corded below in the section "List of Plant Species."

Edge Effect of the Lesser Vegetation

35

study. It would be both impossible and impracticable to include all
of these in a study of this nature. Hence only the most common
species were used — those which were consistently and widely dis-
tributed in sufficient numbers to be considered as characterizing the
vegetation. It is estimated that the species considered constitute over
99 percent of the total herbaceous cover of the area studied.

F. Location of the Transects. Although it was impossible
to completel}- randomize the data, some quality of randomization
was accomplished by two methods. The first has been mentioned
already, that is, the method of locating the plots in a straight line
along a given bearing. The location of the transects was further
randomized by first selecting them on a type map of the Check Area.
After selecting the most representative locations for the transects,
their positions were plotted on paper and the transects were then
established in the field.

The transects were so located that they crossed the transition
between typical sections of the selected forest types. The abruptness
of the ecotone was found to vary for each group of transitions and
there was considerable variation within each group. The transects
together with their lengths and numbers of plots are listed in Table
2. This table includes 126 transects averaging 6.06 chains in length
and if placed end to end would extend 9.55 miles. The flora were
listed from 671 tree plots, 2,031 underbrush plots and 2,031 herb
plots, making a total of 4,733 plots.

IV. ANALYSIS OF DATA— METHODS

A. Preliminary Tabulation. The data in reference to each
transect were transcribed from the field sheets to graphical analysis
forms. Tree distribution data were plotted as shown in Graphical
Analysis Form 1 (Fig. 5). Each dot on this form represents an
individual tree. Where there were more trees to record than avail-
able spaces in the form, the number of trees was entered. It should
be noted that both mature trees and reproduction were recorded, the
former from the tree plots and the latter from the underbrush plots.
In a corresponding manner, the preliminary tabulation of the shrubs
and herbs was as shown on Graphical Analysis Form 2 (Fig. 6).
The various species were recorded by densities, i.e., one dot repre-
sented a density of trace, two dots a density of one, three dots a
density of two, and four dots a density of three. The center of tran-
sition on both forms was indicated as being the one tree plot on
which the transition from one forest type to the other was most
apparent.

GRAPHICAL ANALYSIS FORM
TRANSITION:- MA-fZ? HDWD.

EDGE

TRANSECT NO HI
NORTHERN HARC^VOOD

The tree data from each transect were transcribed from the field
sheets to Graphical Analysis Form 1.

GRAPHICAL ANALYSIS FORM ^
TRANSITION:- MIXED HARDW.

EDGE

TRANSECT NO HI
NORTHERN HARDWOOD

TREE PLOTS

1

2 2

3

4

5

6

BRUSH PLOTS

1 2 3

4 5 6

7 8 9

10 11 12

13 14 15

16 17 18

UNDERBRUSH SF

RASPBERRY. RED
HONEYSUCKLE
WITCH HOBBLE

TOTAL DENSITY

HERB SPECIES
MAIANTHEMUM
CLINTON/A
SHINY CLUBMOSS
WOODS FERN
WOODSORREL
FALSE MITERWORT

Fig. 6. The herb data from each transect were transcribed from the field
sheets to Graphical Analysis Form 2.

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38

Roosevelt Wildlife Bulletin

B. Graphical and Mathematical Analysis of Underbrush and
Herbs. Since the time allotted to this study precluded a mathe-
matical analysis of the distribution of the mature dominant forest
vegetation, the greater part of the work was in reference to the
lesser vegetation. This included herbs, shrubs and reproduction of
dominants up to one inch in diameter.

The analysis of the data in reference to shrubs and herbs was
continued on Graphical Analysis Form 3, a modification of which is
included in the text in the discussion of each of the ten transitions
studied. All of the shrub and herb data in reference to each type
of edge were summarized on these forms. Each shaded line of
squares opposite each species represents an individual transect. The
extent of the shading indicates the length of the transects, and the
plot numbers at the top of the sheet indicate the extent of the tran-
sect in each direction from the center of transition. The heavy dots
represent the distribution and density of the species across the tran-
sect. The transects were so placed on the sheet that their centers of
transition were vertically aligned. This procedure resulted in a
graphical summation of the data, and in some cases it revealed con-
centrations of individuals at particular points across the transition
while in other cases an ungraduated distribution was apparent. The
range of each species across the transition as well as its relative
abundance at different parts of the transition can be derived from
this graphical summation.

At this point the analysis was converted from the graphical to
a mathematical basis.* (See Table 3). In the mathematical analysis
the transition was divided into three parts : ( 1 ) the data from the
plots at the end near type "A"; (2) the data from the plots at the
end near type "B" ; and (3) the data from the three underbrush-herb
plots at the center of transition. This arrangement made it possible
to compare the average cover value of each species in each of the
three parts of the transition. As an example let us consider the oc-
currence of witch hobble across the transition between spruce flat
and northern hardwood (Table 4). The names of the parts of the
transition, plus the number of plots in each part, are entered in the
first line across the top of the mathematical analysis sheet. The fig-
ures in reference to witch hobble show that in the 135 plots on the
spruce flat side of the transition, this species occurred 23 times with
a density of trace, 30 times with a densit}' of one. seven times with
I

* It was assumed that if actual square-foot coverage were measured on each
of a large number of densities for each class the average square-foot coverage
would be midway between the upper and lower limit in each instance.

39

TABLE 3. DERIVATION OF FACTORS FOR THE CON-
VERSION OF GRAPHICAL DATA TO A MATHEMA-
TICAL BASIS

UNDERBRUSH
Size of Plot— 6.6' Square or 43.56 Sq. Ft. (0.001 Acre)

Densities

Part of Plot
Covered by
Each Density

Range of Area

Covered by
Each Density

Average Area*

Covered by
Each Density

•

T

0 to 4 sq. ft.

0 to 4 sq. ft.

2.0 sq. ft.

• •

1

4 sq. ft. to 1/3 plot

4 to 14.52 sq. ft.

9.26 sq. ft.

• ••

2

1/3 plot to 2/3 plot

14.52 to 29.04 sq. ft.

21.78 sq. ft.

• •••

3

2/3 plot to 3/3 plot

29.09 to 43.56 sq. ft.

36.30 sq. ft.

HERBS

Size of Plot— 3.3' Square or 10.89 Sq. Ft. (0.00025 Acre)

Densities

Part of Plot
Covered by
Each Density

Range of Area
Covered by
Each Density

Average Area*
Covered by
Each Density

•

T

0 to 1 sq. ft.

0 to 1 sq. ft.

0.5 sq. ft.

• •

1

1 sq. ft. to 1/3 plot

1 to 3.63 sq. ft.

2.32 sq. ft.

• ••

2

1/3 plot to 2/3 plot

3.63 to 7.26 sq. ft.

5.45 sq. ft.

• •••

3

2/3 plot to 3/3 plot

7.26 to 10.89 sq. ft.

9.08 sq. ft.

NOTE — The figures in the last column are the "Factors" used in Table 4.

40

Roosevelt Wildlife Bulletin

a density of two and six times with a density of three. The com-
puted average cover values of each one of these densities are Hsted
as being 2.00, 9.26, 21.78, and 36.30 .square feet, respectively (see
Table 3. Derivation of Factors for Conversion of Graphical Data
to Mathematical Basis). The total coverage of all the witch hobble
in the 135 plots at the spruce flat side of this edge may be determined
by multiplying the frequency of occurrence of each density by the
average area covered by each density and adding the products. The
660 square feet of witch hobble cover in 135 mil-acre plots represents
12 percent of the ground surface as being covered by witch hobble.
The relative percent of composition of the shrubs at the various parts
of the transition can be determined by adding the cover values for
each species and then dividing each by the total. It can be deter-
mined whether a species occurs characteristically in relatively large,
dense patches, or in sparse patches ; or whether it is usually found as
scattered individual plants, by consulting the frequency of occurrence
in the various densities. Furthermore, it can be determined whether
a species is more abundant at one part of the transition than at the
others. This method of analysis shows in mathematical terms how
much of each of these species of food or cover is present at any de-
sired locality or type of locality.*

C. Mathematical Analysis of Arborescent Reproduction. The

data in reference to arborescent reproduction were computed on a
mathematical basis only, because a graphical analysis would have
been too unwieldy. The formula used to derive the number of seed-
lings per acre for each section (tree plot) of the transition was as
follows :

The factor of 1,000 was used because the plots were 1/1,000
acre in size. When the computation was completed the figures (num-
ber of seedlings per acre) were transcribed to mathematical analysis
forms for each of the ten edges. These are presented in the text.

Graphical and mathematical analyses of this sort were made of
each of ten kinds of forest type transitions, and their relative wild-

* Printing limitations precluded the presentation of all of the ten mathematical
analysis sheets ; however, Table 4 is presented as a representative. Copies of
all ten sheets are on file at the Roosevelt Station for purposes of reference. The
tables headed "Summary of Principal Features . . ." in the discussion of each
edge were derived as a summary of each mathematical analysis sheet.

total number of seedlings recorded
in all of tree plots at that
distance from the center

number of seedlings =
per acre

X 1000

total number of plots at that
distance from the center

TABLE 4. .MATHEMATICAL ANALYSIS OF THE SHRUBS AXD HERBS ACROSS THE EDGE
BETWEEN SPRUCE FLAT AND NORTHERN HARDWOOD

Factor9

Densities and
coverages

Spruce Flat— 135 plots

Per
cent
of

total

Area
covered
(sq. ft.)

Area
covered
(sq. ft.)

21,78

.Area
covered
(sq. ft.)

covered
(sq. ft.)

Sq. ft.

cover-
age
per
acre

Center— 45 plots

Per
cent
of
total

2.00

Area
covered
(sq. ft.)

covered
(sq. ft.)

Area
covered
(sq. ft.)

36.30

Area
covered
(aq. ft.)

Sq. ft.
cover-

Northern Hardwood— 147 plots

Per
cent

Area
covered
(sq. ft.)

(sq. ft.)

Area
covered
(sq. ft.)

Area
covered
(sq. ft.)

SHRUBS

Alder

Blueberry

Elderberry

Iliizclnut

, miniiitain.

liiispl.c'iTv, ihvarf..
Ui.spl>i'iTy, rwl ..

Sweet gale

Witch hobble

1.1
1,0

'i.6
.4
8,1
.1

isi

25,8
.6
43.0

6.00

s'.bo

6-00
28.00
2.00

'e'.oo

52.00

9.26
9.26

18.52
lbi.86

9.60
138.90

9.26
277.80

87,12
152.46

108.90
72.60

1,029
113

197
44

962
15

2.183
3.082

6 ;217.80

Total.

11.7
23.7

12 00
24.00

37,04
83.34

21
21

87.12

6,00

2».ix)

2,00
.54 00
54.00

21.78

Factors

Aster, sp

Bedstraw

Clintonia

Club-moss, shiny. . .

Cucumber root

Dogwood

Fern, beech

Kern, bracken

Fern. New York , . .

Fern, oak

Fern, sensitive

Fern , wood

(loldenrod, hairy. . .
: (li)l(lenrod, wjods. .

(loldtbread

(jv-dss sp.

.lack-in-the-pulpit..

j Maianthemuvi

I .Miterwort, false . . .

Nettli'

Nightshade

Partridge berry

Piroh

Hjiitmcuvi

I Sarsaparilla

I Sedge sp

SnowbeiTy

Solomon's seal

Starflower

Trillium, painted .
I Trillium, red

Twisted stalk

Violet

Violet, false

Wintergreen

Wood sorrel

Total

.4

.6
2.4

.7
1.2
4.3
1.1

.1

.6
1.0

.4
31.3

.6
1.0
1.4

.4

,9
1.:
1 2
.2
.2
.1
1,0
1.,
8.0
.1
.1
1.4
3
.1
.5
2.7
1.4
.4
30.1

99 S

3.50
14.00
2.00
7.50
13.00
2.00
0-50
1.00
1.50
2.50
23.50
1,50
1.50
3.50
2.50

21 ,50
4.00
2.50
1.50
1.00
0.50
1.50
6.00
7.00
0.50
0.50
8.50
2.00
0,50
3.00
6.50
4.00

20.50

2-32
4,64

81.20
2,32
4,64
4.64

4.64
4,64
4,64

9.20
4.64
2.32
85,84

0.50

2.32

5-45

9-08

0..50

2.32

5,45

9.08

69

89

.7

8

4.00

109

101

.6

2

1,00

178

1 .6

13

6 .50

1

2,32

240

415

1,3

4

2 00

340

1.5

12

6 00

1

2-32

226

128

2.4

3

1 .50

1

2 32

267

2-9

27

13.50

1

2 -32

430

222

1,9

6

3,00

473

.6

2

1-00

1

2 32

|91

753

3.4

6

3,00

1

2.32

44

-8

4

2.00

1

2-32

117

197

.3

1

0.50

15
98

5-0

1.00

1

2.32

5.45

is.ie

773

182

' .9

3

1.50

' ' 133

.4

1

2,32

63

74

.1

'0.5b

13

5,, 524

33.1

17

8.50

15

34.80

'9.08

4^656

34.0

45

22.50

48

lii.36

49.65

4,970

113

182

'i.8

0.50

'i

2-32

251

"'.8

2.00

i

2.32

117

241

.3

0.50

44

.2

1.00

27

74

.9

3

1..50

133

.6

3-50

95

.6

2

1.00

89

2-0

12

6-00

2

4^64

289

159

4.6

10

5-00

1

2.32

651

.9

10

5-00

135

325

5,9

14

7,00

1

2.32

828

9-2

57

28.50

'9

20^8

1,344

212

44

3.4

'5;45

484

"".8

2.00

2.32

117

30

.3

1

0.50

44

.2

1-00

27

15

182

315

' .3

1

0-50

'44

1.3

4-50

2-32

"m

1.421

.3

1

0.50

44

1-0

5-50

150

15

.9

4 64

126

15

"'.3

1

0.50

44

1.3

4-50

2-32

186

252

.9

3

1.50

133

1.6

8-50

236

59

.3

1

0.50

44

.6

2

1-00

'2.32

90

15

.3

1

0.50

44

.4

4

2.00

54

89

! .6

2

1.00

89

1.1

12

6.00

164

468

; 3.4

1

6

3,00

I

'2^32

473

4.5

33

16. .50

2.32

5.45

660

256

2

1,00

89

1.9

7

3-50

3

6.96

284

69
5.304

30.9

ii

5.50

14

32.48

2

io^go

4.345

22^8

oi

28.50

28

64.96

2

io.bb

2

is. 16

3^333

17.636

9!) 6

14.0.53

69-7

14,6.52

Edge Effect of the Lesser Vegetation

41

life habitat values in reference to the arborescent reproduction,
underbrusli and herbs was estimated in this manner. For example,
it was found on this basis that the edge between spruce flat and
northern hardwood provided a maximum variety of underbrush and
herbs, for here there were 12 species of shrubs and 36 species of
herbs. The total shrub cover was about 13,000 square feet per acre
and that of the herbs about 16,000. In other words, about 30 per-
cent of the ground surface was covered with shrubs and 37 percent
with herbs. Several valuable fruit-bearing shrubs as a group were
more abundant at this edge than at any other part of the forest.
Among these were blueberry, elderberry, mountain holly, honeysuckle,
red raspberry and dwarf raspberry as well as several species of
fruit-bearing and succulent green herbs. It was found that both
summer and winter cover were abundant. By referring to the sum-
marized mathematical analysis tables, it is possible to determine not
only what species of plants characterize each edge, but also the
relative amounts of each species at each portion of each edge. The
figures are given as average cover values, as percent of total compo-
sition, as well as square foot coverage per acre.

V. ANALYSIS OF DATA— EDGE COMPOSITION WITH
SPECIAL REFERENCE TO GROUSE AND DEER

Rather than involve the reader in a detailed recapitulation of the
data, the author has attempted to first briefly summarize and inter-
pret the more easily recognized vegetative characteristics of each
edge and then to relate the importance of these features to the wild-
life concerned.

The data are interpreted with special reference to the require-
ments of rufifed grouse and deer, since these are the two principal
herbivorous game species on the Huntington Forest. Deer are in-
cluded not because of any cruising radius limitations, but rather be-
cause examination of distribution data revealed that they are more
frequently observed near the edge between certain forest types than
in the centers of large blocks of single types. They apparently ex-
hibit a certain degree of edge effect in spite of their relatively un-
limited cruising radius and some of the reasons for this will become
evident in the discussion which follows. Since the habitat require-
ments of deer and grouse have not been completely determined for
this region, most of this discussion must be based on personal ob-
servations made locally, and habitat requirements of these species as
determined by others in various localities. As the specific habitat
requirements of the local species of wildlife are more fullv revealed,

42

Roosevelt IVildlife Bulletin

the data gathered in the course of this study can be reinterpreted in
the Hght of these findings. Observations and conclusions, espt-ciall)-
in reference to the distribution of deer and grouse in relation to th.
vegetation, are those of the author, and although based in part on the
work of the Roosevelt Station, do not represent conclusions reached
by that organization.

Certain limitations of a study of this type must be recognized.
Among thesf may be listed the impossibility of gathering all of the
data at a time when all of the many species of seasonal herbs are at
the peak of abundance. However, it is believed that the relative
amounts in the different forest types are accurately portrayed. An-
other condition difficult to interpret is the true value of certain very
scantily distributed plant species which are considered by some to
constitute favorite foods. Such fruits as creeping snowberry, par-
tridge berry, and wintergreen are present in very small amounts, but
it is impossible to attach the proper wildlife value to them since in-
tensive studies of local food habits of grouse have not b"en completed.
It must be taken for granted that although these species compose a
very small proportion of the vegetation they may constitute a very
important part of the diet of grouse.

The four seasons of the year, as mentioned below, do not con-
form to calendar seasons, but are set up on the basis of weather
conditions and the condition of the vegetation. Cakndar equivalents
and definitions are as follows: spring — April 1 to June 1, the period
after the snow has left, new leaves are being formed and plants start
to grow; summer — June 1 to October 1, deciduous plants are in leaf,
maximum cover is afforded by hardwoods, summer fruits ripen and
fall; fall — October 1 to December 1, period of leaf fall and bare
ground prior to the permanent snow cover of winter; winter — De-
cember 1 to April 1, period of deep snow, deciduous woody plants
devoid of foliage, minimum cover afforded by hardwoods.

The tables summarizing the more abundant vegetative features
of value to wildlife were derived from the graphical and mathematical
analysis forms at the end of each edge discussion. The classifica-
tions as to presence in "abundance" and "medium to small amounts"
were based on the arbitrary scale presented in Table 5. "Scarce"
species were not listed in these tables.

A. The Edge Between Bracken Fern and Aspen. The
bracken fern and aspen types, which developed after a post-logging
fire in 1903, are restricted to the northeast corner of the Check Area.
Most of the burned area already has progressed to the aspen and
second growth hardwood stages but portions still remain with a
cover dominated by bracken fern. It may be of significance to note

Edge Effect of the Lesser Vegetation 43

TABLE 5. NUMERICAL EQUIVALENTS OF GENERAL
ABUNDANCE GROUPS USED IN TABLES SUMMARIZ-
ING THE PRINCIPAL VEGETATIVE FEATURES OF
VALUE TO GROUSE AND DEER

Form of
Vegetation

Size Class

Abundant

Moderate

Reproduction

(expressed in
numbers per
acre)

24" high to r'diam.
6" high to 24" high
0" high to 6" high

250 or more
1 , 000 or more
5 , 000 or more

50 to 250
250 to 1,000
1,000 to 5,000

Shrubs

10,000 or more

1,000 to 10,000

(in square feet of
cover per acre)

Herbs*

5 , 000 or more

500 to 5,000

(in square feet
of cover per
acre)

* In the case of shrubs and herbs these values were occasionally modified by
a consideration of densities and frequencies.

that the great bulk of bracken types are along streams which, until
recently, were used by beaver ; i.e., it would appear that beaver have
had some part in the continuance of this stage by their intermittent
removal of aspen and young hardwoods.

The total area of bracken fern type is 37.6 acres, or 0.9 percent
of the Check Area, while that of aspen is 57.8 acres, or 1.4 percent
of the total. The study of this edge is based on eight transects con-
sisting of 186 underbrush and herb plots. Seventy-nine of these plots
were in the bracken fern type, 56 at the center of transition, and 51
in the aspen type.* In most cases this was a relatively abrupt transi-
tion. The transects averaged 250 feet in length but in most cases the
effective transition occurred over a zone averaging 225 feet or less.

1. Vegetative Characteristics. Although the species composition
of the lesser vegetation did not vary markedly across the transition,
there was a definite change in dominants among the underbrush
species and arborescent reproduction. The most abundant species of
reproduction on the bracken side was trembling aspen, but seedlings
of sugar maple and red maple were much more numerous than those

* Occasional discrepancies will be noted in the references to the number of
plots used; these discrepancies are due to the fact that certain transects were
extended across several edges and thus parts of a single transect plotted in
more than one edge.

44

Roosevelt Wildlife Bulletin

of aspen at the aspen side of the edge. This may be taken as demon-
strating the advance of the aspen type into the bracken fern and the
advance of the second growth hardwood into the aspen type. The
fact that seedHngs characteristic of the mature forest occur here is
significant in that it points out that in small denuded areas the cur-
rent nature of the vegetation is controlled not so much by the seed
supply as by site conditions. Although characteristic climax forest
species will continue to seed into this area, the vegetation probably
will not start to assume the nature of the climax forest until site
conditions change sufficiently for these seedlings to develop. The
amount of reproduction at this edge was so small as to be of no
great value to wildlife. Potential browse species were so small and
sparsely distributed as to be almost obliterated by the other vege-
tation in the summer and were covered by snow in the winter. Cover
in the form of conifers was extremely sparse and occurred in widely
separated small patches.

The most abundant species of underbrush on the open bracken
side of the transition were blueberry and red raspberry, while red
raspberry alone dominated the underbrush under the aspen. Bracken
fern was by far the predominant herb species at all parts of the
transition, while hairy goldenrod and dwarf dogwood alternated in
secondary predominance. One species of underbrush and one herb
species revealed a marked preponderance at the center of transition.
The reproduction decreased in size from the open bracken to the
cover of the aspen ; the underbrush coverage was most abundant
under the aspen ; and there was a definitely decreasing trend in the
total amount of herbaceous cover from the open bracken to the aspen.

2. U'ildllfc J 'allies. The increased vegetative complexity of the
entire transition over any of its parts is indicated above, as is the fact
that the width of transition is well within the cruising radius of
grouse. The most conspicuous features of value to wildlife contrib-
uted hv the bracken fern side of the edge were small amounts of late
summer and early fall fruits in the form of blueberry (averaging
3,051 square feet per acre), and small amounts of browse in the form
of aspen reproduction (averaging up to 5,000 stems per acre).

Features contributed by the aspen side of the edge were a ver\-
small amount of sugar maple and striped maple deer browse, and
winter grouse food in the form of aspen buds. Catkins of aspen also
provided abundant earl}- spring food for grouse. A small amount of
early fall fruit was contributed bv the aspen side in the form of false
lily-of-the-valley whicli averaged up to 1,058 square feet per acre at
this side, and tliis species also supplied some early spring greens.

Edge Effect of the Lesser Vegetation

45

Moderate amounts of greens in the form of wood fern and sarsa-
parilla also were provided.

Among the features characteristic of both types may be Hsted
the large amounts of late spring food in the form of bracken fern
"fiddle heads" which are avidly taken by deer. Less conspicuous but
probably also of value was the goldenrod, sedge, and other succulent
herbs. As the season progresses, all of these become more important
as cover than as food, and serve in this capacity from July through
September. However, adequate grouse cover is not available for
the remaining nine months of the year. Goodly amounts of fall fruit
were found on the dwarf dogwood which was abundant throughout
both types. Other fruits, found in smaller quantities in both types,
were : red raspberry, dwarf raspberry, honeysuckle, and hazelnut.
Greens present throughout both types were violet, shiny club-moss,
bedstraw, and browse-size red maple seedlings. (See Fig. 7 and
Tables 6, 7 and 8.)

