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A STUDY OF RUFFEB GRGUSE DBUMMENG SITES
IN NORTHERN MICHIGAN

 

Thesis for the Degree of M. S.
MICHIGAN STATE UNEVERSI'I‘Y
WALTER LAWRENCE PALMER

1961

IIIIIIIIIIIIIIIIIIIIIIIIIII

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L 13 R A R Y
Michigan State
University

 

 

 

 

 

 

 

 

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A STUDY OF RUFFED GROUSE DRUMMING SITES

IN NORTHERN MICHIGAN

BY

Walter Lawrence Palmer

AN ABSTRACT

Submitted to the School of Graduate Studies of Michigan
State university of Agriculture and Applied Science
in partial fulfillment of the requirements

for the degree of

MASTER OF SCIENCE

DEPARTMENT OF FISHERIES AND WILDLIFE

1961

ABSTRACT

Forty ruffed grouse (Bonasa umbellus L.) spring courtship sites

 

were studied in 1959. Sites consisted of drumming logs and surround-
ing vegetation, and had been located on the Rifle River Area, Ogemaw
County, Michigan, during a nine-year period from 1951 to 1959.

I recorded log dimensions, azimuth of log axis, species of log
whenever possible, and distances from the drumming position to the
nearest end of the log. Vegetation surrounding the logs was studied
quantitatively and qualitatively by utilizing a series of quadrats and
transects.

Logs averaged about 12 inches high, 13 inches wide, and almost 20
feet long. Logs used most often were larger in diameter than those
used infrequently, but length did not appear to be important. Almost
three-fourths of the logs were white pine (Pinus strobus) and about half
of the remaining logs were other coniferous species. The drumming loca-
tion was usually about five feet from the larger end of the log.

Vegetation except in the 0-2 feet tall size class, was usually
more dense immediately surrounding the logs than elsewhere in the habitat.
In the 0-2 feet tall size class, fewer stems were tallied near the logs
than elsewhere.

No significant vegetational differences were found between logs
used several years as compared with logs used less frequently.

All forty drumming logs were located in lowland vegetation types,
but usually were near a lowland-upland type boundary. Although 44 woody
plant species were tallied on the sites, speckled alder (Alnus rugosa),

red-osier dogwood (Cornus stolonifera), willows (Salix $22.), balsam fir

ii

(Abies balsamea), and mountain holly (Nemopanthus mucronata) were most

 

 