B. The Edge Between Bracken Fern and Second Growth
Hardwood. The location and origin of these types were included
in the discussion of the previous edge. Although the total area of
the second growth mixed hardwood comprises 88.6 acres (2.2 per-
cent of the Check Area), only about ten acres lie adjacent to bracken
fern type. The analysis of this edge is based on nine transects com-
posed of 295 plots. Of these, 96 were in the open bracken, 36 at the
center of transition, and 133 in second growth hardwood. The aver-
age length of the transects was 350 feet, and this may be taken to
represent the average width of the transition in the lesser vegetation.

1. Vegetative Characteristics. The most abundant species of
arborescent reproduction at this edge, in most cases, was sugar
maple. The fact that it occurred in numbers exceeding 32,000 per
acre under the second growth indicates that this site may possibly be
tending toward a northern hardwood climax rather than mixed hard-
wood. However, this species constitutes potential deer browse only
until winter because most of the seedlings are stagnating at less than
two feet. Red maple seedlings were more abundant under the bracken
fern than were those of aspen, indicating thereby that the aspen
stage may be by-passed in some places. The absence of reproduction
of beech, striped and mountain maples, and white ash in the bracken
area as opposed to their presence under the cover of the young hard-
wood is probal)ly an indication of the inability of these species to
become established in pioneer societies of this type.

The more important features relative to the shrubs and herbs
may be summarized as follows : Although the same species prevailed

TRANSITION -

OPEN BRACKEN

CENTER ASPEN

TREE PLOT NOS.

3

2

1

1

B

■<UDn Q HtHB PLOT NOS

9

6

4

3

2

1

t

2

3

1,

BLUEBERRY

■

»

•»
V

r"

-

-

2

HAZELNUT

•

3

HONEYSUCKLE

1

*

•

t

-1

-

-

4

MEADOW SWEET

5

OSIER. RED

RASPBERRY, DWARF

—
**

1

•

J,

6

•

1

•

1 !b

I.'

IT

w

■

7

RASPBERRY, RED

r

1

1
t

i*

L*

L

a

WILLOW

9

WITCH HOBBLE

•

•

— 1

1.

ASTER. SP

BEDSTRAW

*-

3"

•

t

r
1

•

2

3

CLINTONIA

•

— 1

-

4

CLUBMOSS, SHINY
'

•

5.

DOGWOOD, DWARF

<%-

f-

i

h

9

a

h

f

i

6

FERN, BRACKEN

Bi

s

ca

1!

i

M

m

1

7

FERN, HAY-SCENTED

•

e

FERN, NEW YORK

— 1

~

9

FERN, WOOD

-1

. r

^7

10

GOLDENROD, HAIRY

ft

t
r

L

K

IT

.

II

GOLDENROO, WOODS

V

i

12

GOLDTHREAD

r-

13

GRASS, SP

*'

14 ' HIERACIUM

V—

15 JACK-IN-THE- PULPIT

1

16. MAiaiVTHeMUM CAN.

{

17

MITERWORT. FALSE

H

#

$

1

-

18 1 SARSAPARILLA

•

-

•

19

SEDGE SP

f

IT

1

I

1

i-

•

•

i

1

T-

20

STARFLOWER

*

»

21.

STRAWBERRY SP

22.

THISTLE. CANADA

Fig. 7. Graphical an
shrubs and herbs aero
Fern

al>sis of the di:
ss the edge betw
and .\spen.

Uribntion of
een Bracken

[46]

Edge Effect of the Lesser J'cgetatioii

47

TRANSITION -

OPEN BRACKEN

CENTER

ASPEN

TREE PLOT NOS^

3

2

1

1

BRUSH a HERB PLOT NOS

9

6

7

6

3

2

1

1

2

3

25

TDII 1 II lUi DAIMTPn

1

m

24.

TRILLIUM. RED

f

25

TWIN FLOWER

26

VIOLET SP

E

}
i

{

!

•

27.

VIOLET, FALSE

w

-

28.

WINTERGREEN

29

WOODSORREL

Fig. 7 (Concluded)

in dominance across the edge they dropped considerably in abundance
from bracken fern into second growth hardwood. The characteristic
species of imderbrush were hazelnut, dwarf raspberry, red raspberry,
honeysuckle, blueberry, and witch hobble. The characteristic species
of herbs under the second growth were wood fern, false lily-of-the-
valley, and wood sorrel ; species characteristic of the bracken fern
type were bracken fern, hairy goldenrod, dwarf dogwood, grasses,
and sedges. Species found in moderate abundance in both types were
false miterwort, starflower, bedstraw, and violets. It is obvious that
the total number of species across the transition was greater than
that at any part. The greatest concentration of total underbrush
cover was at the center of transition and there was a definitely de-
creasing trend in the total amount of herb cover from the open
bracken to the wooded side of the transition.

2. Wildlife Values. Features of importance to wildlife con-
tributed by each portion of the edge are listed in Table 9. This
edge, in spite of the increased diversity of vegetation, produced wild-
life foods during only certain seasons and provided practically no
cover. The bracken fern side provided large amounts of late spring
greens ("fiddle heads", grass and goldenrod), and excellent summer
ground cover. It also provided small amounts of summer fruit
(blueberry) and fairly large amounts of early fall fruit (dwarf dog-
wood). The only species of arborescent browse occurring in signifi-
cant quantities was red maple, but this probably was not sufficient to
maintain even a small population of deer throughout the winter.

The principal feature provided for grouse on the second growth
side was hardwood buds for winter food, and four species of herbs
for spring and summer food (with wood fern being available until

■a

O

3

4?

C TO
t. o ?

O ^

C3

a- 2 S - —
rt E t p =

tA ir, > (/)

o o s: 5.2

_ o

2 o s

X3

a

XI
T3

X <f

X o

<CQU.

>
o
U

X

3

P o

.H X

148]

49

TABLE 7. MATHEMATICAL ANALYSIS OF THE ARBOR-
ESCENT REPRODUCTION ACROSS THE EDGE BE-
TWEEN BRACKEN FERN AND ASPEN EXPRESSED
IN NUMBERS PER ACRE

Species

Size class*

Bracken Fern

Center

Aspen

Plot
No. 3

Plot
No. 2

Plot
No. 1

I lot
No. 1

24 diam.
6 "-24"
0"- 6"

167
3,668
2,166

167
5,330
2,166

91
1,227
182

250
1,583
333

45
1,044
45

Black cherry. . .

24"- 1" diam.
6 "-24"
0"- 6"

42
85
375

333

546
227

45
546

Sugar maple. . .

24"- 1" diam.
6 "-24"
0"- 6"

136

127
958

91

4,315

Red maple

24"- 1" diam.
6 "-24"
0"- 6"

-

408
500

85
2,090

91
2,590

Beech

24"- 1" diam.
6 "-24"
0"- 6"

45
136

136
364
318

167

Shadbush

24"- 1" diam.
6 "-24"
0"- 6"

42
125
136

167

91
42

Striped maple..

24"- 1" diam.
6 "-24"
0"- 6"

85
292

1,181

Yellow birch . . .

24"- 1" diam.
6 "-24"
0"- 6"

85

182

182

Red spruce.

24"- 1" diam.
6 "-24"
0"- 6"

91

45
45

Balsam

24"- 1" diam.
6 "-24"
0"- 6"

45

84

* All figures refer to height unless followed by term "diam." which refers to
diameter.

50
TA

No.

1

2
3
4
5
6
7
8
9

1
2
3
4
5
6
7
8
9
10
11
12
13
14
15
16
17
18
19
20
21
22
23
24
25
26
27
28
29

?LE 8. SUMMARY OF MATHEMATICAL ANALYSIS
OF THE SHRUBS AND HERBS ACROSS THE EDGE
BETWEEN BRACKEN FERN AND ASPEN

Open Bracken
34 plots

Species

Per
cent

of
Total

Sq. Ft.

Coverage
per
Acre

Center
24 plots

Per
cent

of
Total

Sq. Ft.

Coverage
per
Acre

Aspen
22 plots

Per
cent

of
Total

SHRUBS

Blueberry

Hazelnut

Honeysuckle

Meadowsweet ....

Osier, red

Raspberry', dwarf.
Raspberry, red . . .

Willow

Witch hobble

Total .

43

17
29
1
1

3,051

294
334

1,219
2,042
59
59

7
7
31

i

7
47

417
473
2,012

""83
417
3,033

6
17
65

7,058

6,435

HERBS

Aster sp

Bedstraw

Clintonia

Club- moss, shiny. ,
Dogwood, dwarf. .

Fern, bracken

Fern, hay-scented.
Fern, New York. .

Fern , wood

Goldenrod, hairy. ,
Golden rod, woods.

Goldthread

Grass sp

Hieracium

Jack-in-the-pulpit .

Maianthemum

Miterwort, false. . .

Sarsaparilla

Sedge sp

Starflower

Strawberry sp

Thistle, Canada. . .
Trillium, painted..

Trillium, red

Twin flower

Violet sp

Violet, false

Wintergreen

Wood sorrel

.1
.8
.1
.3

7.0
46.3
.1

" '. 'l
27.7

' .2
.8
.4

.3
2.4
.3
5.9
.4
.2
.1

Total .

59
412
59
176
3,706
24,674
59

"lis

14,716

"lis

449
235

" 176
1,268
176
3,042
235
118
59

59
2,668
59
546
59

53 , 246

8

417

3

167

3

0

1,462

16

7

8,137

54

0

26,283

2

3

1,107

9

5

4,635

2

83

i

2

583

2

83

3

4

1,680

1

4

666

2

83

1

0

500

2

5

1,220

.2
.2

2". 7
" .2

83
83

1,303

83

48,658

2.0

" ' .2
13.5
47.5

1.5
17.9
.5
.2
.5
.2
1
6

Edge Effect of the Lesser Vegetation

51

covered by snow). Fruit production by honeysuckle and false lily-
of-the-valley has been notoriously low on the Huntington, therefore
moderate quantities of these species do not imply moderate amounts
of fruits.

A fairly large variety of food species was common to both types.
Bedstraw and violet provided early spring greens, supplemented later
in the season by starflower and false miterwort. This last species
over-winters in a green condition and can be used in the fall until
covered by snow. Red and dwarf raspberries provide a limited
amount of summer fruit, and hazelnut a very small amount of early
fall mast. No late fall and early winter fruits were found. (See
Fig. 8 and Tables 9, 10 and 11.)

C. The Edge Between Second Growth Hardwood and Ma-
ture Northern Hardwood. The ten transects across this edge
were taken in the northeast part of the Check Area between the
previously burned area and an apparently unburned elevation. The
data were derived from 344 underbrush and herb plots, of which
138 were in the second growth, 60 at the center of transition, and
146 in the mature hardwood. The average length of the transects
was 425 feet, but the data indicate that the actual transition occurred
within a zone of 225 feet.

1. Vegetative Characteristics. Sugar maple was by far the most
abundant species of arborescent reproduction. x\ll three size classes
of this species exhibited a gradual increase from the second growth
toward the mature northern hardwood. Striped maple was the next
most abundant species but it showed a reverse relative abundance
tendency, i.e., much more abundant in all size classes in the second
growth than in the mature stand. Beech reproduction was the third
most abundant. As will be noted throughout, beech reproduction ex-
hibited an abnormal size class distribution in that it was exactly the
reverse of all other species : the largest size classes were the most
abundant, the smallest size classes the least abundant. This condition
poses an interesting question the answer to which may be one or more
of the following: It may indicate relatively rapid growth up to two
feet in height and then a slowing down of growth rate. It also may
indicate a large proportion of root sprouts, which normally grow
very rapidly for the first few years. Then again, relatively few small
individuals possibly may be due to the intensive feeding of wildlife
on mast, while the abundance of larger size classes may be due to
lack of browsing and low mortality due to competition. Reproduc-
tion of red maple was somewhat less than that of beech and it showed

"TOEE PLOT NOS.

4

3

2

1

2

3

4

BRUSH a HERB PLOT NOS.

12

1 1

10

9

8

7

6

5

Z 1

7

3

g

10

I

12

1

ALDER

, ....

2

BLUEBERRY

_l

3

DOGWOOD. ALT. LEAR

1

4

ELDERBERRY

—

1—

5

HAZELNUT

1**

•

n

T
1,
1

•

-

6

HOLLY, MOUNTAIN

7

HONEYSUCKLE

-

-

—

—

r

1—

1-1

—

1—

e

MEADOW SWEET

•

-

—

—

—

RAISIN, WILD

9

10

Raspberry, dwarf

-

a

1-

:

»

*■

_

1 1

Raspberry red

tt

f

t

r

1.

t

r

{

•

:

•

*

1

1

12

virgin's BOWER

*

«

F-

— i

1 3

WITCH HOBBLE

•

..

•

•

•

•

1

tt

♦

•

#

r

:

f

{

i

■

—

14

BEDSTRAW

1

*

— 1

15

CLINTONIA

»

•

— 1
•

16

CUCUMBER ROOT

•

t

*

— i

17

DOGWOOD DWARF

t

I*

m-

t

I

:

r

♦

•

•

a

r

1

—

U-

— 1

18

FERN, BEECH

—

—'

L

**

*

r

:

19

FERN, BRACKEN

20

FERN, NEW YORK

21

FERN, OAK

—

22

FERN, WOOD

•

b

1

— .

— ,

—

23

utji_ucnnvu, riAinT

m

I

f

£

1

•

1

I

24

L-

GOLDTHREAD

«

r

25

.1

26

GRASS SP

r

T

•

»

r

27

MERACIUM

t

-

Fig. 8, Graphical aiial\sis of the distrihuti(.)n ui shruh.-; and herbs across
the edge between Bracken Fern and Second Growth.

[52]

Ed(ic Effect of the Lesser J'cf/etatioii

S3

TRANSITION-

OPEN BRACKEN

CENTER

SECOND GROWTH

TREE PLOT NOS

4

3

2

1

1

2

3

4

BRUSH a HERB PLOT NOS

lalii

10

9

8

7

6

5

4

3

2

1

1

2

3

4

5

6

7

8

9

10

1 1

12

28

t

X

r

*•
*

♦

•

*

MAIANTHEMUM CAN

Ah

V

*

s

A-

29

MITERWORT. FALSE

•
*

t

a

r
*
«

*

»

•

A.

i.

30

NIGHTSHADE

31

PIROLA

32

SARSARARILLA

»

*•

»

*

t

33

SEDGE SR.

i

r

1
*

1

X

t

i

—

i

r

1

•

•

•

•

34

SOLOMON'S SEAL

-J

\

1

t

t

*

35

STARFLOWER

*

■

36

STRAWBERRY

37

THISTLE. CANADA

*

•

38

TRILLIUM, PAINTED

39

TWIN FLOWER

40

TWISTED STALK

♦

t

t

fc

1

r

r

4!

VIOLET SR

{

t

X..

S

s

42

VIOLET. FALSE

43

WINTERGREEN

**

*

44

WOODSORREL

Fig. 8 (Concluded)

a marked increase in abundance toward tlie second growth side of
the edge. Deer I:)rowsing probably has intensified the diminution of
larger size classes normally attributable to competition. The rela-
tively small amount of yellow birch reproduction probably was due
to the closed canopy although deer browsing, too, may be indicated.
It was significant to note that small amounts of trembling aspen re-
production up to two feet in height were found as far as the center
of transition. On the other hand, white ash was mostly confined to
the mature hardwood side of the edge. Species occurring with too
low a density to show any site preference were : black cherry, spruce,
balsam, mountain maple, shadbush, and hop hornbeam.

I-* <

< w

H

w
<o
>Q

<; w

<^
wo

[t, I— I

o w

ffi ffi

^pqQ
P W _

cK>0

r 1 O U

<

rt; O

o) O

rt,>.

o

3

o 2 «

o E ^

? , o

3. c/)

act; « o J

« « 3 u > _

•a

o

= i

u. ?

C ~ u

4J C

-a U X i/

o a X

2 > 2 o

2^

J3 V
(J tao
3 T3

CQc/2

C c8

C cs

[34]

o
O

CU V

O

vO m O
»-i ro lo

IT)

r*^ p^

0.

00 >^ O

o o

• 00 O

3 O

lO vO
t 00 0\

i.4

00 o

T-l PT)

o

IT) CN

OO

P^ Ov
O 00

• PO

• PO

PD

00 O
^ PO

Cl. yr

PO

O

vO PO

PC
PO

^2

PO o
pn o o

PD O O

C/5

T-. O

^1 ^1 ^1
^ o o

O

^1 ^1 ^1
^ o o

>-H O

^ o o

.2
-5

^ ^ o

^ MS O

E
.2
■5

-H Tj" O

J ^1
o o

.2
-5

a

be
3
C/3

?3

03

[55]

rn 00

O 0\

"T! 00

00 rl-

-H Tj- VO

vC O

fN

^ o

^1 ^1 ^1

o o

^ -c

J -' ='

« O

C/2

-3

cs

[56]

TABLE 11. SL AIMARY OF MATHEAIATICAL ANALYSIS
OF THE SHRUBS AND HERBS ACROSS THE EDGE
BETWEEN BRACKEN FERN AND SECOND GROWTH

Open Bracken
40 plots

Densities and Co\erages

Fer
cent

of
Total

Sq. Ft
Coverage
per
Acre

Center
27 plots

Per
cent

of
Total

Sq. Ft.

Coverage
per
Acre

Second Growth
57 plots

Per
cent

of
Total

SHRUBS

Alder

Blueberry

Dogwood, alt. leaf

Elderberry

Hazelnut

Holly, mountain . .

Honeysuckle

Meadowsweet

Raisin, wild

Raspberrs', dwarf.
Raspberry,', red . . . .
Mrgin's bower. . . .
Witch hobble

Total

.4
11.1
.4
.4

13.2
.4
5.1
.4
.9
19. S
39.3
6.0
3.0

50
1 .242
50
50
1,478
50
568
50
100
2.172
4,413
668
334

38.1
19 0

13.5
17.7
.5
11.2

5,221
'2,607

1,855
2,424
74
1,535

22.1

23.3
16.8
3.1
12.6

11,225

13,716

HERBS

Bedstraw

Clintonia

Cucumber root . . . .
Dogwood, dwarf. .

Fern, beech

Fern, bracken

Fern , New York . .

Fern, oak

Fern , wood

Goldenrod, hair>-. .
Goldenrod, woods

Goldthread

Grass sp

Hieraciuni

Maianthemum

Miterwort, false. . .

Nightshade

Pirnla

Sarsaparilla

Sedge sp

Solomon's seal

Starflower

Strawberry

Thistle, Canada. . .
Trillium, painted. .

Twin flower

Twisted stalk

Violet sp

Violet, false

Wintergreen

Wood sorrel

1.0
.6
.1
10.4

47.9

'l
. 7
9.8
.1
.3
9
,3
6
0

13

Total.

650
400
50
6 , 390

29^520

" "so

450
6,058
50

200
8,575

200

350
1,246

' ' '.SO
300
4,057
50
500
50
100
50
50
150
1,677
350

61,573

2.6
1.5
.4
12.0

35' 8

.t
3.5
2.9
.6
.6
3.9

'2^9
4.5

5.2
7.5

'5^7

A
.2

4.6
2.7

518
158
4, 178

12^433

"222
1,206
1,010
222
222
1,354

'loio

1,550

1 ,791
2,606

L994

148
74

1,603
862

640

34,689

6.2
2.6
4.2
4.4
1.7
14.1
.5

13

6.1

3.7
5.7

'4'2

'3

12

1.0
.3
4.7

[57]

58

Roosevelt Wildlife Bulletin

The data in reference to shrubs and herbs may be summarized
as follows : In general, the same species were dominant across the
transition, but there was some shift in relative dominance. The
underbrush was composed principally of witch hobble and honey-
suckle. The herb species consistently most abundant across the tran-
sition were wood fern, false miterwort, wood sorrel, and violet. Two
species of underbrush and eleven herb species occurred with greatest
abundance at the second growth side while seven herb species and
witch hobble indicated a preference for the mature hardwood side.
The total number of herbaceous species across the transition was
greater than at any part of the transition, but in the case of the
underbrush the greatest number of species was found on the second
growth side (Table 12).

2. Wildlife Values. The more obvious and abundant features
of value to wildlife provided by each of the two adjacent types are
summarized in Table 12. Striped maple browse was more abundant
on the second growth side but since most of it was less than two
feet in height it could be used only in the early part of the winter.
The same could be said for red maple, except that it was only about
one-tenth as abundant. The only other foods confined to this type
were early spring greens in the form of violet and bedstraw.

The mature hardwood side of the edge provided large numbers
of young beech from two feet tall to one inch in diameter, the only
value of v/hich, however, might be a small amount of fall and winter
cover. Mature beech, on the other hand, served as an excellent po-
tential source of mast. Shiny club-moss was characteristic of this
type, but the extent to which it is used by deer and grouse is un-
known even though it remains green all winter. Early spring greens
in the form of dogtooth violet and spring beauty were especially
abundant in this type.

Present in both types were hardwood buds for winter grouse
food and sugar maple browse for deer. Witch hobble provided only
scant cover and browse, due to its occurrence in such small densities
(see graphical analysis form, Fig. 9). Honeysuckle, too, provided
very little browse and even less fruit, although present throughout
and fairly abundant. Overwintering greens common to both types
were wood fern, false miterwort, and wood sorrel, while goldenrod
constituted a fairly abundant spring food.

It would appear, therefore, that in some respects this edge had
even less to offer than the previously discussed two, for there were
fewer fleshy fruit-bearers than at those edges. Furthermore, the
very scant underbrush in either type provided a minimum of sum-

TABLE 12 SUMMARY OF THE PRINCIPAL FEATURES ADVANTAGEOUS TO DEER AND GROUSE
PROVIDED BY THE LESSER VEGETATION OF THE EDGE BETWEEN SECOND GROWTH AND
NORTHERN HARDWOOD

Features Contributed by
Northern Hardwood

Food

Dogtooth violet
Spring beauty

Shiny club-moss

Cover

Beech

(24" high to
to 1 " diam.)

Beech
(6 "-24")

Features Characteristic of
Both Types

Food

Sugar maple
Hardwood buds

Witch hobble
Honeysuckle
Wood fern
False miterwort
Wood sorrel
Goldenrod

Cover

Features Contributed by
Second Growth

Food

Striped maple

Violet
Bedstraw

Cover

Red maple

Present in abundance. . . .

o
S

.E ^
1^

0) r
a) t
1)

[59]

TRANaTION-

SECOND GROWTH

CENTBR

TREE PLOT NOS

4

3

2

1

1

2

3

4

BRUSH 8 HERB PLOT NOS.

12

1 1

10

9

8

7

6

5

4

3 2

1

2

3

4 5 6

7

8

3

10

II

.

1

1

HAZELNUT

!

•

1

tl

.

—

.

2

HONEYSUCKLE

•

•

3

RAISIN, WILD

•

i

—1—

t.

•

•

s

4

AASPBERfV^, DWARF

•

1

•

t

1

•

— f—
• 1

5

RASPBERRY. RED

•

1

»

1 •

•

•

•

:

t

„

1

r

I:

g

}

r

t

.

•

I

*

•

•

BEDSTRAW

1

1

1

I

1.

U

•

.

•

Q

PI IMTnuiA

1

1

i

t

9

CLUBMOSS, SHINY

*

1

t

H

t

:

♦

10

CUCUMBER ROOT

•

•

•

•

*

•

•

II

DOGWOOD, DWARF

1

•

12

FERN BEECH

lJ

13

FERN. BRACKEN

14

FERN, NEW YORK

—

•*

•

L

*

I.

t

*.

•

•

15

FERN, WOOD

**

a

t

•

16

GOLDENROD HAIRY

•

■*-

•

•

•

17

GOLDENROD. WOODS

■1

*

■

— j —

r

16

GRASS SP.

.1.

•

1

•

•

i

•

•

■

1 9

JACK-IN -THE- PULPIT

—

u

1

20

MAIANTHEUIM CAN.

*

•

!

t
1

-

1

21

MITERWORT, FALSE

1

k

•

•

j

I

i—

■n

*

•

22

PARTRIDGE BERRY

1

23

SARSAPftRILLA

L-

—

—

24

SEDGE SP

1

25

SOLOMON'S SEAL

NORTHERN HAROWOOO

Fig. 9. (Irapliical an;

the edge betw

:il\sis of the distribution of shrubs and herbs across
een Second Growth and Xorthern Hardwood.

160]

Edge Effect of the Lesser Vegetation 61

TRANSITION-

SECOND

GROWTH

CENTER

NORTHERN

HARCWOOO

TREE PLOT NOS.

4

3

2

1

1

2

3

4

BRUSH a HERB PLOT NOS

12

II

10

9

8

7

6

5

4

3

2

1

1

2

3

4

5

6

7

8

9

10

1 1

12

•

*

26

STARFLOWER

•

1

•

•

•

27

TRILLIUM, FftlNTEO

•

28

TRILLIUM, RED

•

29

TWISTED STALK

♦

•

•

{

1

I

r
i

*

k

1

*

*

«

•

30

VIOLET SR

1

!

t

t

2

t

I.

t

1

-»

«

r
#

t

31

WOOOSORREL

♦

4

U

Fig. 9 (Concluded)

mer ground cover for young birds and the relative absence of coni-
fers proved a similar deficiency for other seasons. (See Tables 13
and 14.)

D. The Edge Between Spruce Flat and Mixed Hardvi^ood.

These are two of the three major forest types on the Check Area,
spruce flat occupying 778 acres (19.1 percent) and mixed hardwood
1,212 acres (29.8 percent). The spruce type occupies the lowlands
while the mixed hardwood type usually is topographically between
the northern hardwoods and the spruce types. The separating zone
in certain places becomes quite narrow and some may consider it
arbitrary. However, since the mixed hardwood type appears fairly
stable and widespread and since it constitutes an important distinc-
tion in relation to wildlife, these narrow zones were so indicated on
the type map.

The analysis of the edge between these two types was based on
seven transects composed of 252 plots. The average length of the
transects was 450 feet but the average width of the transition in the
lesser vegetation was about 375 feet.

1. Vegetative Characteristics. Six species of trees were repre-
sented with moderate to abundant amounts of reproduction. As
would be expected, sugar maple was much more abundant on the
mixed hardwood side, but it was fairly abundant even to the center
of the transition. Beech did not reveal a strong preference for either
type near the edge, but this should not be interpreted as indicating
uniform preference for both types because beech is much more
abundant in the mixed hardwood. It does show, however, that beech
does occur in fair abundance on the upper edge of the spruce flat

KW
H CO

CO
CO W

2o

Q <

O It:
W pi!

uo

CO

5^
<<

^§

K Wco

5 pq CQ

<

ro
O vO
O 00

-o
o
o

•a

u.

c«

X

O

Z

^1

■ o o

■ o o

• O 1^

■ o

00 ro ^
1^ r«o ^

O r*: O
O O

\0

o

o

•a
c
o

Oh ^

4-.

C/2

tS O

\0 0^

(vj IT) lO

f»2
C-l O <^

O
O

sO lO

vO

l/^ (v^ rvN
CM

• O

t~- O o
vO lO O
^ f^l O

■ 1^ O

• oo

■ O

,-. Tf< \0

J ^'

o

— Tj- VO

='

o o

cs

« -o

^'

^ nC o

_2
•5

— O

rs

^1 ^1
O

.2
•5

3

in

a:

[62

ro 1^ O
O 1^
CO sO O

■* O O
ir; rr;
O

■ 00 o

<-l

r<^ O "-I

•oo

■ o ^

o 0\
rrj O

. o

• o o

00

in «^

m o <n

E
.2

o

csi o
vO ^ <n

o o <n
o <n
o <n

O I-;

o o

O — '

■ o

■ o
• o

.2

^1 ^1

o o

— Tt" o

^1 ^1 ^1
■* o o

^ o

O O

— vO
CM

^1 ^1

^ o o

o

c
o

[63]

64

TABLE 14. SUMMARY OF MATHEMATICAL ANALYSTS
OF SHRUBS AND HERBS ACROSS THE EDGE BE-
TWEEN SECOND GROWTH AND MATURE NORTH-
ERN HARDWOOD

Densities and Coverages

Second Growth
69 plots

Per
cent

of
Total

Sq. Ft.
Coverage
per
.•\cre

Center
30 plots

Northern
Hardwoofl
73 plots

Per
cent

of
Total

Sq. Ft. Per j Sq. Ft.
Coveragel cent 'Co\erage
per of I f>er
.•\cre Total I Acre

SHRUBS

Hazelnut

Honeysuckle

Raisin, wild

Raspberry, dwarf.
Raspberry, red . . . .
Witch hobble

Total .

1.9
40.6

1.0
16.5
14.0
26.0

58
1.239
29
503
426
793

25.6

13.2
25.6
34.5

645 15.6

333
645
891

13.0
13.4
58.0

3.048

2.514

384

320
329
1,427

2.460

HERBS

Bedstraw

Clintonia

Club-moss, shiny.
Cucumber root . . .
Dogwood, dwarf. .

Fern, beech

Fern, bracken ....
Fern , New York . .

Fern, wood

Go! den rod, hairy.
Goldenrod, wood.

Grass sp

Jack-in-lhe-pulpit.
Maianthemiim ....
Miterwort, false. .
Partridge berry. . .

Sarsaparilla

Sedge sp

Solomon's seal . . . .

Starflower

Trillium, painted.

Trillium, red

Twisted stalk. . . .

\'iolet sp

Wood sorrel

Total.

6.5
.8
1.4
3.4
3.0

667
87
145
348
308

8.2
1.7

20.2
1.1
4.8
4.7
1.4
2.2

14.0
.6
.6
2.8

3'i

.3
.6
15 3
3.4

842
174

2.078
116
501
482
145
232

1,444
58
58
290

319

29
58
1.581
348

10,310

5.3
3.2
7.4
1.1

\.\

\.\
14.1

3.2
3.2
2.1
3.2
20.4
1.1
1.1
3.2

2.'\
1.1
1.1
3.2
12.7
9.2

333 3.1

200 i 1.6

467 I 9.9

67 6.3

67

67
885

200
200
1.S3
200
.285
67
67
200

5.0
8.3

\3.9
3.5

133
67
67
200
800
576

6.281

10

4

3

6

1

0

5

5

i

6

2

6

7

3

12

7

164

82
520
329

264
438

728
182
456

548
192
55

50

136
384
665

5,252

Edge Effect of the Lesser Vegetation

65

types. It will be noted that here too, as discussed in the previous
section, beech had an inverse ratio of size classes. Red spruce re-
production was of course much more abundant on the spruce flat
side of the edge. This increased quantity of spruce reproduction near
the edge may be an indication of an effort by this sjjecies to advance
into the mixed hardwood. Yellow birch revealed noticeably more
reproduction near the edge in .spruce flat than in the center of either
type. This was correlated with a similar distribution of mature in-
dividuals of this .species. Red maple and balsam fir showed a marked
abundance on the spruce side as opposed to the mixed hardwood side.
The paucity of hemlock reproduction poses a serious question rela-
tive to the ability of this species to maintain itself under existing
conditions. The seeds api^ear to have a high germination percent on
disturbed sites such as occur along the truck trail, but germination
is very poor under the mature forest canopy. Small quantities of
arbor vitae on the spruce side usually are associated with certain site
conditions such as lake shores or calcite soils, while characteristically
small amounts of striped maple and mountain maple are found on
the hardwood side.

The principal features relative to the distribution of slunibs and
herbs across the edge may be summarized as follows: Although tliere
were six species of underbrush on the spruce side and only four on
the hardwood side, the total amount of underbrush cover was greater
on the mixed hardwood side. The most abundant species of under-
brusli were witch hobble, honeysuckle, red ra.spberry. and dwarf
rasj)berry. There were 28 species of herbs on the spruce flat side,
but only 20 on the mixed hardwood side. The most abundant herb
species were wood fern, wood sorrel, violet, false lily-of-tlie-valley,
and false miterwort, plus dwarf dogwood and goldthread on the
spruce flat side. Two species of underbrush and eight species of
herbs occurred with greatest abundance at the spruce flat side. Only
one species of underbrush (no herbs) occurred with the greatest
abundance under the mixed hardwood. The spruce flat side of the
transition had a greater variety of underbrush and herbs than either
of the other two ])arts of the transition.

2. JVildlifc Values. The principal vegetative features of impor-
tance to wildlife are summarized in Table 15. This table may give
the erroneous impression that the mixed hardwood provides but little
wildlife food and cover. Actually the amounts of sugar maple browse
and beech mast are very important. A greater variety of necessities
are provided by the spruce flat side of the edge, but the central part
of the table should not be overlooked. Year-round cover is provided

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[66]

Edge Effect of the Lesser Vegetation

67

by spruce and balsam thickets of the spruce type, and also by the
young beech occurring throughout the edge. Browse species restricted
to the spruce type are red maple and balsam fir, while red raspberry
is more abundant here than in the other portions of the edge. The
dwarf dogwood and goldthread of the spruce type provide fall fruits
and overwintering greens, but wood sorrel, violet, and false miter-
wort, which are found in both types, also serve as overwintering and
early spring greens. Spring beauty is present as an early spring
green. The only abundant food-producing species confined to the
mixed hardwood is sugar maple, the seedlings of which provide fall
and winter browse. Mature beech is more abundant in the mixed
hardwood type and serves as an important potential source of mast.

\\'itch hobble is the only species of underbrush abundant in both
types, with somewhat greater amounts in the mixed hardwood : the
data indicate a greater proportion of dense patches of witch hobble
in the mixed hardwood type. This species is the most important
winter deer food, therefore its occurrence in large quantities so close
to good winter cover should be of great importance to deer. Honey-
suckle also is characteristic of both types, but it occurs in such small
scattered amounts as to be of secondary value — both as browse and
early summer fruit.

The relative scarcity of browse-size arborescent reproduction
available to deer is quite apparent. Balsam, arbor vitae, hemlock, and
all maples are practically absent in the two feet high to one inch in
diameter size class, while non-browse species — beech and spruce —
reveal relativeh^ large numbers of individuals in this size class. This
is a pertinent commentary on the size of the deer herd, for it appears
that this species of wildlife is seriously influencing the vegetation
towards a spruce-beech subclimax on the mixed hardwood sites. (See
Table 16.)

It is obvious from the above that not only does this edge possess
greater wildlife potentialities than either type alone, but also that
here, at last, is a near ideal edge for deer because year-round food
and cover are provided. Also, this edge more nearly fulfills the re-
quirements of grouse than any of the preceding because it provides
winter buds, spring greens, summer and early fall fruits, and year-
round cover both on the ground and above. (See Fig. 10 and Tables
16 and 17.)

E. The Edge Betvi^een Spruce Flat and Northern Hardwood.

These two types together comprise over half of the Check Area
(56.05 percent) — northern hardwood covering 1,500 acres and spruce
flat occupying 778 acres. Although the amount of edge directly be-

II

TRANSITION -

SPRUCE FLAT

CENTER

MIXED HARDWOOD

TREE PLOT NOS.

6

5

4

3

2

1

1

2

3

4

5

E

iRUSH 8 HERB PLOT NOS

18

17

6

15

14

13

12

1 1

10

9

8

7

6

5

4

3

2

1

2

3

4

5

6

7

8

9

10

12

,4

15

1

BLUEBERRY

'"

-

2

HAZELNUT

-J

-

-

•

9

u

3

HONEYSUCKLE

•

*•

u

Mi

4

RASPBERRY, DWARF

,,

,

5

RASPBERRY, RED

s

•

r

-

•

.

-

s

"E

-

-

6

WITCH HOBBLE

>

V

•

:

r-i

— 1

7

BEDSTRfllW

•

•

M

6

CLINTONIA

•

I

.

.J

9

CLUBMOSS. SHINY

1 —

•

*

u

10

CUCUMBER ROOT

•

'.

•

•

•

•

1 1

DOGWOOD, DWARF

s;

r

:

r

•

•

H

—

n

12

FERN, BEECH

13

FERN, NEW YORK

rw

-

i

14

FERN, OAK

-

W

IS

FERN, SENSITIVE

H

r=-

r

i>

r
S

16

FERN, WOOD

—

-

I.

s
B

1

5

-

•

1,

-

u

;

r

17

GOLDTHREAD

•

•

•

1

T

•

.

•

18

GRASS, SR

— 1

•

19

JACK-IN-THE-PULPIT

•

20

MAIANTHEMUM CAN.

-r

•

{

•

1

i

21

MITERWORT, FALSE

•

h

•

22

'■-

P

NIGHTSHADE

23

PARTRIDGE BERRY

■

24

HYPERICUM

h

25

SARSARftRILLA

•

•

-

\

26

SEDGE SP

•

„

27

STARFLOWER

r

i

r

•

.

28

TRILLIUM, PAINTED

|_

u

29

TRILLIUM, RED

j—

•

(

30

TWIN FLOWER

•

31

TWISTED STALK

•

ii-

i-

u

32

V

*■

■f ■

h

VIOLET SR

.

w

33

VIOLET, FALSE

••-

1 ■

34

WINTERGREEN

u

35

WOOD SORREL

•

r

e

i

w

t

L

1

I

r

f

L

Fig. 10. Graphical analjsis of tlie distribution of the shrubs and herbs across the edge between

Spruce Flat and Jklixed Hardwood.

68

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[70]

71

TABLE 17. SUMMARY OF MATHEMATICAL ANALYSIS
OF THE SHRUBS AND HERBS ACROSS THE EDGE
BETWEEN SPRUCE FLAT AND MIXED HARDWOOD

Densities and Coverages

Spruce Flat
60 plots

Per
cent

of
Total

Sq. Ft

Coverage
per
Acre

Center
21 plots

Per
cent

of
Total

Sq. Ft.

Coverage
per
Acre

Mixed
Hardwood
46 plots

Per
cent

of
Total

SHRUBS

Blueberry

Hazelnut

Honeysuckle

Raspberry, dwarf.
Raspberry, red . . .
Witch hobble

Total .

156
33
1,203
1,138
1,020
7,522

6.2
4.0
10.7

79.1

987
636
1,718
12,645

11,072

15,986

22.9
2.8
1.8

72.5

HERBS

Bedstraw

Clintonia

Club-moss, shiny. .
Cucumber root. . . ,
Dogwood, dwarf. .

Fern, beech

Fern, New York. .

Fern, oak

Fern, sensitive. . .

Fern, wood

Goldthread

Grass sp

Jack-in-the-pulpit .
Maianlhemiim . . . .
Miterwort, false. .

Nightshade

Partridge berry. . .

Hypericum

Sarsaparilla

Sedge sp

Starflower

Trillium, painted.

Trillium, red

Twinflower

Twisted stalk. . . .

Violet sp

Violet, false

Wintergreen

Wood sorrel

2.3
2.9
1.7
2.1
8.2
5.3
3.2

Total.

.2
28.5
4.1
.8
.4
4.7
4.0
.2
.4
1.4
1.1
3.7
1.1
1.1
.2
.4
.2
1.7
.6
.2
19.0

355
455
267
333
,279
833
497

33
4,441
643
133
67
733
631
33
67
221
167
576
167
167
33
67
33
267
100
33
2,963

15,594

1.2
1.8
.5
1.8
2.4
2.8

'r.2

31.4

'3.4
.5
4.2
4.0

9.9
1.2
.5

16.4

'2.8
13.9

190

286
86
286
381
442

190

4,990

" '537
86
667
632

1,575
190
86

2,613

" '442
2,210

15,889

3.4

"2.8
.5
.9

1.8
1.4
2.3
.5
28.8

3'

.

7

348

5

44

1

8

174

1

8

174

5

44

5

44

35

5

3,345

1

8

174

9

87

5

5

521

5

.1

478

174
130
217
44
2,713

9,426

72

Roosevelt li'ildlije Ihilletin

tween these two types (spruce flat and northern hardwood) is not as
great as originally presumed, their juxtaposition at these narrow
zones of mixed hardwood is, for practical purposes, close enough to
be considered an edge.

The analysis of the transition between these two types is based
upon 15 transects composed of 654 underbrush and herb plots of
which 270 were in the spruce flat, 90 at the center of transition, and
294 in the nortliern hardwood. The average length of the transects
was 540 feet but the effective transition in the lesser vegetation oc-
curred over a zone 450 feet wide. Although this was somewhat
wider than any of the other transitions studied, it was still well
within tlie cruising radius of ruffed grouse.

1. Vegelative Characteristics. The most abundant species of
arl)orescent reproduction at this edge was sugar maple which in
places attained numbers averaging in excess of 28,000 per acre in
the smallest size class (0"-6"), over 2,000 per acre in the middle size
class (6"-24"), and over 600 per acre in the largest size class (24"-
1" diam.). Although all of these large numbers occurred on the
hardwood side of the transition, considerable quantities of this species
also were found under the cover of the spruce type. The fact that
spruce and balsam were found under the edge of the hardwood type
demonstrates the zone of tension between the two types. Beech again
exhibited an inverse size class-numbers ratio, but a greater abun-
dance of this species was found on the hardwood side of the edge.
Yellow birch reproduction appeared to prefer the edge between the
two types rather than either one of the types alone. This is corre-
lated with a greater abundance of mature individuals of this species
at this zone. Reproduction of red maple, of spruce, and of balsam
was more abundant on the spruce side of the edge and gradually
diminished in numbers into the hardwoods. Occasional species of
reproduction in the spruce flat were hemlock, arbor vitae, and moun-
tain ash, while sucli species as white ash, basswood, and hop horn-
beam occurred scattered through the hardwoods. Black cherry and
shadbush were not indicated as exhibiting an}- preference, probably
due to the small number of occurrences. It is interesting to note
that mountain maple and striped maple, both favorite winter deer
foods, gradually increased in abundance from spruce to hardwood.

The same species of shrubs and herbs dominated the lesser vege-
tation at all parts of the transition, but there were minor shifts in
their relative abundance. The most abundant species of shrubs were
witch hobble, red and dwarf raspberries, and honeysuckle. The most
abundant herbs were wood fern, wood sorrel, false lilv-of-the-vallev.

c: <

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[73]

74

Roosevelt Wildlife Bulletin

false miterwort and violet. None of the shrub or herb species showed
a significant superabundance at the center of transition but two shrub
species and six herb species were most abundant on the spruce flat
side while two other shrub species and six herb species were most
abundant at the hardwood side. The total number of species across
the transition was greater than the number at any of the three parts.
There was a progressive increase in the total amount of shrubs from
the spruce flat to the northern hardwood. The greatest amount of
herb cover was found at the spruce flat side of the edge.

2. Wildlife Values. It is evident from the above that these two
types supplement one another in fulfilling the habitat requirements of
rufl^ed grouse and deer. This is further demonstrated in Table 18,
which summarizes the distribution of the more abundant species
across the edge. It is shown that features peculiar to spruce flat are
a year-round source of cover in the form of conifers, both large and
small, while browse species were furnished in the form of reproduc-
tion of balsam and red maple. Other important species more abun-
dant here than in the hardwoods were red raspberry (summer browse
and fruit), sedge (browse), dwarf dogwood (early fall fruit), false
lily-of-the-valley (early spring greens and late summer fruit), and
clintonia, violet and spring beauty (early spring greens).

Food species peculiar to the hardwood side of the edge were
sugar maple (browse and seed), beech (mast), and lesser amounts
of striped maple (browse). Hazelnut provided minor amounts of
browse and occasional mast while dogtooth violet and New York
fern provided spring greens but false miterwort served as a year-
round supply of greens.

The relative scarcity of browse-size (two feet high to one inch
diameter size class) reproduction of such species as balsam, arbor
vitae, and all maples on the spruce side of the edge may indicate more
than merely site preference or seed viability, i.e., winter concentra-
tions of deer in the cover of the spruce probably have had some
effect in holding back these species. This condition also is reflected
in the shrub cover, for although witch hobble is characteristic of
both types, the plants are bigger and occur in larger patches in the
hardwood. The witch hobble in the spruce t3-pes is so low that it
is often completely covered by snow, but a large proportion of the
witch hobble in the hardwood remains exposed for deer long after
that in the softwoods has been covered. This is a very fortunate cir-
cumstance for deer, and therefore one might well expect winter con-
centrations near the zone of juncture between these two tA'pes.

Other food species abundant in both types were wood fern and

TREE PLOT NOS.

6

5

4

3

2

1

Z

3

4

5

B

RUSH a HERB PLOT NOS

18

17

16

15

14

1 3

12

0

9

8

7

6

5

4

3

2

1

2

3

4

5

6

7

8

9

10

1 1

12

13

14

15

mm

^■

'V

»>

-

1

ALDER

•

2

BLUEBERRY

"

•

-

3

ELDERBERRY

-

_

— 1

♦

•

4

HAZELNUT

It

♦

*

•

•

5

HOLLY, MOUNTAIN

f—

»

•'

>

•

6

HONEYSUCKLE

;

*

•

•

•

-

1 — 1 —

_

'"4

7

LEATHERLEAF

•

8

OSIER, RED

•

•*

•

P

I

1.

-

-1

r

r-

%

8

r

-

-

-

-

9

RASPBERRY, DWARF

♦

-

•

Si

n

■K.

ft!

r

r

«•

;

-

L

*

•

I

•

10

RASPBERRY, RED

i

■M.

•

r

«.

*

■n

1 1

SWEET GALE

k

—

12

WITCH HOBBLE

-J"

1

_

-ir

fa-

J,
t.

s,

L

I"
s

I

3

JL

-n

M.

IT
f

-

i

1

- ■

1

13

ASTER SP

-]

1

S

•

•

•

r

14

BEDSTRAW

!

t

I

r

15

CLINTONIA

—

J

16

CLUBMOSS, SHINY

itt

4

P"ig. 11. Graphical analjsis of the distribution of the slirubs and herbs across the edge between
Spruce Flat and Northern Hardwood.

[75]

TREE PLOT NOS.

6

5

4

3

2

1

1

2

3

4

5

BRUSH a HERB PLCfT NO;

18

17

16

15

14

13

12

1 1

10

9

6

7

6

5

4

3

2

1

2 ' 3

r '
'i 1 e

3

2

r

13 14 15

•
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*

*

1

*:

U

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*

•

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f

1

4 :

u

W

1 7

CUCUMBER ROOT

•

-

f

r

r

r

1

J

■

1

18

DOGWOOD, DWARF

•

4-

i-

•

.

• ■

1 -

■ i

•

.

i

I

19

FERN, BEECH

-I-

20

FERN, BRACKEN

1

1

«

•Ml

i*

1
1

21

FERN, NEW YORK

1

■i-

•

22

FERN, Oak

*

•

J

-1

1

23

FERN, SENSITIVE

•

■

•

"L

«

«*

*

9

f

L
m

»

r

f

1

*-

•

_ I

V

i

t

1

t

i

r

t

I

24

t

•

t

i

*

■-

1

25

GOLDENROD. HAIRY

I—

f

■

f

1

26

GOLDENROD, WOODS

" -

¥

f

27

GOLDTHREAD

...

- ,

•

•

•

r

r

1

I

28

GRASS SR

•

-

—

,k

29

JftCK-IM-THE-PULPIT

*

*

•

•

\

i

f

•

•

t

♦

-

t

•

1

1

30

MAIANTHEMUU CAN.

•

•

i

1

♦

-4-

T

+-

w

t

i

«

t

t

\i

II

t

t

r

•

I

•

♦ •

tli

m

31

MITERWORT, FALSE

1

*

«

«

f]

32

NETTLE

Fig. 11 I Continued)

[76]

TRANSITION-

SPRUCE FLAT

CENTER

NORTHERN HARDWOOD

■ TREE PLOT NOS

6

5

4

3

2

1

1

2

3

4

5

BRUSH a HERB PLOT NOS.

18

1

:I7 16

15

14I13

1 1

10

9

6

5

4

3

2ll

2

3

4

6

9

10

1 1

14I15

J

1

»

♦

*

Nlun 1 sMAUL

*

4

WRTRIOGE BERRY

5

PIROLA

-

t.

,g

HYPERICUM

•

J

t

*

♦

*

SARSAPARlLLA

-

♦

. j

1-

— 1

•

u.

a

1

t

•

*

•

>a

SEDGE SP

....

*

—

-J

—i

9

SNOWBERRY

•

»

•

i

■

•

•

♦

-

•

0

SOLOMONS SEAL

*

♦

IT

■»

1 — 1

«

*

•

■

3

•

•

'1

STARFLOWER

♦

—

-I

— 1

■

1 — i

'Z

TRILLIUM, PAINTED

-

•

-nr

3

TRILLIUM, RED

1 —

—

' ■

*

•

—

*

*

•

*

t

*

4

TVyiSTED STALK

*

i

•

V

f

» ■

—

—

1

1

•

IK

5

VIOLET SP

-

—

•

T

.

*•

•

-

*

6

VIOLET, FALSE

'

♦

♦

—

—

— t—

m

7

WINTERGREEN

r

\

f

J.

t

i

f

f

I

t

1

r

1

L

a

F

•

i

n

a

1

*■

r

i

8

WOODSORREL

«

i

r

as

m

Fig. 11 (Concluded)

[77

78

Roosevelt Wildlije Bulletin

wood sorrel, both of which overwinter green. It is a well known fact
that on the Huntington the deer seek the clumps of wood fern in late
fall and early winter. They browse the tender "fiddle-heads," occa-
sionally also consuming the evergreen fronds and portions of the
rhizome. Grouse also feed on the wood fern and probably on wood
sorrel. Buds of yellow birch and sugar and red maples serve as the
principal winter food for ruffed grouse, while dwarf raspberry and
honeysuckle supplement the variety of summer fruits. Violets prob-
ably serve as early spring greens as does dwarf raspberry. Minor
amounts of several other species which provide both greens and
fruits are provided by each of the two types. However, due to in-
complete data on species habits and requirements it is impossible to
state the value and extent of use of these plants at this time.

Not shown in the data to any appreciable extent is the early
spring occurrence of dogtooth violet and spring beauty both of which
are very abundant in the hardwood and mixed hardwood types within
a very few weeks after the snow leaves. These are the two earliest
spring plants and probably serve grouse as the first spring greens
in the change from an all-bud winter diet.

In summary, it may be said that this edge surpasses even that
between mixed hardwood and spruce flat in \ariety and abundance
of food and cover for grouse and deer. In spite of this great diver-
sity, however, fall and overwinter fruits are lacking at this edge also.
(See Fig. 11 and Tables 19 and 20.)

F. The Edge Between Mixed Hardwood and Northern
Hardwood. These two types cover more of the Check Area (67
percent) than all of the other types combined; northern hardwood
occupies 1,500 acres, while mixed hardwood covers 1,212 acres. The
northern hardwood type occupies the well-drained lower slopes and
benches and represents the most fertile soils. The mixed hardwoods
are topographically above or below the northern hardwood and in
either case represent a trend toward conditions favoring spruce and
balsam fir rather than sugar maple. The larger amount of sugar
maple and relative scarcity of red spruce are the principal features
distinguishing the northern hardwood from the mixed hardwood type.

The edge between these two types was analyzed by ten transects
composed of 358 underbrush and herb plots. Of these, 138 were in
the mixed hardwood, 60 were at the center of transition, and 160
were in the northern hardwood. The average length of transects was
445 feet, but the eif'ective transition in the lesser vegetation occurred
over a zone about 375 feet wide.

1. Vegetative CJiaracteristics. The most abundant species of

■o

o
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2

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00 ro
00

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o o
o o
o o

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^ rj< \0
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^ vb o

■* o

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T-l Tt< so

^1 ^1 ^1

^ sO o
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.1 ^1 ^1

^ SCO
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— ^ -o

SO O

C-1

— I so

J J J

o o

I I I

^ so

rsi

^1 ^1 ^1

O O

■>* so

i'

sbo

St
■3l

J3 . V

_2

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^ o 5

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^' J£
^ sc c .5?

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80

81

TABLE 20. SUMMARY OF MATHEMATICAL ANALYSIS
OF THE SHRUBS AND HERBS ACROSS THE EDGE
BETWEEN SF^RUCE FLAT AND NORTHERN HARD-
WOOD

Northern

Spruce Flat

Center

Hardwood

135 plots

45 plots

147 plots

No.

Species

rer

bq. bt.

rer

isq. rt.

1 er

oq. rt.

cent

Coverage

cent

Coverage

cent

Coverage

of

per

of

per

of

per

Total

Acre

Total

Acre

Total

Acre

SHRUBS

1

Alder

1.1

137

1 . U

1 14

3

Elderberrv

.6

89

J.
•*

1 . 0

1 OQ

A 1
4 . 1

ooz

5

Hollv, mountain

If

.4

A A

44

6

Honeysuckle. . , .

y /u

Q 1

O . J

1 10 0

1,1//

' - ' Z.

0 . /

7
8

Leatherleaf

. 1

1 c

\j

Osier, red

1

. 1

1 A

9

Raspberr\-, dwarf

18 . 6

I , lyu

1 CQ 2
1 , 56o

y . u

10

Raspberrv, red

25.8

3,092

23.7

3,212

6.6

1,059

11

Sweet gale

.6

69

12

Witch hobbie

4j . U

5 , 104

55 . /

/ , 508

7/1 C
/4 . 5

1 J. y UUU

Total

1 1 , yyo

13,574

16,101

HERBS

1

.4

oy

2

Bedstraw

. 0

1 CiA

. 0

QO

8y

7
. /

1 no

3

0 A

L . 4

/I 1 c
410

1 7
1 . 5

1 7Q
1/8

1 A
1 . 0

4

Club-moss, shin\'. .

. /

I/O

0 A

L . 4

o4U

1 r
1 . 5

5

Cucumber root

1 . L

ILL

1 o

1 . y

0 A7

/o/

0 o

/ . y

430

6

Dogwood

4.3

753

3.4

473

.6

91

7

Fern, beech

1 1

1 . i

1 Q7

ly /

1
. o

44

Q

. 6

1 1 7
11/

8

Fern, bracken ....

1

. i

lo

9

Fern, New York

.6

98

5.6

773

10

Fern, oak

1.0

182

.9

133

.4

63

11

Fern, sensitive

A

. 4

/4

1

. i

10

12

Fern , woo l

O 1 . ^3

7.% 1

JO . 1

4 , 000

04 . U

4 , V / u

13

Goldenrod, hairy. . .

.6

113

14

Goldenrod, woods...

1 A

loz

1 Q

1 . 8

0 C 1
/5 1

Q

. 8

117
11 /

15

Goldthread

1 1

Z41

1

. J

A A
44

0

. /

07
/ /

16

Grass sp

A

. 1

1 A

1 4

Q

. y

17 7

loo

. 0

yo

17

Jack-in-the-pulpit

. 0

8y

0 o
/ . U

OQQ

/8y

18

Maianlhemum

1 CO

A A

4 . 0

AC 1
05 1

. y

1 7 C
105

19

Miterwort, false

1 o
1 . 8

"3 O C

3/5

5 . y

QOQ
8/6

n 0

y . -i

1 1A A

1 , 044

20

Nettle

1 . Z

21/

21

Nightshade

.2

44

3.4

484

.8

117

22

Partridge berry

.2

30

.3

44

.2

27

23

Pirola

.1

15

24

Hypericum

1.0

182

25

Sarsa pari 11a

1.8

315

' ' .3

44

r.3

186

£.\y

8.0

1,421

.3

44

1.0

150

27

Snowberry

.1

15

.9

126

Z8

.1

15

' ' .3

44

1.3

186

29

Starflower

1.4

252

.9

133

1.6

236

30

Trillium, painted

.3

59

.3

44

.6

90

31

Trillium, red

.1

15

.3

44

.4

54

32

Twisted stalk

.5

89

.6

89

1.1

164

33

Violet

2.7

468

3.4

473

4.5

660

34

Violet, false

1.4

256

.6

89

1.9

284

35

Wintergreen

.4

69

36

Wood sorrel

30.1

5,304

36'9

4,345

22.8

3,333

Total

17.636

14,053

14,652

82

Roosevelt li'ildlife Bulletin

reproduction was sugar maple, and it was much more abundant on
the northern hardwood side of the edge. Beech again showed a re-
verse size class-numbers ratio and not much preference of either
type, although its reproduction was slightly more abundant in the
northern hardwood. The most interesting feature relative to yellow
birch reproduction was the absence of trees in the size class two feet
tall to one inch in diameter. Yellow birch in this size class has been
very scarce at all edges, and this probably indicates a high degree of
environmental resistance, both in the form of vegetative competition
and wildlife feeding. Since it does occur in fair abundance in the
overhead canopy of mature trees, its scarcity on the ground removes
it from the group of major climax dominants, and indicates that its
continued presence is dependent upon repeated environmental dis-
turbance. Although not overly abundant in mixed hardwood type,
red spruce reproduction nevertheless was more abundant there than
in the northern hardwood type. Balsam fir reproduction was rela-
tively scarce at this edge. It is not as well adapted to mixed hard-
wood as it is to spruce flat, and is practically absent in the northern
hardwood type. Red maple, though somewhat more abundant in the
mixed hardwood, was nevertheless relatively rare at this place. The
occurrence of white ash at this edge w^as very spotty. This species
usuall)- is correlated with a highly organic mull soil and therefore
occurs in greatest al)undance in the northern hardwood type. The
absence of hemlock reproduction at this edge is further indication
of the poor ability of this species to reproduce itself. Apparently
the same is true of black cherry. Striped maple reveals no obvious
preference for either type but mountain maple apparently finds the
rockier and more shallow soils of the mixed hardwood more suitable
for development.

The principal features relative to the distribution of shrubs and
herbs across this edge may be summarized as follows : The most
abundant shrub species was witch hobble and this w-as supplemented
by honeysuckle, red raspberry, and dwarf raspberry. The most abun-
dant species of herbs were wood sorrel, wood fern, and false lily-of-
the-valley, although false miterwort and violet were fairly common.
While there was no change in dominant species across the transition,
there were minor shifts in relative abundance. Two species of shrubs
and five species of herbs were most abundant at the center of transi-
tion ; another shrub species and four species of herbs were most abun-
dant at the side of the edge toward mixed hardwood; and one shrub
species and si.x herb species were most abundant on the side toward
northern hardwood. The variety of herbaceous species across the
transition was greater than that at any of the three parts of the

WQ

o >

T3 O
0) O

cX

in _c

i; "
= o

c/) 'V

u (K TO

3 oj 1) n O

Qu.>u

cn
O

- H
S o =.2

^w. ■« o >

•= § § §1-5

V

!= tJ
- 3

o

c

.9 >■

O ^ D

a..2 a

o oj -4- a
c

3^ «

o.

■•3 c

<U 3

.E «i
% E

[83]

84

Roosevelt Wildlije Bulletin

transition, but all recorded species of shrubs were represented at the
mixed hardwood side. Both shrubs and herbs occurred with the
greatest abundance at the center of transition and with the least
abundance on the mixed hardwood side.

2. Jl'ildlife Values. The principal vegetative features of value
to wildlife which are contributed by each of the two adjacent types
are summarized in Table 21. From this table it is evident that neither
type provided abundant winter cover, for the conifers in the mixed
hardwood type were somewhat scattered. To be sure, occasional
small clumps of reproduction were present, but more often these size
classes occurred as scattered individuals. Red raspberry browse and
fruit were more abundant in this type than in the northern hardw^ood,
and small amounts of clintonia and false lily-of-the-valley provided
additional early summer fruits and greens.

Features contributed principally by the northern hardwood side
of the edge were scattered amounts of dwarf raspberry (greens and
fruit), especially in moist spots, while liberal amounts of false miter-
wort provided overwintering greens. Violet, spring beauty, dogtooth
violet, and cucumber root were more abundant in this type, and
represent a potential source of early spring greens.

Among the features present in both types was a liberal amount
of w^itch hobble, which serves as excellent summer cover and an im-
portant source of winter deer browse. It should be noted that this
.species attains its best height growth and greatest density in these
two types. Where the ground is not taken up with patches of witch
hobble, large numbers of sugar maple reproduction usually dominate
the lesser vegetation (especially in the northern hardwood), thus
supplementing the amount of available browse. Fairly abundant
beech reproduction in the largest size class serves as additional sum-
mer and fall cover. Mature sugar maple and beech provide ma.^t
while buds of maple and yellow birch also are used as winter food
by grouse. The browse supply is supplemented in both types by
striped and mountain maples as well as a small amount of honey-
suckle. Wood fern, wood sorrel, shiny club-moss, and false violet,
all of which overwinter green, occur sparingly throughout both types.
Spring beauty and violets provide abundant supplies of early spring
greens. Although it is known that the wood fern is used by both
deer and grouse, the extent to which the other three species are used
has yet to be determined. Small amounts of several other species of
herbs supplement the variety, if not the quantity, of greens and fruits.
In spite of the increased variety of buds, browse, fruits, and greens
at this edge, nevertheless winter cover and fall and overwinter fruits
are still deficient. (See Fig. 12 and Tables 22 and 23.)

i

TRANSITION-

MIXED HARDWOOD CENTtR

NORTHERN HARDWOOD

TREE PLOT NOS.

I

BRUSH a HERB PLOT NOS

HONEYSUCKLE

4 RASPBERRY, DWARF

5 RASPBERRY, RED

5if

1i 2 ■

6 WITCH HOBBLE

7 ASTER, LARGE-LEAVED

8 iBANEBERRY

CLINTONIA

II CLUBMOSS, SHINY

12 I CUCUMBER ROOT

13 I DOGWOOD, DWARF

14 I FERN, BEECH

FERN, NEW YORK

FERN, WOOD

GOLDENROD, WOODS

GOLDTHREAD

JACK-IN-THE-PULPIT

UAIANTHEMUM CAN.

MITERWORT, FALSE

NIGHTSHADE

PARTRIDGE BERRY

Fig. 12. Graphical analysis of the distribution of shrubs and herbs across
the edge between Mixed Hardwood and Northern Hardwood.

[85]

86

Roosevelt Jf'ildlije Bulletin

TRANSITION -

MIXED HARDWOOD

GENTER

NORTHERN HARDWOOD

TREE PLOT NOS.

4

3

2

1

1

2

3

4

BRUSH S HERB PLOT NC5S.

12

1 1

10

9

8

7

6

5

A

3

z

1

1

2

3

4

6

7

8

9

10

1 1

t2

•

•

-.

26

SARSAPARILLA

•

t

•

—

27

SEOGE SP

•

•

S

•

•

•

•

j

28

SOLOMON S SEAL

* •

%

•

1

29

STARFLOWER

—

•

1

30

TRILLIUM, PAINTED

•

31

TRILLIUM, RED

32

TWIN FLOWER

t

33

TWISTED STALK

•

1 —

t

34

VIOLET SP

V

—

\

.

■

35 VIOLET, FALSE

r

r

T

1

r
t-

r

f

!

f

36

WOOOSORREL

r

V

L

r.

i

1

l-'ig. 12 (Concluded)

G. The Edge Between Spruce Flat and Spruce Swamp. Al-
though the spruce swamp type covers an aggregate of only 49 acres,
its wildlife value is greatly increased by its being scattered widely
throughout the Check Area in small blocks. The most distinguishing
feature of this type, other than the conifer cover, is the presence of
sphagnum moss. This species represents a year-round high water
tal)le and usually is the most abundant non-woody species. Since the
presence of the sphagnum usually marks the edge of the swamp, the
width of this edge usually can be measured in inches rather than in
feet or yards.

The vegetation across the edge between spruce flat and spruce
swamp was analyzed by a series of 15 transects. The swamps were
so small that the transect was continued across each one into the
spruce flat on the opposite side. Thus, 30 edges rather than 15 were
actually plotted. These transects were composed of 438 underbrush
and herb plots of which 168 were in spruce flat, 180 at the center of
transition, and 90 in the swamp.

•V

o
o

u

a)

X

o
2

o
o

-o

X

-a

D-c V

o .

u

^z

^ 6

a

o i*<

o —

O

o o
©•no
— < 00 00

o o

\0

U-)

o o
— o

(N 00

o o o
o o o

00

1^ trj o

•O O

lO o
r~i

, o o

( O
I o o

O

o o
o o

.2
-5

^ ^

J ^1 ^1
^ o o

O

O 'O

O 00

IT) O ^

00 — ' fN
vO ^

t-~ o
o o o
o ^ <^

O

ro O IT)
lO fT)

O

o o o

^"0 1^
O ^

O
O

-H rf O
CM

v'

■ in O
• "1 M

■ 00 o

O r~

O O ro

O O t~0
O O IT)

■rt to Tj<
— ' 00 O
^ t-^

• in m

■ M O

■ tN 00
• CM

. OS
, -rf OS
■ T-l Ov

• I>a (N

■ r<i tN

■ CM CN

• m in

■ O

■ o o

■ o o

.2
■5

VO

CN

J

ij< SO o

CN

.2
-5

^ '*isb

CN

v' ^1

^ so o

CN

.2
-3

^^sb

^'

if o o

CN

I so

^1 ^1

Tt< so O
CN

-a
Pi

[87]

OS
go

I— I

c/)0

< ^

<

< H

rg
<

I I I

fN

Tt< — 1^
« «^

O »0
"7 O

r-4 .-H

pr> O

<r) o >o
^ t-~ "*

IV5 O
O O

CN

PC

— I 00

— rc

< CM —

PO O

o

PC c

— o

^1 ^1 ^1

^ o o

o o
o o

oo
oo

— Tf O

^1 ^1 ^1

^j" o o

in r*5

1^ o
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— so

^1 ^1 ^1

— Tj< >0
CM

a'

■* OO

E

XI

c
o

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O

o

3

[88]

89

TABLE 23. SUMMARY OF MATHEMATICAL ANALYSIS
OF THE SHRUBS AND HERBS ACROSS THE EDGE
BETWEEN MIXED HARDWOOD AND NORTHERN
HARDWOOD

Mixed

Northern

Hardwood

Center

Hardwood

69 plots

30 plots

80 plots

No.

Species

Per

Sq. Ft.

Per

Sq. Ft.

Per

Sq. Ft.

cent

Coverage

cent

Coverage

cent

Coverage

of

per

of

per

of

per

Total

Acre

Total

Acre

Total

Acre

CUIDT IRC
OillvU DO

1
1

2

29

2

Hazelnut

1

1

136

67

1

I

. \j

150

3

Honeysuckle ....

25

9

3,151

10

7

1 ,693

12

.4

1 ,832

4

Raspberry dwarf

5

0

610

7

6

1 , 193

13

0

1 ,914

5

Raspberry' red

10

8

1 ,317

11

1

1 , 750

4

.2

618

6

57

0

6,940

2

11,054

A

10,222

Total

12,183

15,757

14,736

1 1 1^ i\ f > . J

1

Aster, large-leaved

3

7

473

2

/5

3

Bedstraw . . . .

4

2

490

1

fi

o

267

1.

o

1

400

4

Clintonia

3

2

ill

7

4

1 ,085

8

100

5

Club-nioss shiny

4

4

519

5

1

752

2

7

350

o

1

5

174

1

4

200

3

7

475

7
8

Dogwood, dwarf

1

0

116

Fern , beech

2

29

4

50

9

Fern, New York

4.

67

1
i

1

141

10

Fern, oak

2

29

4

67

4

50

11

Fern wood

31

5

3,690

20

6

3 ,032

28

1

3 , 632

12

Golden rod wood

2

29

2

1

309

13

Goldthread

2

4

279

2

1

309

6

75

1 A.

4

67

4

50

15

Jack-in -t he-pulpit

1

0

116

2

6

376

5

2

673

16

Maianthemiim

6

2

725

3

6

533

1

7

225

17

Miterwort false

2

7

319

7

8

1,152

10

6

1,370

18

Nightshade

5

58

2

25

19

Partridge berry

2

29

4

67

4

50

1

2

145

2

6

376

8

100

21

Sedge, sp

7

87

9

133

3

8

498

22

Solomon's seal

5

58

4

67

8

100

23

Starflower

7

87

9

133

8

100

24

Trillium, painted

1

0

116

4

67

4

50

25

Trillium, red

2

29

4

67

2

25

26

Twinflower

4

50

27

Twisted stalk

1

2

145

"l

3

333

1

7

225

28

Violet, sp

2

6

308

2

3

333

8

9

1,146

29

Violet, false

1

7

203

4

4

643

2

7

350

30

Wood sorrel

30

4

3,556

28

9

4,244

16

3

2,112

Total

11,713

14,679

12,920

90

Roosevelt Wildlife Bulletin

1. Vegetative Characteristics. Red spruce was the most abun-
dant species of reproduction in these tj'pes. It was most abundant in
the swamp and least abundant in the spruce flat. (For the sake of
those who would expect black spruce here, it should be explained
that this species is of very limited occurrence on the Huntington.)
Balsam fir was the second most abundant species and it also was
most abundant in the middle of the swamp and center of transition
rather than in the spruce flat. But even here, the paucity of individ-
uals in the largest size class (two feet tall to one inch in diameter )
was striking. This cannot be taken as a measure of vegetative com-
petition, rather, it is an indication of the intensity of deer browsing.
Red maple, too, was quite abundant at this edge, but it showed a
progressive increase in numbers from the swamp outward. Hemlock
reproduction was more abundant at this edge than at any other, and
increased in abundance near the swamp, but even at that it was rep-
resented by very few individuals in even the smallest size class, while
there were none at all in the largest size class. Arbor vitae and
black ash are not characteristic of all spruce swamps, as is indicated
I;y their very low incidence.

Hardwood reproduction (beech, birch, and the three maples)
was surprisingly abundant ; however, each of these species except
yellow birch, which was erratic in occurrence, increased in abundance
from the swamp outward. The presence of sugar maple seedlings
even in the center of the swamp may be explained by the extensive
sources of seed supply in adjacent areas, and by the small size of
these swamps. Furthermore, it will be noted that the numbers of
this species were incomparabl}- lower at this place than on favored
sites, and that they died before reaching two feet in height. The
presence of beech reproduction even in the largest size class is char-
acteristic of this species as displayed on other spruce flat sites.

Although the shrubs at this edge were more varied than at many
others, they were, nevertheless, not especially abundant. In total
amount, they decreased from the spruce flat into the spruce swamp,
being supplanted there by coniferous reproduction. \\'itch hobble
was the most abundant species at the spruce flat side and the center
of transition. Honeysuckle and red raspberry were the next most
abundant species on the spruce flat side, but dwarf raspberry ex-
ceeded even witch hobble in abundance in the swamp. Blueberry
also was relatively more abundant in the swamp. Unlike the shrubs,
the total amount of herb cover increa.sed progressively into the
swamp. On the spruce side the most abundant species were wood
sorrel, dwarf dogwood, and wood fern, while the most abundant

o

e

C M

n -a

a

3 ^ o i;
Sp>u

11 0)

US

ECO

-•uS §

I 8 s

• - T3 o O ra r;

13 o ^ Ol2

J3

3

c <u
o

^ a.

tao £ O

Da >

u

[91]

92

h'uoscvcll li'ildlijc lUillctin

species in the swamp were sphagnum moss, cinnamon fern, sedge,
wood sorrel, wood fern, and dwarf dogwood. Four shrub species
and ten herb species evinced a preference for the swamp while three
species of shrubs and six of herbs were most abundant at the spruce
flat side. The variety of both herbs and shrubs was greater across
the transition than at any of its three parts.

2. Wildlife Values. The principal vegetative features of im-
portance to wildlife contributed by the two adjacent forest types at
this edge are summarized in Table 24. Cover in the form of mature
conifers (spruce, balsam, hemlock), and spruce of all sizes was
abundant in both types, but reproduction was especially heavy in the
swamp. Young balsam which have grown beyond the reach of deer
still present lower limbs that are browsed when weighted down with
snow. A small amount (jf red mai)le browse was found in both types,
although most abundant in the spruce flat. It cannot be said with
certainty whether the absence of the largest size class of reproduction
of this species is due to overbrowsing by deer or whether it is purely
a vegetational phenomenon. W itch hobble was the most abundant
browse species on the spruce flat side and this was supplemented
lightly with sugar maple and striped maple reproduction and red
raspberry. Mature yellow birch and red maple produced buds for
winter grouse food on the spruce side while wood fern as an over-
wintering green and clintonia and red raspberry as summer fruits
were present in limited amounts.

Wood sorrel and goldthread were a source of overwinter greens
in both types, and false lily-of-the-valley provided early spring greens
and late summer fruit. Dwarf dogwood provided late summer and
early fall fruit in both types.

Sphagnum moss is listed as the most abundant species of herb
in the svvamp side of the edge but its wildlife value has not been
determined. Cinnamon fern fronds probably are taken by deer in
the spring before they are unfolded. Sedges also are available for
spring and early summer browse. Small amounts of fruit are pro-
vided by blueberry, dwarf raspberry, and occasional clumps of creep-
ing snowberry. \'iolets were more abundant at this portion of the
edge and may serve as early spring greens. Several other species
supplemented the variety of greens and summer fruits in both types.
This edge appears to have most of the vegetational requirements for
deer, but it apparently is lacking in fall and overwinter fruits for
grouse. (See Fig. 13 and Tables 25 and 26.)

TRANSITION -

SPRUCE
FLAT

CENTER

SPRUCE
SWAMP

CENTER

SPRUCE FLAT

TREE PLOT NOS.

1

1

1

2

BRUSH a HERB PLOT NOS

3

2

1

2

3

5

6

1

ALDER

m

1

•

2

BLUEBERRY

•

•

3

DOGWOOD, ALTERNATE

4

HOLLY, MOUNTAIN

I.

' u

t

.

1

r

*

.

5

HONEYSUCKLE

t

-

•

~

6

LEATHERLEAF

7

MEADOW SWEET

-

t
r

s

•

•

e

RASPBERRY, DWARF

}

■y-

9

RASPBERRY, RED

'

-

T

r

10

WITCH HOBBLE

^
u

r

j-
I
t

_

,3a

?

s

j_

f-

3s.

f

t

1 1

BEDSTRAW

I

:

12

CLINTONIA

t

•

4-

-

t

.

•
•

•
•

13

CLUBMOSS, SHINY

•

-1

•

•

14

CUCUMBER ROOT

•

•

*

♦

r
a

f

I*

t

t

1

IS

DOGWOOD, DWARF

r

\

—

—

16

FERN, BEECH

Fig. 13. Graphical analysis of the distribution of the
shrubs and herbs across the edge between Spruce Flat and
Spruce Swamp.

[93]

TRANSITION -

SPRUCE
FLAT

CENTER

SPRUCE

CENTER

SPRUCE FLAT

TREE PLOT NOS.

1

1

2

BRUSH a HERB PLOT NOS.

3 U 1 1

V

— [

.23

5 1

6

w

1

■

1

m

I

17

FERN, ONNAMON

1

i

i

-

1

-

~_:

■ i

18

FERN, NEW YORK

0:

-4

J

-

i

19

FERN, SP

J

20

FERN, OAK

1

-

t

21

FERN, SENSITIVE

1

-

L

f

i

22

FERN, WOOD

1

J.
-

f

1

.

5

a.
-

s.
-

I*

♦ "

f

r

*f
L

L

L

L

•

•

.

•

23

GOLDEN ROD, WOODS

\

•

t

i

:

^

•

•

•

r

24

GOLDTHREAD

*

r

i.

s

Mm

•

r

•

■

•

•

*

25

GRASS SP

•

26

JAGK-IN-THE- PULPIT

27

JEWELWEED

•

•

f

1

I

1

r

1

1

1

•

*

f

28

MAIAf/THEMUM

t

•

•

1

1

t

1

■t

29

MITERWORT, FALSE

30

NIGHTSHADE

#

31

SARSARARILLA

32

SEDGE, SP

!"■

Fig. 13 f Continued)
[94]

TRANSITION-

SPRUCE
FLAT

CENTER

SPRUCE
SWftMP

CENTER

SPRUCE FLAT

TREE PLOT NOS.

1

1

1

2

BRUSH 8 HERB PLOT NOS.

3

2

1

2

3

4

5

6

4

w

r

•

33

SNOWBERRY

•

%

• b •! ttt 1

**

34

SPHAGNUM

1

9

*

1

1

«

■

&

35

STAR FLOWER

J

*

*

•

•

•

36

STRAWBERRY

-

--"

*

37

TRILLIUM, nwNTED

•

*

*

•

^ -

38

TWIN FLOWER

1

39

TWISTED STALK

1

{

•

z

z

r

*.

*

40

VIOLET SR

%

*

¥

41

VIOLET. FALSE

h

i

?■

t

f

\

a

1.

m

1

42

WOODSORREL

i

r

k

r

s

h

1

Fig. 13 (Concluded)

[95 1

96

TABLE 25. MATHEMATICAL ANALYSIS OF THE ARBOR-
ESCENT REPRODUCTION ACROSS THE EDGE BE-
TW EEN SPRUCE FLAT AND SPRUCE SWAMP EX-
PRESSED IN NUMBERS PER ACRE

Species

Size class*

Spruce
r lat

Center

Spruce 1
Swamp j

Center

Spruce Flat

Plot
No. 1

Plot
No. 1

Plot 1 Plot
No. 1 No. 2

Spruce

24"- 1" diam.
6 "-24"
0"- 6"

67
533
600

400
1,200
5,133

845
1.510 1
5,780

578
1,465
6,320

178
666
4,360

286
1.000
6,480

Balsam

24"- 1" diam.
6 "-24"
0"- 6"

" 400
23,870

22
132
11,900

67
6,133

67
12,030

111

4,580

10,940

Red ma])le. . .

24"- 1" diam.
6 "-24"
0"- 6"

533
7,540

22
7,410

'266
3,990

67
5,000

89
6,490

1,048
10,230

Hemlock

24"- I" diam.
6 "-24"
0"- 6"

22
111

400

733

533

356

238

Arbor vitae . .

24"- 1" uiam.
6 "-24"
0"- 6"

200

67

22

Beech

24"- 1" diam.
6 "-24"
0"- 6"

2,674
200

133
67

44

67
22

156
244
133

238
191
95

Sugar maple .

24"- 1" diam.
6 "-24"
0"- 6"

200

1,667

877

44
488

200
378

111

1,265

191

762

Alountain
maple

24"- 1" diam.
6 "-24"
0"- 6"

933

1,000

66
488

176
445

44
445

191
95

Striped

maple

24"- 1" diam.
6 "-24"
0"- 6"

"67
1,600

"22

333

155

"89
400

1,376

48
952
2,333

Yellow birch .

24"- 1" diam.
6 "-24"
0"- 6"

800

667

88
1,733

155
1 1,920

88
822

572
1,333

Black ash . . . .

24"- 1" diam.
6 "-24"
0"- 6"

' " "44
22

67

44

Mountain ash

24"- 1" diam.
6 "-24"
0"- 6"

267
133

22
111

22
178

190

Shadbush. . . .

24"- 1" diam.
6 "-24"
0"- 6"

"266
133

"466
267

"22

" 66
22

1 "266

" "48

* All figures refer to height unless followed by term "diam." which refers to
diameter.

97

TABLE 26. SUMMARY OF MATHEMATICAL ANALYSIS
OF THE SHRUBS AND HERBS ACROSS THE EDGE
BETWEEN SPRUCE FLAT AND SPRUCE SWAMP

No.

Densities and Coverages

Spruce Flat
81 plots

Center
60 plots

Spruce Swamp
45 plots

Per
cent

of
Total

Sq. Ft.

Coverage
per
Acre

Per
cent

of
Total

Sq. Ft.

Coverage
per
Acre

Per
cent

of
Total

Sq. Ft.

Coverage
per
Acre

1

2
3
4
5
6
7
8
9
10

SHRUBS
Alder

'3.1

' ' .3
11.9

19.3
8.5
56.9

1.2
16.4

'2.3
13.9
.6
.6
35.8

29.3

89
1,262

178
1,068
44
44
2,749

Blueberry

Dogwood, alt .-leaved...

Honeysuckle

Leatherleaf

.2
.2
.2
11.4

25
25
25
1 , 556

317

33
1,231

Raspberry, dwarf

Raspberry, red

1.9
8.5
77.7

264
1,168
10,602

1,992
880
5,881

Witch hobble

Total

2,251

13,665

10,334

7,685

1

2
3
4
5
6
7
8
9
10
11
12
13
14
15
16
17
18
19
20
21
22
23
24
25
26
27
28
29
30
31
32

HERBS
Bedstraw

.2
6.2
1.3
1.0
18.5

.1
9.0

17.1
.2
3.8
.1

49
1,328
272
222
3,971
25
1,933

3,669
49
828
25

"2^1
1.6

.5
9.6

.8
7.8

.1

.1
1.5
14.0

. 1
4.9

.2

.1

'2^9
.7
.1
.6
5.2
2.4
.1
12.6
1.0

' ". Z
.3

"ZA
1.0
25.8

.3

'4^6
.4

22.0
.1
.3
.4
5.0
.3
3.5
.2

.1
1.0
.9

' ' .3
11.6
1.6

31.8
.4

"a

.7

"z.Q

.7
9.6

Clintonia

Club-moss, shiny

Cucumber root

700
551
167
3,261
228
2,635
33
33
518
4,750
33
1,661
67
33

473
133

Fern, beech

Fern, cinnamon

Fern, New \ ork

Fern, oak

Fern, sensitive

Fern, wood

Goldthread

Grass, sp

Jack-in-the- pulpit

Jewelweed

2,037
178

9,604
44
133
178

2, 183
133

1 , -'^35
89

44
444
384

Maianlhemum

Nightshade

4.5
.5

963
99

967
233
33
200
1,766
818
33
4,277
333

Sarsaparilla

Sedge, sp

Snowberry

Solomon's seal

.5
.2
.2

99
49
49

133
5,060
695

Sphagnum

Starflower

Strawberry

1.7
.1
.6
.1
.2
.7
.8
32.3

370
25

124
25
49

148

173
6,958

13,891
178

Trillium, painted

Twinflower

Twisted stalk

100
100

44
311

Violet, sp

Violet, false

Wood sorrel

Total

1,140
355
8,726

1,314
311
4,204

21,502

33,751

43,733

98

Roosci'cU Wildlife Bulletin

H. The Edge Between Spruce Flat and Opening. Natural

openings in mature forest t}pes sometimes occur due to windthrow
and various other causes. A few natural openings have been created
on the Check Area but they are very small, aggregating only 2.3
acres in the spruce flat type. These openings are very temporar}- in
nature, and may be obliterated in as few as 20 years by rapidly grow-
ing new trees. However, during the period that they remain open
they acquire a characteristic vegetation which is different from that
of the surrounding forest and thus enhance the variety and abund-
ance of wildlife food and cover. In this regard, it should be pointed
out that forest openings vary considerably in size and characteristic
vegetation and that the eight sampled in this study represent only
one of these many kinds.

In estal)lishing transects across this group of edges, the strip
in some cases was laid entirely across the opening and continued into
the forest on the far side. Thus, the 12 transitions plotted were de-
rived from only eight transects. These consisted of 114 underbrush
and herb plots on the spruce flat side, 70 at the center of transition,
and 84 in the opening, making 268 plots in all. Although the average
length of the transects was about 275 feet, the effective transition in
the lesser vegetation often was completed over a zone less than 100
feet wide.

I. J'cgctath'c Characteristics. The very effective seeding of
sugar maple is demonstrated by the abundance of its seedlings at
these openings in the centers of spruce flat types, for it surpassed
each of the typical spruce flat species in quantity. Xot shown by the
data, however, is the very limited number which succeeded in attain-
ing a diameter of one inch or more. Beech reproduction also was
fairly abundant on these sites, with a greater number in the larger
size classes, as usual. The typical aspect of these clearings exhibited
a number of young beech and spruce in a matrix of red raspberry
canes. Reproduction of most trees increased in number from the
clearing into the forest, but ash and black cherry appeared to be more
abundant in the clearing.

The total amount of shrub growth nearly doubled in quantity
from the forest into the opening. The most abundant species in the
forest was witch hobble, followed by dwarf raspberry, red raspberry
and hazelnut. Whereas red raspberry increased tenfold and domi-
nated the site in the opening and dwarf raspberry doubled in quan-
tity, the other species decreased in amount. The herb cover also
decreased b\' about a third in both quantity and variety from the
forest into the opening. The most abundant species in the forest

J3

(LI

01

J3 „

rt J;
u O

2^ tuO
2 ^ "O

P ^ O m r/5

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<D

J3

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ri.T3 o >- V;
in -n S. O O

X)

o

o

T3 c

S ?

_ c

(U —

[99 1

100

Roosevelt Wildlife Bulletin

were wood sorrel, wood fern, dwarf dogwood, and false miterwort,
while in the opening false miterwort, wood fern. New York fern,
violet, and grass were the most abundant. In the case of both under-
brush and herbs, the variety of species across the entire edge was
greater than that at any portion.

2. Wildlife ['allies. The increased variety of the principal vege-
tative features of value to wildlife at this edge is summarized in
Table 27. Year-round cover is indicated on the spruce side, but
abundant highly effective summer cover is provided by the red rasi>-
berry of the clearing. Mature yellow birch and red maple scattered
through the spruce tiat provide ample winter buds, and wood sorrel
and wood fern provide overwintering greens. False lily-of-the-valley
furnishes a small amount of late summer fruit and dwarf dogwood
retains some fruit into the fall. The spruce side also supplies browse
in average amounts of witch holible and hardwf)od reproduction, as
were present in limited amounts.

Red raspberry grows prolifically in the openings and thus pro-
vides large amounts of summer and early fall browse as well as
al)undant summer fruit. The other si>ecies of underbrush and herbs
in llie clearing serve principally as browse and greens. Hazelnut
occurs near the edge in both types, but it normally provides a very
limited amount of mast. Violet and false miterwort, also common
to both types, further diversify the variety of spring and overwinter-
ing greens. Additional variety of greens and fruits are provided by
several other species of herbs wliich occur in very small quantities
in both types.

From the al)ove, it is ol)vious that the clearing provides food
and cover during the summer and early fall, but the increased variety
due to the edge is still lacking in fall and winter fruits. (See Fig.
14 and Tables 28 and 29.)

I. The Edge Between Northern Hardwood and Opening.

Tlie total area (1.9 acres) of the few small openings in the northern
hardwood type was even less than that of those in spruce flat. !Most
of these openings too were due to windthrow and, as in spruce, rep-
resent but one of several different possible types of revegetation. By
extending the plots entirely across some of the openings 14 edges
were sampled by nine transects. These were composed of 268 under-
brush and herb plots of which 110 were in the northern hardwood,
84 at the center of transition, and 74 in the openings. The
average length of the transects was 240 feet, but in most cases the

TABLE 28. MATHEMATICAL ANALYSTS OF THE AR-
BORESCENT REPRODUCTION ACROSS THE EDGE
BETWEEN SPRUCE FLAT AND OPENING EXPRESSED
IN NUMBERS PER ACRE

Species

Size Class*

Spruce Flat

Center

Opening

1 lot
INo. 0

rlot
No. 2

rlot
No. 1

rlot
No. 1

riot

Snrii r*p

24"- 1" diam.
6 "-24"
0"- 6"

111

500
3,945

222
167
417

29
147
88

67

AA'7
001

O , 00/

24"- 1" diam.
6 "-24"
0"- 6"

28
1,834

235
688

67
166

3,333

6,450

333

Red nitiple.

24"- 1" diam.
6 "-24"
0"- 6"

29
647
1,940

556

0, IzU

1,415
5,190

67

667

\'ellow birch.

24"- 1" diam.
6 "-24"
0"- 6"

167
723
778

56
472
583

67
300
133

472
1,058

3,000

3,333

T-ffnilorU'

24"- 1" diam.
6 "-24"
0"- 6"

333

28

Beech

24"- 1" diam.
6 "-24"
0"- 6"

111

639
278
138

264
324
176

433

56

24"- 1" diam.
6 "-24 "
0"- 6"

333

167
3,330
2,667

2,945
6,725

147
1,712
2,175

33
467
1,365

Striped nitiple

24"- 1" diam.
6 "-24"
0"- 6"

"l67
444

500
333

235
294

IVIountain
maple

24"- 1" diam.
6 "-24"
0"- 6"

"222
389

"555
611

235
88

" "76
133

IVIountain ash

24"- 1" diam.
6 "-24"
0"- 6"

"667

111

29

White ash ,

24"- 1" diam.
6 "-24"
0"- 6"

56
56

83
56
111

57
147
117

33
267

Black ash ....

24"- 1" diam.
6 "-24"
0"- 6"

29

33
33

Black cherry .

24"- 1" diam.
6 "-24"
0"- 6"

' "56
56

472

lis

440

67

Shadbush

24"- 1" diam.
6 "-24"
0" -6"

56

28
56

29
176

233
33

Arbor vitae . .

24"- 1" diam.
6 "-24"
0"- 6"

1,333

* All figures refer to height unless followed by term "diam." which refers to
diameter.

102

TABLE 29. SUMMARY OF THE MATHEMATICAL ANALY-
SIS OF THE SHRUBS AND HERBS ACROSS THE
EDGE BETWEEN SPRUCE FLAT AND OPENING

Densities and Coverages

Spruce Flat
57 plots

Per
cent

of
Total

Sq. Ft
Coverage
per
Acre

Middle
35 plots

Per
cent

of
Total

Sq. Ft.
Coverage
per
Acre

Opening
42 plots

Per
cent

of
Total

SHRUBS

Alder

Hazelnut

Honeysuckle

Raspberry, dwarf.
Raspberry, red . . .

Sweet gale

Witch hobble

Total.

2.6
14.2
11.3
16.2
13.4

42.2

382
2.056
1,629
2,329
1,938

936
6,084

11.9
7.7
29.0
33.2

18.2

2,609
1,692
6,343
7,255

'3^987

1.0
3.8
4.1
17.8
67.8

"5'.4

15,354

21,886

HERBS

Bedstraw

Clintonia

C^lub-moss, shiny.
Cucumber root . . .
Dogwood, dwarf. .

Fern, beech

Fern, cinnamon. .
Fern, New York. .

Fern, oak

Fern , royal

Fern, sensitive. . .

Fern , wood

Goldenrod, hairy.
Goldenrod, wood .

Goldthread

Grass sp

Hieracium

Jack-in-the-pulpit .

Jewelweed

Maianthemum ....
Miterwort, false. .

Nettle

Nightshade

Partridge berr>'. . .

Pirola

Sarsaparilla

Sedge sp

Solomon's seal ....

Starflower

Thistle

Trillium, painted. ,

Trillium, red

Twisted stalk

Violet sp

Violet, false

Wood sorrel

1.3
2.1

2.5
.3
.5
.1
6

19

Total.

1.7

1.6

.2

.7
3.4
.6
27.6

386
632
351
561

3,047
686
765
742
105
163
35

5,893

70
571
588

"l75
35
982
2,466
163
233
70
140
246
501
233
491

"76

216
1,015

175
8,324

30,124

4.7
1.3
1.0

.8
3.1
3.6
2.8
11.6

.2

20.2
2.8
.2

"5.3

i.5
.2
2.3
12.5

.2

' ". '2
2.0

.2
1.5

.2

10.0
.5
9.5

1,044
286
228
171
702
816
623

2,606
57

4,527
623
57

i , 195

'343
57
514

2,807

" 'l7i
57

""57
457

57
343

57

7.3

.2
1.3
. 7

57

2,240
114
2,120

22,386

15.8

11.0
" ". '9

17

2

3,691

1

0

221

9

190

4

95

9

9

2,128

2

44

4

5

966

1

0

221

1

6

334

21

7

4,650

4

95

2

1

442

4

95

4

95

928

4

95

4

95

TREE PLOT NOS.

3

2

1

1

2

3

4

5

BRUSH a HERB PLOT NOS

9

8

7

6

5

4

J

2

1

1

2

3

4

5

6

7

8

9

10

1 1

12

13

^4

15

1

ALDER

-

J"

2

HAZELNUT

3

HONEYSUCKLE

P

♦

-

•
«

-

•

4

RASPBERRY, DWARF

-

r

-

-

5:

r
1

9

«

•

p

a

5

RASPBERRY, RED

s*

•A

-

r

•A

6

SWEET GALE

7

WITCH HOBBLE

-

-

-

V

W

9

?i

C
fi

-

1

.

r

•

8

BEDSTRAW

•

r

I

9

CLINTONIA

T

♦

1

,p
J.

1

10

CLUBMOSS, SHINY

•

»

-

II

CUCUMBER ROOT

•

.

t
t

12

DOGWOOD, DWARF

_,
*

•

•

♦

t

-

13

FERN, BEECH

t

-

—

— 1

14

FERN, CINNAMON

z

-

t

15

FERN, NEW YORK

16

FERN, OAK

17

FERN, ROYAL

■

IS

FERN, SENSITIVE

t

h.

9

t

it

m

19

FERN, WOOD

« '

I

g

I.

m

\r

20

GOLDENROD, HAIRY

_

1

j

21

GOLDENROD, WOODS

Fig. 14. Graphical analysis of the distribulion of the shrubs and herbs across
the edge between Spruce Flat and Opening.

[103]

Fig. 14 (Continued)

[104]

lidijc llffccl of the Lesser I'eiietation 105

TRANSITION-

SPRUCE FLAT

CENTER

OPENING

TREE PLOT NOS.

3

2

1

1

2

3

4

5

E

)RUSH a HERB PLOT NOS

9

8

7

6

5

4

3

2

1

1

2

3

4

5

6

7

8

9

KD

II

12

13

14

15

42

VIOLET, FALSE

i

43

WOODSORREL

K

Mi

1

i

I

P

Fig. 14 idnu-ludcd)

effective transition in the lesser vegetation was complete within a
zone of 100 feet or less.

1. ]' cgctatk'c Charaeteristics. Sugar maple reproduction was
more abundant than that of any other species at all parts of this
edge. The numbers increased progressively from the opening to the
forest and ])rowse-size seedlings were more abundant near this edge
than at any other northern hardwood edge sampled. The large num-
ber of vigorous young trees in the largest size class even in the
centers of the clearings attests to the shortness of the period that
these clearings remain open. Beech also was relatively abundant,
especially in the largest size class. It is very interesting to note that
not a single individual in the smallest size class was recorded. Al-
though present in relative abundance, yellow birch seedlings were a
poor third in the largest size class, adding further proof of the in-
ability of this species to conii>ete with the others. Balsam was ab-
sent and spruce was scarce, but red maple was present in small
numbers up to the two-foot size class. Striped maple was relatively
abundant but the scarcity of mountain maple was further indication
of its greater success on more rocky and shallow soils. The great
numbers of white ash seedlings may be an indication of a tendency
to take advantage of such openings as well as an indication of the
excellent soil at these particular locations. The large amount of ma-
ture white ash on fornu-r clearings (Ackerman and Catlin) may very
well be proof of the habit of this species to take special advantage of
clearings. Seedlings of black cherry, hop hornbeam, hemlock, and
shadbush were characteristically scanty.

Shrub cover more than doubled in quantity from the forest into
the clearing but there was a complete change of dominants. Witch
hobble was the most abundant species in the forest, while red rasp-
berry and hazelnut, the two next most abundant species, together

AND GROUSE
^HERN HARD-

■a

-o
o
o

Red raspberry
Dwarf raspberry

53 -f.
% 2

tiO O 'Jl t«

3i.lJ

Features Contribute
Opening

oi ^
go

w

Cover

>>

u

u
XI
C

u

oS

O

< W
HO

< a

^<
^>

Features Characteristic of
Both Types

Food

1 ilse niitcrwort

Wood fern
Wood sorrel
Jaek-in-thc-pulpit

Cover

Features Contributed by
Northern Hardwood

Food

-= 13X! =

2. I- x; w
ts 0 Cj
P -^^X!

--2.
MO — .— C

3 O ^ C

Striped maple
(6" high—
1 " diam.)

White ash

Honeysuckle

1 la/.einut

Shiny club-moss

TABLE 30. SUMMARY OF P
PROVIDED liY THE LESS
WOOD AND OPENING

Cover

Beech

(24" high—
r'diam.)

Witch hobi)le

0)

o
c
nj

•a
c

3

XI
nJ

_c

-t->

c

o •

3 ^
O 3

a.

Zj ^
3

til

rio6]

TABLE 31. MATHEMATICAL ANALYSIS OF THE ARBOR-
ESCENT REPKODIXTION ACROSS THE EDGE BE-
TWEEN NORTHERN 1LARD\V(X)I) AND OPENLNG
EXPRESSED IN NUMBERS PER ACRE

Species

Size class*

Northern
Hardwood

Center

Opening

Plot
No. 2

Plot
No. 1

Plot
No. 1

Plot
No. 2

-Sugar maple . . .

24'- 1' diam.
6 '-24"
0"- 6"

2 200
16,350
32^800

1 , 250
10,000
40^590

1,275
6.320
16,430

852
2 618
5 !490

2 333

Beech

24"- 1" diam.
6 "-24"
0'- 6"

1,067
200

700
200

572
233

111

296

Yellow birch . . .

24"- 1" diam.
6 "-24"
0"- 6'

25
25
350

71

333
238

74

296
296

133
67

Spruce

24"- 1 " diam.
6 "-24"
0'- 6"

75

47

1

Red maple

24"- 1 " diam.
6 "-24"
0'- 6"

==^==

200
133

75
300

214
71

37
111

Striped maple..

24'- 1" diam.
6 "-24"
0'- 6'

133
1.133
867

236
572
762

37
37
333

325
775

=====

Mountain
mapl e

24'- 1" diam.
6 '-24"
0"- 6"

25
175

94

Hemlock

24"- 1 " diam.
6 "-24"
0'- 6"

75
50

Black cherry. . .

24"- 1 " fliam.
6 "-24"
0"- 6"

47

24
405
1 ,741

111

37
185

W hite ash

24"- 1 " diam.
6 "-24"
0"- 6"

267
1 .600

50
851

200

1 ,665

Black ash

24"- 1" diam.
6 "-24"
0"- 6"

24

Hop hornbeam.

24"- 1' diam.
6 "-24"
0"- 6"

67
200

47
714

37

167

Shadbush

24'- 1" diam.
6 "-24"
0"- 6'

67

Crataegus

24"- 1" diam.
6 '-24'

50

0'- 6'

* All figures refer to height unless followed by term "diam." which refers to
diameter.

108
TA

No.

1

2
3
4
5

1

2
3
4
5
6
7
8
9
10
11
12
13
14
15
16
17
18
19
20
21
22
23
24
25
26
27
28
29
30
31
32
33

?LE 32. SUMMARY OF THE MATHEMATICAL ANALY-
SIS OF SHRUBS AND HERBS ACROSS THE EDGE BE-
TWEEN NORTHERN HARDWOODS AND OPENING

Densities and Coverages

Northern
Hardwood
55 plots

Per
cent

of
Total

Sq. Ft.
Coverage
per
Acre

Center
42 plots

Opening
37 plots

Per
cent

of
Total

Sq. Ft.
Coverage
per
Acre

Per
cent

of
Total

SHRUBS
Hazelnut

Honeysuckle

Raspberry, dwarf.
Raspberry, red . . .
Witch hobble

10.9

6.6
6.7
22.6
53.2

1,320
801
813
2,749
6.456

8.4
23.4
42.6
25.6

1,728
4,821
8,787
5,285

1.2
37.0
58.4

3.4

Total.

12,139

20,621

HERBS

Beclstraw

Cljntonia

Club-moss, common.
Cluh-moss, shiny....

Cucumber root

Fern, bracken

Fern, cinnamon

Fern, hay-scented.. .

Fern , New York

Fern, oak

Fern, sensitive

Fern , wood

Goldenrod, hairy. . . .

Goldthread

Grass sp

Jack-in-the-pu:pit. . .

Jewelweed

Maianlhemum

Miterwort, false

Nettle

Nightshade

Partridge berry

Pirola

Hypericum

Sarsaparilla

Sedge sp

Solomon's seal

Starflower

Strawberry

Trillium, painted. . . .

Twisted stalk

Violet sp

Wood sorrel

1.6
1.3
1.0
8.6
1.3

182
145
109
942
145

73
36

16.7

" .7
1.5
9.2

. I
11.4

1.0
1.0

2.9
15.2

i.6

.3
1.3
10.6
11.2

Total.

1,836

'73
169
1.014

73
1.210

109
109

314
1,672

"182

" ' 36
145
1,161
1,233

10,968

3.0

1.8
1.1

"t.2
8.5

.3
10.8
1.3

524

316
190

.3
.3

16.4
.3
2.4
.5

.3
13.8
7.9

1,078
1,479

48
1,867
221

364
381
286
190
2.750
48
143
48
48

"143
2,851

48
411

95

48
2,404
1,377

17,358

4.1

.8
19.0
5.1

.9
.2
13.3

13.1
4.9

3

8

1.178

5

2

1.597

8

251

2

54

4

0

1,218

1

9

594

6

8

215

2

54

12

4

3,852

"2

7

846

108

4.050
1,522

28,964

(L>

U c

-a
o
o

o

u

O ^

dj b o

O S

2 S

rt >.

J2

° a.

o
U

St/)

o
o

:r °^

"t: -G -T3

n o o
o o

o

I

o

S a I ^'S^i^^ : o o

— — r- ^ O 5^-^ t/^ C

rt cj <u o oi^; rt— i2

be rt

o ^ o
-rt-^SS

?N ^ 'X —

3 vO

01 ^

3 i2

■■5 3

<D 3

c rt

[109]

110

Roosevelt Wildlife Bulletin

constituted less than lialf as much cover. The oi>ening, however, was
dominated by a rank growth of red raspljerry, underlain in the moist
spots with dwarf raspberry and very lightly interspersed with witch
hobble.

The total herb cover also increased from the forest mto the
opening, nearly tripling in amount. Most species on the forest side
were scattered sparingly, wlrile dense patches of several of the same
species characterized the opening. The most abundant species on
the forest side were wood fern, sedge, wood sorrel, false miterwort.
and violet, while in the opening the herb cover was characterized
by larger patches of sedge, violet, false miterwort. and in moist places
cinnamon fern and jewelweed.

2. Wildlife J^alites. The most abundant sjx-cies of food and
cover contributed by each portion of the edge are listed in Table 30.
From this it is evident that unlimited amounts of buds were pro-
vided for winter food by the mature hardwoods of the forest, as
well as abundant browse in the form of witch hobble and arborescent
reproduction. The usual summer cover of witch hobble was supple-
mented by a limited amount of fall cover in the form of young beech,
but winter cf)ver was noticeably absent. Small amounts of hazelnut
snd honeysuckle augmented the variety of fruit, and shiny club-moss
was present as a year-round green.

The opening side of the edge also provided abundant summer
cover, but in the form of red raspberry rather than as witch hobble.
This also served as an important deer browse as well as an important
source of summer fruit. Uwarf raspberry provided a much smaller
amount of both types of food, but was of no value as cover. Sedge,
jewelweed, bedstraw, and grass supplemented both the quantity and
variety of summer food on this side.

Species common to l)Oth sides of the edge provided small
amounts of spring and summer greens in the form of violet and
jack-in-the-pulpit and liberal amounts of overwinter greens in the
form of false miterwort, wood fern, and wood sorrel. The jack-in-
the-pulpit, of course, also provided a small amount of fruit. Several
other species of herbs, occurring in small quantities, supplemented
the variety of food and in some instances the cover.

It appears that this edge provides an excellent variety of summer
food and cover both of which are lacking in the winter. Although
buds are plentiful, fall and overwintering fruits are entirely absent,
as is escape cover after the leaves have fallen. (See Fig. 15 and
Tables 31 and 32.)

TRANSITION-

NO. HARDWOOO CENTER

TREE PLOT NOS.

2

1

1

z

3

BRUSH a HERB PLOT N0&

6

5

4

3

2

1

1

2

3

4

5

6

7

e

9

J J

-

«

■

1

HAZELNUT

■MM

2

HONEYSUCKLE

1

I

-

_

r

«

1

«...

, ....

4

RASPBERRY. RED

t.

IT

ii

1

T

ri

1

T

*

H

-

«.

H-

•

5

WITCH HOBBLE

m

a

r

t

4

I

•

6

tl&U3 I nAVf

*

7

CLINTONIA

8

CLUBMOSS, COMMON

9

CLUBMOSS, SHINY

*

■

10

CUCUMBER ROOT

i

L i

1 1

PPPU ROAri^CM

—

•S

12

'»5

13

rcnn^ nAvovC^'tu

•

14

PERN, NEW YORK

i

15

FERN, OAK

— !

16

FERN, SENSITIVE

r

s

r.

»i

•

f

t

1

P

i

1^

17

FERN, WOOD

s

lb

OPENING

Fig. 15. Graphical anahsi.s of the (Hstrihiition of the
shnih.s and herbs across tlie edge between Northern Hard-
wood and Opening.

[Ill]

TRANSITION-

NO HARDWOOD

CENTER

OPENING

TREE PLOT NOS.

2

1

1

2

3

BRUSH a HERB PLOT NOS

6

3

2

1

1

2

7

8

9

I—.

18 1 GOLOENROD, HAIRY

-

19

GOLDTHREAD

*

20

GRASS SR

—

X

*

f

•

•

t

i

•

21

OACK-IN-THE-PULPIT

u
1

«

•

•

i

•-.

22

JEWELWEED

23

UAIANTHEMUM CAN.

*

•

I

1.

i

V

S

- jp

i

1

a

•J

1

•

M

24 j MITERWORT, FALSE

25 1 NETTLE

\

♦

J

•

.1:

t

:

26

NIGHTSHADE

1

i

27

PARTRIDGE BERRY

1

1

28

PIROLA

29

HYPERlCim

30

SARSAftVRILLA

■

«
•

•

.

•*

C

:

B

31

SEDGE SP

•

a

•

32

SOLOMON'S SEAL

33

STARFLOWER

i

-H

34

STRAWBERRY

Fig. 15 (Continued)

1112 1

Edge Effect of the Lesser Vegetation

113

TRANSITION-

NO HARCWOOO

OPENING

TREE PLOT NOa

2

1

2

3

BRUSH a HERB PLOT NOS

6

5 |4

3

2

1

1

2

3

4

5

6

7

8

9

r

35

TRILLIUM, PAINTED

•

» :

36

TWISTED STALK

•

\

•

k

m

I

37

VIOLET SR

\

•

* ^

%

.»

t

«

I r
r

<t

a

_

\

38

WOODSORREL

•

1

t

f-

u

I

2l

1

«

Fig. 15 (Concluded)

J. The Edge Between Spruce Flat and Open Swamp. All

of the few optn swamp types on the Check Area are a resuh of
beaver flooding of previously wooded areas; they are located on
low, spruce flat climax sites. The trees have been killed and the
vegetation has taken on the aspect of open sedge-covered marsh with
the water level fluctuating seasonally from about two feet above the
surface to slightly below the surface. The total area of the open
swamp types is 44.6 acres. During the past few years the largest of
these marshes (K-9 Swamp) has become partly overgrown with
speckled alder. Most of the data in reference to the edge between
open swamp and spruce flat were secured along the south shore of
this large swamp. The data were collected from a series of seven
transects composed of a total of 166 underbrush and herb plots of
which 72 were in spruce flat, 42 at the center of transition, and 52
in the open swamp. The average length of the transects was 295
feet, but the actual transition in the lesser vegetation was complete
in most cases within a zone of 75 feet.

1. Vegetative Characteristics. The amount of lesser vegetation
at this edge was somewhat less than might be expected, due probably
to the dense shade created by the abundance of pole-size conifers
and the low crowns on most of the trees at the edge of the forest.
Thus, the tallied size classes of reproduction were all relatively light.
Balsam was especially scarce in the largest tallied size class, probably
as a result of concentrated deer browsing. Red maple and yellow
birch were the next most abundant, but they were relatively few as
compared to other edges. The occurrence of other species was ex-
tremely low.

114

Roosevelt IVildlije Bulletin

Total shrub cover increased nearly 50 percent from the spruce
flat into the swamp. The most abundant si)ecies on the forest side
were red raspberry, dwarf raspberry, honeysuckle, and witch hobble.
The relatively small amount of witch hobble probably was due to
the dense cover of conifers. Species most abundant on the swamp
side of the edge were sweet gale, meadowsweet, and alder, but the
amount of alder has increased greatly since these data were gathered
and now dominates not only the margin of the swamp but also most
of the central portion.

The total amount of herbaceous cover increased but slightly
from the forest to the open swamp. Whereas the herb species in the
forest were more uniformly abundant, three-fourths of the herba-
ceous cover in the swamp was composed of sedges and grasses. These
were supplemented by small amounts of other species, chief among
which were violets, wood sorrel, sensitive fern, goldthread, nettle,
and St. John's-wort. On the spruce flat side, dwarf dogwood, wood
fern, and wood sorrel were about equally abundant and there was a
greater variety of si)ecies of intermediate abundance such as violet,
sedge, false niiterwort, grass, beech fern, false lily-of-the-valley,
goldthread, clintonia, and starflower. The total number of species
of both underbrush and herbs across the entire transition was greater
than that at any part.

2. Wildlife Values. The more abundant vegetative features of
value to wildlife are listed in Table 33. From this it is apparent that
there were a relatively large number of moderately abundant food
and cover plants at this edge. The spruce flat contributed ample ex-
cellent year-round cover and winter buds, as well as moderate
amounts of a variety of summer fruits : honeysuckle, red raspberry,
blueberry, false lily-of-the-valley. and clintonia. Abundant overwin-
tering greens were provided by wood sorrel, wood fern, false miter-
wort and goldthread, while early spring greens were furnished by
false lily-of-the-valley, violet, and a small amount of dogtooth violet.

The open swamp side of the edge probably is more valuable to
deer than to grouse. Sedge and grass furnished early summer
browse, the several herbs and the abundant aquatic plants in the
streams and sloughs provided midsummer food, while young balsam
and the various species of underbrush provided fall and winter food.

Although most of the food and cover requirements for deer are
provided in the vicinity of this edge, fall and winter fruits are lack-
ing for grouse. The total amount of blueberry, honeysuckle and rasp-
berry was greater at the center of transition than in the adjacent
forest, but the swamp plants did not greatly improve the site for the
last species. (See Fig. 16 and Tables 34 and 35.)

TRANSITION-

SPRUCE FLAT CENTER OPEN SVMtWP

TREE PLOT NOS

3

2

1

1

2

BRUSH 8 HERB PLOT NOS.

9

8

7

6 5

3

2

1

1

2

1

6

ALDER

-

-

1"

1

BLUEBERRr

•

•

2

J

HAZELNUT

**

-

•

4

HONEYSUCKLE

**

-

-

-

•

—

•

1

•

LABRADOR TEA

S

LEATHER LEAF

•

6

MEADOW. SWEET

r

—

7

M.

p-

T

•

8

RASPBERRY. DWARF

t

\

•■

9

RASPBERRY. RED

L

-

t

"7"

10

WILLOW

8

II

SWEET GALE

12

WITCH HOBBLE

-

•

13

BEDSTRAW

•

T"

•

•

14

CLINTONIA

*

1

t

-

•

•

15

CLUBMOSS, COMMON

16

CLUBMOSS, SHINY

•

17

CUCUMBER ROOT

•

18

DOGWOOD, DWARF

\

r

1^
«1

I-

-

•

•

1

19

FERN, BEECH

•

-

20

FERN, BRACKEN

21

FERN, CINNAMON

22

FERN, NEW YORK

■r"

23

FERN, OAK

24

FERN, SENSITIVE

•w-

25

FERN, WOOD

m

r
t.

TT

i

8

k

-

-

26

GOLDENROD. HAIRY

27

GOLDTHREAD

28

GRASS (MARSH)

•

-

-

';

-

29

JACK-IN-THE -PULPIT

1 —

+

1

30

JEWELWEEO

1 1

1

31

MAIANTHEMUM CAN

"r-

1

!
t

32

MITERWORT, FALSE

T~

It

33

NETTLE

34

NIGHTSHADE

35

PIROLA

Fig. 16. Graphical analysis of the distribution of the
shrubs and herbs across the edge between Spruce Flat
and Open Swamp.

[115]

TRANSITION

SPRUCE FLAT

|C£NTER

OPEN SVMAMP

TREE PLOT NOS.

I

BRUSH a HERB PLOT NOS

36 HYPERICUM

SARSAPARILLA

38

SEDGE SP

39

SNOWBERRY

SOLOMON^ SEAL

STARFLOWER

43 THISTLE

44 TWIN-FLOWER

TWISTED STALK

VIOLET SR

VIOLET. FALSE

fcliTT

48 WOODSORREL

Fig. 16 (Concluded)

[116 1

117

TABLE 34. MATHEMATICAL ANALYSIS OF THE AR-
BORESCENT REPRODUCTION ACROSS THE EDGE
BETWEEN SPRUCE FLAT AND OPEN SWAMP EX-
PRESSED IN NUMBERS PER ACRE

Species

Size Class*

Spruce Flat

Center

Open Swamp

Plot
No. 3

Plot
No. 2

Plot
No. 1

Plot
No. 1

Plot
No. 2

Spruce

24"- 1" diam.
6 "-24"
0"- 6"

333
333

167
250
833

143
810
1,190

619
323
143

48

200

Balsam

24"- 1" diam.
6 "-24"
0"- 6"

47
371
858

191

3,475

48
238

1,000
800

2,333

2,750

Red maple . . .

24"- 1" diam.
6 "-24"
0"- 6"

2,667
1,667

83
1,167
2,800

48
371
1,000

47
572
288

143

Hemlock

24"- 1" diam.
6 "-24"
0"- 6"

48

Sugar maple.

24"- 1" diam.
6 "-24"
0"- 6"

'416'

' "95
2,500

48
333

48

Beech

24"- 1" diam.
6 "-24"
0"- 6"

167

83
48

Yellow birch .

24"- 1" diam.
6 "-24"
0"- 6"

333
167
667

810
1,250

190
1,095

190

800
48

Striped
maple

24"- 1" diam.
6 "-24"
0"- 6"

250
333

167

Mountain

24"- 1" diam.
6 "-24"
0"- 6"

333

"'83
583

478
428

286
285

95

Shadbush

24"- 1" diam.
6 "-24"
0"- 6"

48

95

143
48

Mountain ash

24"- 1" diam.
6 "-24"
0"- 6"

143

* All figures refer to height unless followed by the term "diam." which refers to
diameter.

118

TAP.l.E 35. SUMMARY OF THE MATHEMATICAL ANALY-
SIS OF THE SHRUBS AND HERBS ACROSS THE
EDGE BETWEEN SPRUCE FLAT AND OPEN SWAMP

Densities and co\erage

Spruce Flat
36 plots

Per
cent

of
Total

Sq. Ft
Coverage
per
Acre

Center
21 plots

Per
cent
of

Sq. Ft
Coverage
per

Total Acre

Ojjen Swamp
26 plots

Per
cent

of
Total

SHRUBS

Alder

Blueberry

Hazelnut

Honeysuckle

Labrador tea

Leather leaf

Meadow sweet . . .
Raspberry, dwarf.
Raspberry, red . . .

Willow

Sweet gale

Witch hobble

3.9
9.0
10.9
18.3

27,0
17.4

13.5

388
909
1,096
1,838

2,715
1,744

1,356

7.4
13.4

1.6
19.6

2.4
27.7
19.0

's.i

5.3

891
1,623

190
2,365

236
3,351
2,290

446
636

12.4

3.

18.

11.3
3.1
6.4

31.8

10.5

Total.

10,046

12,028

HERBS

Bedstraw

Clintonia

Club-moss, common.
Club-moss, shiny. . . .

Cucumber root

Dog\\'ood, dwarf. . . .

Fern, beech

Fern, bracken

Fern, cinnamon

Fern, New York

Fern, oak

Fern, sensitive

Fern, wood

Golden rod, hairy. . . .

Goldthread

Grass (marsh)

Jack-in-the-pulpit. . .

Jewelwced

Maianthemum

Miterwort, false

Nettle

Nightshade

Pirola

Hypericum

Sarsaparilla

Sedge sp

Snowberr>-

Solomon's seal

Sphagnum

Starflower

Thistle

Twin- flower

Twisted stalk

Violet sp

Violet, false

Wood sorrel

1.0
1.8
.5

' '. 2
21.4
3.6

333
611
167

.8
.2
1.0
.5
19.2

2^8
3.4
.2
.3
3.1
4.9

Total.

.3
5.7
.5
.2
1.7
2.4

.8
.5
4.2
1.0
17.7

56
7,281
1,232

"258
56
333
167

6,531

960
1,162
56

111
1,055
1,677

56

111

1,948
167
56
571
833

' "278
167
1.440
333
6,024

34,030

1.4
2.3
.2
.2
.2
12.8
2.2
1.4
.2

571
918
95
95
95
5,046
884
537
95

1.2
.2

.4

\A
1.0

1

4

537

8

3

3,282

i

8

728

3

8

1,516

2

95

1

1

442

1

9

762

5

190

4

1

1,611

2

95

5

190

7

286

27

5

10,855

2

95

1

1

442

6

9

2,708

1

4

571

2

95

2

1

823

2

95

5

2

2,051

2

95

9

0

3,533

39,433

2
1

2
11

2.0
63^1
' .2
.2

5.9

I 37,056

Edge Effect of the Lesser J^egelolioii

119

VI. COMPARATIVE VALUES OF THE VARIOUS EDGES

The comparative coniploxity and dcnsily of ihe vegetation at the
various edges are summarized in T^ible 36. This, however, is offered
only for special reference for the sake of tliose interested in further
analysis. Certain specific trends have l)een isolated from these and
previous data and are presented in graph and list form in this section.

More attention will be given to the distribution of grouse than to
the distributit)n of deer. The principal interest is grouse because of
their apparent scarcity in what is considered to be the center of their
eastern range (the .\dirondack region). Although this scarcity is
undoubtedly related to a multiplicity of factors, one cannot help but
feel that the vegetation, basic as it is to the needs of the species,
must play an important part in the abundance of the bird. At any
rate, the vegetation at the edges has been analyzed in rather con-
siderable detail and the summary of that analysis is presented in an
attempt to throw some light on the reason for the .scarcity of this
species. There is no such scarcity of deer ; rather, as will be indicated
below, it would appear that the deer (together with mice) may be the
principal competitors of grouse for food, and due to their numbers
may have caused a significant deterioration in tlie range for grouse.

A. Comparison of the Vegetative Characteristics. The rela-
tive variety and abundance of the lesser vegetation at the various
edges is summarized in Figure 17 and Table 36. From these illus-
trations several over-all relationships become clear. It is obvious that
although there is a great deal of variation in the abundance of the
three different forms of vegetation, there is not nearlv as much vari-
ation in number of species. Of striking significance is the very scant
variety of species occurring with sufficient abundance to be .sampled.
Eighteen species of trees were recorded but the tree cover at any
particular site usually consisted of not more than six species : sugar
maple, beech, yellow birch, spruce, balsam fir, and red maple. All of
these species except spruce serve as important sources of food and
cover for grouse and deer.

The variety of shrub species was even more limited. Although
18 species were recorded, the variety at any edge usually was le: s
than that of the trees or reproduction. Four species composed the
great bulk of shrub cover : witch hobble, red raspberry, dwarf rasp-
berry, and honeysuckle. Of these, only witch hobljle and red rasp-
berry normally bear a1)undant fruit, which usually is shed by early
fall.'

As would be expected, the herbs were present in greatest varietv.
Forty-nine species were recorded, and alxnit lialf of them were .suffi-

120

TABLE 36. SUMMARY OF THE PHYSICAL AND VEGETA-
TIVE CHARACTERISTICS OF THE TEN EDGES

Edge

Repro-
duc-
tion

Underbrush

Herbs

Aver-
age
length

No.
of
species

No.
of
species

Average
Cover-
age
sq. ft.

jjer acre

No.
of

species

Average
Co\er-
age,
sq. ft.

per acre

1. Bracken Fern to Aspen. . .

225

10

9

7,000

29

47,000

2. Bracken Fern to Second
Growth

350

12

13

10,500

31

36,600

3. Second Growth to North-
ern Hardwood

225

13

6

2,700

25

7,300

4. Spruce Flat to Mixed
Hardwood

375

11

6

13,600

29

13,600

5. Spruce Flat to Northern
Hardwood

450

16

12

13,900

36

15,500

6. Mixed Hardwoodto North-
ern Hardwood

375

12

6

14,200

30

13,100

7. Spruce Flat to Spruce
Swamp

less
than
100

14

10

10,600

32

33,000

8. Spruce Flat to Opening.. .

less
than
100

16

7

21,400

36

25 , 100

9. Northern Hardwood to
Opening

less
than
100

13

5

19,500

33

19 , 700

10. Spruce Flat toOpcn Swamp

less
than
100

11

12

12,400

36

37,800

Ed<jc Effect of the Lesser Vegetation

121

ciently abuiulant to be termed characteristic. The species most widely
distributed in greatest numbers were wood fern, wood sorrel, violets,
false lily-of-the-valley, false miterwort, dwarf dogwood, sedges,
grasses, bedstraw, startlower, clintonia, cucumber root, and sarsapa-
rilla. It is of especial significance that of these species, only dwarf
dogwood normally produces abundant fruit in this region, and that
it is limited in distribution to coniferous or partially coniferous loca-
tions.

The bulk of the fruit and seed of value to grouse (on the Check
Area) is borne by only 19 of the 761 species of vascular plants found
on the Forest. Five of these species occur at all edges in sufficient
abundance to supply a considerable share of the grouse food. Six
other species occur in like abundance in all but one edge, three occur
in all but two edges, and one in all but three edges. The other four
species are widely distributed but they occur in significant quantities
only at from one to six edges. The fact that this study was confined
to the Check Area does not alter this condition seriously, for it is
believed that even if the more open portions of the Forest were in-
cluded in the study, the great bulk of all fruit and seed would be
produced by not more than about 25 species.

Another interesting fact is that these fruit- and seed-bearers
are not restricted to certain sites, i.e., it could be said that each of
these staple food-bearers is characteristic of the entire vegetation
and each occurs at all but a very few places. Stated still another
way, we can say that on the whole there is no "special" group of
fruit-bearers restricted to each particular site or edge. To be sure,
there are several species, such as mountain holly, winterberry and
wintergreen, that occur only on certain restricted sites, but their
distribution is so scattered or scant they cannot be considered as
being important in the general economy of the rufifed grouse.

Although the more abundant species mentioned above probably
serve as the staple supply of fruit and seed for grouse, and are there-
for very important in the present arrangement of the vegetation,
the distribution of other fruit-bearers occurring in lesser quantities
may prove of significant potential value. Furthermore, most of the
currently "staple" fruit-producers shed their fruit by early fall, while
a considerable number of the "minimum" fruit-producers retain their
fruit well into the fall and winter. Among these latter are such
species as mountain holly, winterberry, wintergreen, snowberry, par-
tridge berry, hazelnut, and wild raisin.

Based on the total variety of species occurring in sufficient quan-
tities to be sampled, the various edges may be listed as follows :

122

Roosevelt WUdlife Bulletin

1. Spruce Flat — Northern Hardwood (64 species)

2. Spruce Flat — Opening (59 species)

3. Spruce Flat — Open Swamp (59 species)

4. Spruce Flat — Spruce Swamp (56 species)

5. Open Bracken — Second Growth Hardwood (56 species)

6. Northern Hardwood — Opening (51 species)

7. Mixed Hardwood — Northern Hardwood (48 species)

8. Open Bracken — Aspen (48 species )

9. Spruce Flat — Mixed Hardwood (46 species)

10. Second Growth Hardwood — Northern Hardwood (44 spe-
cies.

Although this list is admittedly the easiest one to compile, the
reader may question its value because it fails to take into considera-
tion the cover or food value of the species recorded. Since the
general nature of the mature vegetation is that of a dense forest,
with fairly abundant underl)rush, we can scarcely consider cover to
be the principal limiting factor in the grouse population over most
of the range. Furthermore, there is a comparative abundance of
greens at all seasons when not covered with snow and also a limitless
supply of winter buds. The only vegetative requirement not present
in comparative abundance is fruit- — especially that of late fall and
early winter. If then, we arrange the list of edges in order of de-
creasing variety of fruit- and seed-bearing species, the following
sequence results.

1. Spruce Flat — Northern Hardwood (31 species)

2. Open Bracken — Second Growth Hardwood (30 species)

3. Second Growth Hardwood — Northern Hardwood (27 spe-
cies)

4. Spruce Flat — Spruce Opening (26 species)

5. Mixed Hardwood — Northern Hardwood (25 species)

6. Northern Hardwood — Opening (25 species)

7. Spruce Flat — ]\lixed Hardwood (24 species)

8. Spruce Flat — Spruce Swamp (24 species)

9. Spruce Flat — Open Swamp (23 species)
10. Open Bracken — Aspen (23 species)

It is obvious from the data that some seed- or fruit-producing
species in the list occur in such small quantities as to be considered
unimportant in the over-all economy of the grouse, especially when
that species is in competition with much more intensive foragers such
as wood mice. Just how abundant a species of food must be in order
to be of value to grouse is difficult to say. We can assume, however,

Edge Effect of the Lesser Vegetation 123

that the more readily available species would be of greater impor-
tance in the economy of grouse than the scarcer ones. Hence it fol-
lows that it would be more proper to consider the various edges in
regard to the variety of species characteristically occurring in quan-
tities larger than slightly scattered "trace" elements. The edges on
this basis may be listed as follows :

1. Spruce Flat — Opening (20 species)

2. Northern Hardwood — Opening (19 species)

3. Spruce Flat — Northern Hardwood (17 species)

4. Mixed Hardwood — Northern Hardwood (17 species)

5. Spruce Flat — Open Swamp (17 species)

6. Spruce Flat — Alixed Hardwood (15 species)

7. Spruce Flat — Spruce Swamp (15 species)

8. Second Growth Hardwood — Northern Hardwood (18 spe-
cies)*

9. Open Bracken — Second Growth Hardwood (15 species)
10. Open Bracken — Aspen (11 species)

When these three lists are compared, it is obvious that no corre-
lation exists as to their arrangement, i.e., the edges are completely
shuffled so that we cannot say, for example, that the greatest amount
of fruit is to be found at those edges where there is the greatest
variety in the lioristic composition. Because there are more species
of plants at a given edge, or even more species of fruit-producing
plants, it does not follow that there is more food or fruits at that
edge. Thus, although one would like to hold to the old cliche that
wildlife is a product of edges, the evidence indicates that this prin-
ciple may not apply when speaking of certain regions.

As indicated above, the edges did not vary as much in total
number of species as they did in respect to the relative amounts of
each of the three forms of vegetation (herbs, shrubs, and arborescent
reproduction). For example, the herb cover was nearly seven times
as dense at the Open Bracken — Aspen edge as that at the nearby Sec-
ond Growth Hardwood — Northern Hardwood edge (47,000 square
feet per acre as opposed to 7,000). The relative amounts of herbs
and shrubs at the other eight edges as shown in Figure 17 vary
between these two extremes.

The various edges could be easily listed on the basis of total
amount of lesser vegetation by consulting Figure 17. However, it has
already been shown that this would not be a proper listing of wild-

* In this case, although the variety would place this edge higher in the list,
so many of the fruit-hearers were so near the border line of productivity that
it was necessary to phice tlie edge lower in the list.

Ki.,^Ullu:..u^>ilb utrtNDtNT
UPON VEGETATION

RELATIVE ABUTOANCE AT EACH EDGE

Bracken Fern
Aspen

Bracken Fern
Second Growth

Second Growtn
Northern Hardwood

Spruce Flat

:iixed Harawood

-o
o
o

d ie
O

o

Lc

£

V

a a

X u

^ §.

Spruce Flat
Nortnern Harawooa

Spruce Flat
Spruce Swamp

a
-\

'■^

a. Qi

-I o

g
o
s

T3

•a

a i

u u

0 a
z o

Spruce Flat

Open r.wanip

a

Early Greens

DrMairaint. LorS

a,

CO

Cover for Adults

Cover for Younj;

Fr-uits

&

(O

Greens ki^lHB—

Buds

Cover

1 WINTER

BUQS

Greens

Fruits

uover

LEGEND 1 1 — > wmm '^^B

None Present ^oxign Enough C

irer abundani

Fi«. 17. Coinparalive value of tlie vegetation at the various edges for ruft'ed grouse, as
based on general major requirements.

[ l-'4 ]

Edge Effect of the Lesser ]^ eyetalion

125

life value since other considerations must be taken into account.
For example, if the list is to be applied to young grouse, one must
automatically omit such hazardous sites as spruce and open swamp
types. Thus, on the basis of spring cover density usa])le by young
grouse, tlie edges ma}- be listed as follows :

1. Open Bracken — Aspen

2. Open Bracken — Second Growth Hardw ood

3. Spruce Flat — Opening

4. Northern Hardwood — Opening

5. Spruce Flat — Northern Hardwood

6. Spruce Flat — Mixed Hardwood

7. Mixed Hardwood — Northern Hardwood

8. Second Growth Hardwood — Northern Hardwood

Late summer co\er and that of earh" fall probably ct)uld not be
considered as a limiting factor once the young grouse are able to fl\',
hence a listing of relative amounts of cover at these seasons would
be of no value. However, it may well be that when the deciduous
leaves have fallen cover may play an important part in the winter
and early spring distribution of grouse. At these seasons the first
two sites in the above list would be moved to the bottom and conif-
erous types would take the lead. The spruce swamp type would then
be added to the list and placed near the top. However, the value of
the hardwood types should not be overlooked because of two con-
ditions : ( 1 ) scattered patches of beech reproduction in these types
provide escape cover through most of the year, and (2) although
composed principally of deciduous species, they nevertheless do con-
tain scattered clumps of conifers which probably serve very well as
winter cover.

Figure 17 represents an attempt to combine the distribution of
the significant amount of grouse habitat requirements dependent
upon vegetation into a single picture. The distribution of these re-
quirements is shown by edges and by seasons and it approaches as
near as possible to an over-all graphical illustration. The abundance
classification is arbitrary and hence subject to some question. How-
e\er, it cannot be otherwise until we know more about the specific
requirements of the species. It would be especially valuable to know
the specific minimum amount of a given species of food that must be
present in order to be of value to a given species of wildlife.

It is apparent from Figure 17 that although certain edges fur-
nish abundant amounts of certain requirements, they are seriously
lacking in others, either on a seasonal or year-round basis. Tiius
for example, altiiougli the edges of open bracken were bsted as

126 Roosevelt ll'ildlije Bulletin

excelling in summer cover for young grouse, it becomes apparent
thai this cover lasts only a short period. If bro(j(ls emerge before the
middle of June, as they undoubtedly do, this type would offer no
protection either from the bright sun or from predators. Further-
more, the absence of suitable drumming facilities precluded a satura-
tion population of mating males, and the absence of fall fruits con-
fined the adult birds to an almost continuous diet of leaves and buds.
For these reasons the open bracken type, its edge toward aspen and
its edge toward seccnid growth hardwood, must be considered as
falling far short of ideal grouse range. Although somewhat better in
regard to the above-mentioned deficiencies, the edge between second
growth hardwood and mature northern hardwood is not much of an
improvement. A few logs of apprcjpriate size and condition for
drumming are present but they lack the necessary cover. Cover for
young birds likewise is scant, consisting only of hardwood litter over
much of the second growth type but supplemented by relatively small
amounts of herbs and shrubs in the adjacent mature hardwood type.
P"urthermore summer fruits are scarce and fall and winter fruits are
practically absent. Most of the value of this edge derives from the
mature hardwood type, with very little being provided by the second
growth.

The three extensive climax types and the edges between them
and their openings come nearest to fulfilling the vegetative require-
ments of ruffed grouse. They are so much alike in this respect that
it is difficult to arrange them in order or apparent over-all value.
However, each is deficient in late fall and early winter fruits. Ground
cover for young birds is much better in the hardwood types as is the
supplv of buds for winter feeding. Winter cover is especially good
in the coniferous types and, as a whole, the variety of fruit-bearing
species is greater, but the supply of buds for winter use is neither
as abundant nor as varied as in the hardwoods. If the extreme lack
of fruits can be assumed to be one of the limiting factors in the
present population, then edges between these types should be of
especial value. Furthermore, openings in these types also would be
of great value since these openings normally contain dense patches
of red raspberries which bear abundantly almost every year. How-
ever, these fruits usually are gone by the end of the summer so that
fall and early winter fruits are lacking even at their edges.

The swamp types and their edges remain to be considered. The
dense cover of the spruce swamp provides excellent winter cover
but it contains practically no food. Furthermore, the marshy condition
must be considered a hazard to young birds and hence must be
-ivoided in the spring season. Hardwood tyi)es offer equally good

Edge Effect of llie Lesser Vegelailun

127

summer cover l)Ul more food in the form of both greens and fruits.
The open swamp type ofifers large amounts of fall seeds (sedges)
but when these are ripe the cover becomes less effective and birds
expose themselves to avian predation. Furthermore, unless the
swamps have become very dry the birds probably would avoid the
normally wet and marshy substrate. 'Jdie principal feature of the
open swamp type which may be considered of esi)ecial value for
grouse is the fact that several species of shrubs bearing persistent
fruit fre(|uently are found at their margins as well as in their in-
teriors.

Since any wildlife habitat improvement plan must be limited to
a large extent by economic considerations, it follows that improve-
ment practices should be initiated in those places where a unit of
effort will produce maximum results. It becomes apparent, therefore,
that on any range characterized by a relatively low population of the
species being managed, any effort would result in the greatest imme-
diate return if aj^plied in those places where a minimum number of
requirements of the species are lacking. For the most immediate
returns, only tho.se sites which already provide a maximum number
of requirements should be improved. The above analyses have dem-
onstrated that sites most nearly fulfilling these requirements are the
three most extensive forest types and the edges between them (spruce
flat, mixed hardwood, and northern hardwood) and that the principal
apparent deficiency is the lack of fall and early winter fruit. It has
been shown that openings in these types normally provide abundant
supplies of summer fruit. Since the northern hardwood and spruce
flat types more nearly complement one another in fulfilling the re-
quirements of grouse than does association with the mixed hardwood
type, improvement practices would appear logical at the edge l)et\veen
these two types or at places where they come close together. Al-
though the mixed hardwood type does contain a number of elements
common to both spruce flat and northern hardwood, a greater range
of food and cover is available in the two types.

B. Delimitation of Concepts Relative to Edge Effect. Before
going into the final analysis of the relative values of the varicnis edges
and the degree of edge preference shown by grouse and deer for
these edges, it would be well to re-define and delimit the various con-
cepts in relation to edge effect. One new concept and one new term
in relation to edge eft'ect will be introduced and defined in relation
to previously introduced concepts.

The terms juxtaposition and threshold acreage have l)een coined
(King, '38; Beecher, '42) to describe various concepts in relation to

128

Ruoscvcll li'ildlijc Bnllclin

edge effect. Unfortiiiialely, there has been some subsequent confu-
sion in the use of both terms (as well as "edge effect" itself) ; often
they have Ix-en used interchangealjly (jr inai^projiriately, sometimes
with httle regard for the type of edge or the sjx'cies of wildlife con-
cerned. Furthermore, there can be little doubt but that we have to
some extent overplayed the entire concept of edge effect, often in-
discriminantly ascribing to it unlimited powers of increasing all
forms of wildlife. The art of wildhfe management has reached a
stage where it would be wiser to speak more specifically when re-
ferring to edges. The types of edges involved and the species of
wildhfe to be benefited should be made clear. It cannot be over-
enii)hasized that the concept of edge effect is of no value unless used
in reference to specified forms of wildlife for, as pointed out by
Beecher (I.e.), some edges actually repel rather than attract certain
species of wildlife. \\\\\\ these thoughts in mind, the writer has
attempted to re-define and delimit the connotations of the above-men-
tioned concei)ts in relation to edge effect (juxtaposition and threshold
acreage), and has also presented a new concept, namely, "threshold
of edge preference." Furthermore, it is suggested that the term
"edge preference" be used to cover certain situations previously
groui)ed under the term "edge eflfect." This would put an end to
the dual use of this latter term both as a general and specific term,
leaving it free to be used only as a general term, covering all con-
cepts in relation to diverse habitat requirements of wildlife species.

Before going into these definitions it might be well to consider
for a moment a classification of edges in relation to the nature of
the vegetation. On this basis, edges may be divided into two gen-
eral groups :

A. Edges composed only of the intermingling or the juxta-
position of the species of the two adjacent cover types.

A few examples of tliis kind of edge may be listed as
follows :

1. The edge between two crop fields, with no separating
border.

2. The edge between an orchard or plantation and the
surrounding fields, where no special border vegetation
has developed.

3. The edge between two forest types.

B. Edges composed not only of the borders of the two ad-
jacent plant associations, hnt also characterised by a "sf^c-

Edijc Effect of the Lisscr I 'c</ctatioii

129

ciai" edge I'cgctalioti. Examples of this type of edge may
be listed as follows :

1. The edge between a forest and an open field where
there is a border of shrnl)l)y vegetation.

2. The edge between crop fields separated by hedgerows.

3. A woods road, with its "six'cial" or "characteristic"
species.

4. Narrow zones of streambank or lakeshore vegetation.

Additional examples of both of these types of edges undoubtedly
will occur to the reader.

It is suggested that the term "edge preference" (previously one
part of the dual definition of "edge efifect") be used to indicate those
cases where a species of animal shows a preference for the edge be-
tween two or more vegetational associations as opposed to the centers
of these associations. In some cases this preference may be due to
the special vegetation of the edge, as in "B" above ; in other cases
it may be due to the simultaneous access to the two adjacent types,
as in "A" (or "B"). In order to arbitrarily distinguish edge prefer-
ence from juxtaposition in the "A" types of edges, we shall later
on dt.?mt juxtaposition as referring to several (more than two) vege-
tational associations. It should be observed that edge preference is
related principally to species requirements ; that it is only indirectly
related to cruising radius, and not at all related to the nature of the
entire range. It therefore should not be used synonymously with
juxtaposition. Edge preference refers only to the attractiveness of
the edge itself to a certain species of animal. If the edge is very
attractive to a given species, it will be found near that edge, if the
edge is only slightly more attractive than the surrounding vegetation,
the animal will not be found as closely associated with that edge. If,
for example, ruflfed grouse are Hushed more frequently from near
the edge between spruce flat and northern hardwood than from the
center of cither type, then it might be said that rufifed grouse .show
a preference for the spruce flat-northern hardwood edge.

The new term, threshold of edge preference, is a species char-
acteristic which may be defined as being attained when it is possible
to detect the smallest amount of preference for the edge between
two types as opposed to the centers of these types. This concept, there-
fore, is a function of type area and species requirements, but is not
directly dependent upon cruising radius. A method of computing the
threshold of edge preference will be presented below.

The threshold acreage of edge preference has to do with type
areas. It is attained when a given plant community becomes of sufii-

130

Roosevelt IVildllje Bulletin

cient size for a given species of animal to show a preference for its
periphery rather than its center. Thus threshold acreage also is a
function of type area and species requirements, and again, is not di-
rectly dependent upon cruising radius.

The principal distinction between threshold of edge preference
and threshold acreage is that they describe opposite extremes of range
concHtion. Threshold acreage of edge preference occurs in areas with
the greatest possiljle degree of intersix^rsion of different vegetational
associations, i.e., where there are a great many different vegetational
asscjciations of very small size confined in a hmited area. This con-
cept presupposes cover types so small in extent as to preclude a pref-
erence for the ])eriphery of any type as opposed to its center. On
the other hand, threshold of edge preference occurs when the vege-
tation is composed of comparatively large blocks of different plant
associations. If the vegetation of each of certain of these large blocks
is partially deficient in the requirements of the species in question,
it chooses to reside near that edge of the type at which it can fulfill
its requirements from complementary provisions of the edge or of
the adjacent types.

Juxtaposition is defined as the proper distribution of [species']
requirements within the cruising radius of the species in question.
The principal distinction between juxtaposition and edge preference
is tliat tlie former in the case of many species presupposes the neces-
sity of several different plant associations, while the latter has to do
only with individual edges. In the case of an animal with a large
cruising radius, blocks of different types can be more extensive than
in the case of species with a relatively low cruising radius. Proper
ju.xtaposition of required elements for cottontail rabbits may require
much closer spacing tlian the proper juxtaposition of required ele-
ments for ruft'ed grouse. Juxtaposition is largely a function of the
cruising radius of the species involved.

C. Computation of Threshold of Edge Preference. If the

above definition of edge preference is tenable, then it can be derived
in mathematical terms on any given range. The existence of edge
preference can best be determined from random obsen-ations on the
distribution of the species in question. Such random observation can
be accomplished in several ways but one of the most accurate is the
periodic traversing of a grid line of trails such as is found on the
Check Area described above. However, it should be observed that
these trails must furnish representative samples of the cover types
and these samples must be in proportion to the total area of each
type. Examination of Table 37 will reveal that this condition is \tr\

Edge Effect of the Lesser Vegetation

131

well met on the Check Area since "Percentage of Line in Each
Type" very closely approximates "Percentage of Each Type."

TABLE 37. COMPUTATION OF THRESHOLD OF EDGE
PREFERENCE FOR THE MAJOR TYPES* ON THE
CHECK AREA

Forest Types

Feature computed

North-
ern
Hard-
wood

Mixed
Hard-
wood

Spruce
Flat

Lower
Spruce
Slope

Spruce
Swamp

Open
Swamp

A. Number of sections of
line in each type

82

143

102

18

16

5

B. Total length of line in
each type (chains)

1,595.9

1,233.3

831.9

180.0

51.9

36.2

C. Average length of sec-
tions of line in each
type (average width of
type areas, of B/A) . . .

19.5

8.6

8.2

10.0

3.2

7.2

D. Average maximum pos-
sible distance from edge
(type centers to edge,
or C/2)

8.8

4.3

4.1

5.0

1.6

3.6

E. Average median dis-
tance from center of
type to edge (= thresh-
old of edge preference or

cm

4.4

2.2

2.0

2.5

0.8

1.8

* Types occurring in very small blocks are below the threshold acreage of edge
preference for grouse and deer.

Since the threshold of edge preference is defined as a function
of tyi>e areas, the determination of this threshold may be computed
as follows :

A. Count the number of sections of line in each type.

B. Measure the length of line in each type.

C. Compute the average length of sections of line.

D. Then, in any type, the average maximum possible distance
from the edge will be C/2.

E. And, in any type, the average distance halfway to the
center of the type will be C/4. Animal distribution aver-

132

KuoscvcU li'ildlijc Bulletin

agiiif^ closer to the edge than this distance indicates edge
preference. Distribution averaging farther from the edge
than this distance indicates preference for the type. In
other words, the value C/4 may be defined as the threshold
of edge preference, and when a given species of animal
averages a distance less than C/4 from the edge, it may
be said that it exhibits a degree of edge preference.

It should be noted that this method of computing edge prefer-
ence can be applied only in those cases where interspersion of types
is not so extreme as to approach the threshold acreage of edge pref-
erence. Thus, it can be readily applied to the three major forest
types and some of the other types on the Check Area of the Hunt-
ington Forest, but not to the very small types such as the natural
forest openings or to the small areas of post-burn types. Further-
more, since the method depends on randomization, a large number
of samples is required, either a relatively few "censuses"* of a large
population or a large number of "censuses" if the population is small.
Since the population of grouse on the Check Area of the Huntington
Forest is very low, a comparatively large number of "censuses" is
required to indicate the presence or absence of edge preference. For
this reason, the computations and tabulations based on the numl)er
of observations of grouse and deer should be taken more as an ex-
ample of the method rather than as an actual determination, for a
statistically sound determination would require a great many more
observations. However, the method of computing the threshold of
edge preference can be applied to any section of game range since
it is based on size of type areas.

With the computation of the threshold of edge preference com-
pleted, the next step is to find out whether the species in question
exhibit a preference for the edges under consideration. This was
done by plotting 763 observations** of ruiTed grouse and 808 obser\-a-
tions** of deer on a type map of the Check Area and then measuring
the distance of each observed animal from the nearest edge. These
distances were tabulated for each season and the average distance
from each edge computed. The average distances from the edge were
then plotted on a seasonal basis in Figure 18 so as to show their

* For our purposes, a "census" may he defined as the systematic periodic tra-
versin.sr of all of the hne on the Check Area, and tlie recording of the location
of each animal observed.

** These observations were obtained from a long-range study (still in prog-
ress by the Roosevelt Station which involves periodic "censuses" of the Check
Area. The project was initiated in October 1938 but discontinued during the
war and without regularity thereafter.

[133]

134

Roosevelt ll'ildlije Biillelin

relation to the edge (center (jf transition), the threshold of edge
preference (C/4), and the type centers {C/2). Graphs showing the
seasonal distrihution of grouse and deer in the various forest tyix*s
also are presented (Figs. 19 and 20). Although derived from the
same data these latter are only incidental to the co.nputations of edge
preference, but nevertheless they do serve to substantiate some of
the correlations indicated by the computations of edge preference.

D. Wildlife Distribution in Relation to Edges. The detailed

analysis of the vegetation cannot be expected to fully explain the
vagaries of grouse and deer distribution in relation to the various
edges until more s])ecitic and detailed information is available on the
requirements of grouse and deer in relation to the species of plants
ancl the vegetation as a whole. The analysis already has served its
purpose in that it has demonstrated a possible deficiency of the
range for grouse. If lack of fall and early winter fruits had charac-
terized only a few of the edges it would have served as a factor in
comparative edge preference but since all edges were nearly uniform
in this deficiency it is necessary to look for other reasons as the
causes for variation in edge preference. Although the reasons for
the varying types of edge preference may be contained in our detailed
analysis of the vegetation these reasons cannot be traced until more
is known of the detailed requirements of the wildlife involved in
relation to the vegetation. For this reason edge preference in rela-
tion to edge vegetation can be interpreted only in a general wa\-.

In order to determine the extent to which distribution of deer
and grouse was correlated with the forest types and the edges be-
tween them, observations of deer and grouse seen on the Check Area
were plotted on a type map. These observations were made in the
course of periodic "censuses" over a six-year period by the Roosevelt
Station staff. In the course of each of these censuses all of the 54
miles of grid trails on the Check Area were covered in one day
(King, et al. '41).

As was pointed out under "Computation of Threshold of Edge
Preference," these conclusions are based principally on observations
of areas covered by the four major forest types: northern hardwood,
mixed liardwood. spruce flat and spruce slope.* Observations made
in the small areas of post-burn types were so limited in number as
to preclude even tentative conclusions and hence were included under
the "all other" group of types in Figures 19 and 20. Furthermore,

* Unfortunately, the original t\ r-e map did not indicate a sutticiently large
proportion of this type to warrant consideration, hence, its edges were not sam-
pled. Subsequent revision of the type map considerably expanded this type.

10

20

<
en

30
g40

50

SPRING

SUMMER

FALL

NOftTHEPN. HARDWOOD

MIXED HAf?DW0OD

WINTER

10

20
30
140

50
60
70
80

Fig. \9. Grouse distribution in relation to seasons and forest t>pes, based on
763 observations during six \ears on tbe 4,063-acre Check Area. The heavy
liorizontal hues dehniit the proportional areas covered by each of the types ;
the shaded zones s1k)w the proportion f)f the grouse population in each lype.

[135]

Fig. 20. Deer distribution in relation to seasons and forest types, based on
808 observations during six years on tbe 4,056-acre Cbeck Area. The heavy
iiorizontal Hues deHmit the proportional areas covered by each of the types;
tlie shaded zones show tlie proportion of the deer population in each type.

[ 136]

Edge Effect of the Lesser V cgetalioii

137

this latter procedure was justified by the impractical)iHty of deter-
mining a threshold of edge preference due to the small area of these
types which results in an approach of the threshold acreage of edge
preference. Although a threshold of edge preference was determined
for spruce swamp and open swamp types, the paucity of observations
in these types again precluded using these data as a basis for deter-
mining edge preference.

Examination of the vegetative data already has revealed that
none of the edges, nor any of the forest types, appear to completely
fulfill all of the requirements of ruffed grouse to an extent allowing
saturation population characteristic of the best known range. Fur-
thermore, no combination of edges or types, as they now exist, can
be so construed. Of course, the possibility should not be overlooked
that saturation point, although considered a species characteristic,
possibly may be affected to some degree by the quality of the range.
However, if saturation point is to retain any semblance of validity
as a species characteristic, then certainly the low population charac-
teristic of Adirondack range (about one bird to twenty acres) is well
below what might be considered a "minimum" saturation point pos-
sible for the species.

The general deficiency of fruit on all parts of the range as com-
pared to ideal grouse range indicates the conclusion that it may be of
considerable importance in limiting the population. Inasmuch as the
habitat as a whole contains such low potentialities for grouse (as in-
dicated by the population), it would seem logical to expect the birds
to occur most frec[uently near the edges between different plant asso-
ciations, i.e., show some degree of edge preference, since presumably
the wider variety of food at least, if not cover, would better fulfill
their needs. This, however, was not proven to be the case at all
edges.

The distribution of grouse and deer in relation to the edges
between the four major types was plotted in Figure 18. Reference
to this figure indicates at a glance that in spite of their increased
diversity, the edges were not uniformly preferred over type interiors.
In several cases the type centers actually were preferred to the edges.
In other cases distribution at one side of an edge showed edge pref-
erence while distribution on the other side showed edge repulsion, or
type preference. Edge and type preference in other cases alternated
from one season to the next. Distribution in some cases appears to
be consistently correlated with the vegetation, in other cases the
principal consistency was the lack of consistency. Thus the writer
is forced to conclude that wildlife is not always a product of edges,
at least not in the interior of large unbroken stands of timber. In-

138

Roosevelt JrUdlife Bulletin

creased variety is not always synonymous with increased wildlife
value, or at least with increased wildlife preference. Thus the refusal
of deer and grouse to conform to man-made cliches on distribution
becomes somewhat impressive.

Edge preference was indicated by grouse only in the mixed
hardwood type and in the spruce slope type. On the other hand,
grouse flushed in the spruce flat and northern hardwood types aver-
age closer to the center of the type than the threshold of edge pref-
erence. Edge and type preference by deer very closely resembled
that of grouse, although seasonal variation was greater.

.\ consistently strong edge preference was shown by both grouse
and deer in the mixed hardwood type. Grouse appeared to be more
strongly attracted by this edge at all seasons of the year than were
deer. The fact that all three edges of this type (both toward the
northern hardwood type and the two spruce types) attracted grouse
and deer possibly may be ascribed to type repulsion, rather than edge
preference. This possibility with respect to grouse is further sug-
gested in Figure 19. From this figure it is apparent that the mixed
hardwood type was characterized by a subnormal proportion of
grouse throughout most of the year. Deer also showed a subnormal
distribution in this type for all seasons except fall (Fig. 20). It
therefore would appear that our earHer hypothesis to the effect that
northern hardwood and spruce flat types better fulfill the require-
ments of these species than does the mixed hardwood type is borne
out. These conditions point to the logical conclusion that if any por-
tion of the range of a species is in some respect undesirable, then
the species would be expected to occur near the edge of that type
rather than in tlie center of it. From this it appears that edge pre-
erence works not only as a positive attracting force but also as tlK
result of a negative repelling force.

The spruce slope type also caused a strong edge preference on
the part of both grouse and deer. This type borders in all cases on
the mixed hardwood type. The degree of preference for this edge
by both deer and grouse was greatest in the summer and fall. Winter
and early spring distribution, however, was closer to the center of
the spruce, as though in response to more eflfective conifer cover and
easier travel in the more compact snows found under this type of
cover. It might well be that edge preference in this type too was a
result of type repulsion, for this type is comparatively low in both
browse for deer and buds and fruits for grouse. Since the edge pro-
vides a dense cover adjacent to the comparatively more productive
mi.xed hardwood, the combination logically would be expected to
more eft'ectively meet the requirements of these species.

Edge Effect of the Lesser Vegetation 139

DistriI)ution of grouse and deer in the northern hardwood and
spruce flat types was such as to show great seasonal variation in edge
preference. On the whole, both species showed a greater preference
for the centers of these types than for the edges. The seasonal fluc-
tuation in degree of edge preference was greater in the case of deer,
probably as a reflection of greater mobility.

Grouse flushed in northern hardwood types in the spring aver-
aged nearer the center of the type than toward either edge (mixed
hardwood or spruce flat). This possibly may have ])een the result of
attraction for the large amounts of early spring greens and bright
sunshine. With the advent of summer, however, the birds tended to
be closer to the coniferous or partially coniferous types. This ten
dency continued into the fall. Contrary to what might be expected,
however, winter birds flushed in the northern hardwood type oc
curred on the average far in the interior of the type rather than near
the edge toward the shelter of the conifers. The high preference of
grouse for the northern hardwood type is further demonstrated in
Figure 19, which shows above-normal numbers of grouse in this type
at all seasons except fall.

Deer showed a more marked preference for the edge of north-
ern hardwood where it bordered on spruce flat than where it bor-
dered on mixed hardwood. We might logically expect this inasmuch
as the mixed hardwood type does not as well complement the north-
ern hardwood as does the spruce flat. Deer found in the hardwood
type showed a preference for the edge toward mixed hardwood only
in the winter. At other seasons they ranged far into the middle of
the northern hardwood type. The variable preference of deer for th?
edge of the northern hardwood type is reflected in the varying pro
portion of animals seen in this type at different seasons (Fig. 20).
The only season at which there was a superabundance in the type
was in the spring, possibly a reflection of more early greens and
bright sunshine, as in the case of grouse. In the summer and fall
the proportion of deer in this type fell off considerably and to a much
greater extent in the winter.

Both deer and grouse showed a greater preference for the center
of the spruce flat type than for its edges towards either mixed hard-
wood or northern hardwood. This type was the most consistently
preferred at all seasons. At the edge toward mixed hardwood, grouse
approached the threshold of edge preference only in the winter, pos-
sibly in search of buds. However, the birds did not venture as far from
the center of the spruce at edges toward the northern hardwood types.
Deer in this type (spruce flat) showed a similar preference for the
center of the type as opposed to the edge, but their average distance

140

Roosevelt IVildlije Bulletin

from the edge varied considerably from season to season. The gen-
eral reaction of the deer was similar in both the northern hardwood
and mixed hardwood edges of the spruce flat type : edge preference
in the fall, and then back to the centers of the spruce flat in the
winter. The edge tendency in the spring possibly may be correlated
with the warmer conditions, earlier snow removal, and fresh greens.
The apparent summer recession to the spruce to a large extent may be
due to the topographical location of this tyi^e in relation to lakeshores
and streams. Deer feed extensively on aquatic plants in the summer
and use the adjacent spruce flat types for cover. The fall tendency
toward hardwood may be partially ascribed to a search for mast and
the winter preference for the center of the spruce types is correlated
with better cover and easier travel. The high type preference of
deer for the spruce type at all seasons except fall is illustrated in
Figure 20.

VII. SUMMARY

The most striking over-all feature of Adirondack forest vegeta-
tion, as revealed on the Huntington Forest, is the small variety of
dominant species. Most of the tree cover is composed of spruce,
balsam, yellow birch, red maple, beech, and sugar maple. The great
bulk of shrub cover is composed of witch hobble. The major portion
of the herbaceous cover is composed of wood fern, wood sorrel, and
dwarf dogwood. Differences in forest types result from varying
compositions of these species plus additions of small amounts of
other species.

The edges between Adirondack forest types belong to the group
of edges composed only of the constituents of the two adjacent types.
They are characterized by a special flora only to the extent that in
certain edges some species are more concentrated at the center of
transition than toward the center of either type.

The transition from one forest type to another usually is re-
flected in changes of species composition of the lesser vegetation as
well as in the relative amounts of individual species.

The variety of species across any transition is greater than that
at any part of the transition.

Many of those plant species occurring at all parts of the edge
occur in significantly greater amounts on one of the three parts of
the transition.

Although the edges sampled were restricted to the Huntington
Forest, it is doubtful that the variety of species would have been
greatly augmented by extending the study into other parts of the
Adirondacks. Eighteen species of trees, 21 species of .shrubs, and

Edge Effect of the Lesser Vegetation

141

49 species of herbs were recorded. (Some "herbs," as grasses and
sedges, included many undifferentiated species.)

In general, the total amount of lesser vegetation did not vary as
much as the relative amounts of the three types, that is, where there
was a great amount of shrub cover, there was relatively little herb
cover or arborescent reproduction, and where there was a large
amount of herb cover, there was relatively little shrub cover or
arborescent reproduction, or if there was a great amount of arbor-
escent reproduction, there was relatively little shrub or herb cover.

The three components of the lesser vegetation rank in general
cover value (in the summer) as follows: (1) herbs, (2) shrubs, (3)
arborescent reproduction. At other seasons the ranking is : ( 1 )
shrubs, (2) arborescent reproduction, (3) herbs.

The average width of all edges studied was sufficiently narrow
to be easily traversed by animals of low cruising radius, such as
ruffed grouse.

Even with the increased amount and variety of food at the edges
between various forest types, most of the edges sampled nevertheless
lacked a sufficient variety and abundance of late fall and winter
fruits for ruffed grouse.

Grouse and deer distribution data indicate that the centers of
some types are more highly preferred than their edges.

Edge and type preference, as derived from these data, varies
from season to season at each of the edges studied.

The edges most nearly fulfilling the requirements of ruffed
grouse were those between the three major climax forest types;
spruce flat, mixed hardwood, and northern hardwood. Even these
were lacking in substantial amounts of late fall and winter fruits.
Deficiencies at other edges varied from season to season.

The edges showing the greatest apparent year-round attraction
for grouse were those of mixed hardwood, both toward spruce flat
and toward northern hardwood. This appeared to be more strongly
related to a type repulsion than edge preference, for distribution in
the mixed hardwood was very scant.

In general, the distribution data showed a marked preference by
both deer and grouse for the northern hardwood and spruce flat types.

A detailed analysis is presented of the lesser vegetation at the
edges between various Adirondack forest types. The method of
analysis also is presented in the hope that it may prove of value to
others contemplating this type of investigation. The distribution and
abundance of herbs, shrubs, and arborescent reproduction is analyzed
both graphically and mathematically and the major features of value
to deer and grouse at each edge are discussed. Data on distribution

142

Roosevelt Wildlife Bulletin

of deer and grf)use in relation to edges and forest types also are pre-
sented and discussed. Concepts relative to edge effect are brought
together and reviewed, and one new concept (the threshold of edge
effect ) is developed. A list of plant si^ecies encountered is presented.

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Edge Effect of the Lesser Vegetation 143

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144

Roosevelt Wildlife Bulletin

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Edge Effect of the Lesser Vegetation

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LIST OF

1. Ash, black

2. Ash, white

3. Ash, mountain

4. Aspen, trembling

5. Balsam Fir

6. Basswood

7. Beech

8. Birch, yellow

9. Cedar, white, or Arbor vitae

10. Cherry, black

11. Hemlock

12. Hop hornbeam

13. Maple, mountain

14. Maple, red

15. Maple, striped

16. Maple, sugar

17. Shadbush

18. Spruce, red

1. Alder, speckled

2. Blueberry, sourtop

3. Dogwood, alternate-leaved

4. Dogwood, red osier

5. Elderberry, common

6. Elderberry, red-berried

7. Hazelnut, beaked

8. Honeysuckle, fly

9. Labrador tea

10. Leather leaf

11. Meadow sweet

12. Mountain holly

13. Raspberry, dwarf

14. Raspberry, red

15. Raspberry, red

16. Sweet gale

17. Sweet gale

18. Virgin's bower

19. Wild raisin

20. Willow

21. Witch hobble

PLANT SPECIES

Trees

Fraxinus nigra Marsh.

Praximis amcricana L.

Sorbiis amcricana Marsh.

Fopuliis trcnmloidcs Michx.

Abies balsamea (L.) Mill.

Tilia amcricana L.

Fagns grandifolia Ehrh.

Bchda lutca Michx. f.

Tliuja occidcntalis L.

Primus scrotina Ehrh.

Tsuga canadensis (L.) Carr.

Ostrya virginiana (Mill.) K. KocIl

Acer spicatum Lam.

Acer rubrtim L.

Acer pensylvanicum L.

Acer saccharum Marsh.

Aniclanchicr laevis Wieg.

Picea rubra (Du Roi) Dietr.

Shkubs

Alntis incana (L.) Moench.
I'accinium canadense Kalm.
Cornus alternifolia L. F.
Cornus stolonifera Mich.
Santbucus canadensis L.
Sandmcus racemosa L.
Corylus cormita Marsh.
Lonicera canadensis Marsh.
Ledum groenlandicum Oeder.
Chaniaedaphne calyculata (L.) Moench.
Spira-ca latifolia (Ait.) Borkh.
Ncmopantlms mucronata (L.) Trel.
Riibus acaulis Michx.
Rubtis idaeus L. Var. canadensis Rich.
Ridms idaeus L. Var. strigosiis Michx.
Myrica gale L.

Myrica gale L. Var. subgJabra

(Chev.) Fern.
Clematis virginiana L.
J^ihurnum cassinoides L.
Salix sp.

Viburnum alnifolium Marsh.

146

Roosevelt Wildlife Bulletin

1. Aster

2. Baneberry, red

3. Baneberry, white

4. Bedstraw

5. Clintonia

6. Club-moss, common

7. Club-moss, Sliiiiy

8. Cucumber- root

9. Dogwood, dwarf

10. False lily-of-tlie-valley

11. Fern, beech

12. Fern, bracken

13. Fern, hay-scented

14. Fern, New York

15. Fern, oak

16. Fern, royal

17. Fern, sensitive

18. Fern, spiny-toothed shield

19. F'ern, wood

20. Goldenrod, hairy

21. Goldenrod, woods

22. Goldthread

23. Grass, various species

24. Hieracium

25. Jack-in-the-pulpit

26. Jewelweed

27. Miterwort, false

28. Nettle, wood

29. Nightsliade, enchanter's

30. Partridge berry

31. Pirola

32. Sarsaparilla

33. Sedges, various species

34. Snowherry, creeping

35. Solomon's seal

36. Sphagnum moss

37. Spring beautv

38. Starflower

39. St. John's-wort

40. Strawl>erry

41. Thistle, Canada

42. Trillium, painted

43. Trillium, red

44. Twin-flower

45. Twisted stalk

46. Violet, dogtooth

47. Violet

48. Violet, fal.se

49. W'intergreen

50. Wood sorrel

Herbs

Aster spp.

Actca nihra (Ait.) Willd.
Actea alba (L.) Mill.
Galiuin spp.

Clintonia borcalis (Ait.) Raf.
Lycopodium chivatwn L.
Lycopiidmrn luddulum Michx.
Mcdcolo viri/iniana L.
C ornus canadensis L.
Maianthcmum camidcnsc Desf.
Fhegopteris polypodioides Fee.
Fteridiuin latmsculum (Desv. ) Hieron.

ex R. E. F"ries.
Dennstaedtia punctilobula (Miclix.)

Moore.

Dryoptcris novehoracensis (L.) Gray.
Phe<iopteris dryopteris (L.) F'ee.
Osmunda recjalis L. Var. spectabilis

(Willd.) Gray.
Onoclea sensibilis L.
Dryoptcris spinulosa (O. F. Miill.)

Watt.

Dryoptcris intermedia (Muhl.) Gray.

Solidago rugosa Mill.

Solidago caesia L.

Coptis trifolia (L.) Salisb.

// icracium spp.

Arisaema triphyllum (L.) Schott.
Iinpaticns bi flora W alt.
Tiarella cordi folia L.
I.aportea canadensis (L. ) Gaud.
Circaca alpina L.
Mitcliclla re pens L.
Pirola sccunda L.
Aralia nudicanlis L.

Chioi/enes hispidula (L.) T. & G.
Polygonutum pubescens Pursh.
S phagnum sp.

Claytonia caroliniana Michx.

Trien talis boreal is Raf.

Hypericum sp.

Fragaria virginiana Duch.

Cirsium arvcnse (L. ) Scop.

Trillium undulatum Willd.

Trillium erectum L.

I.innaca borcalis L. var. a)nertcafia

(Forbes) Rehder.
Streptopus roseus Michx.
Erythronium americanum Ker.
Viola spp.

Dalibarda repens L.
Gaultheria procumbcns L.
Oxalis montana Raf.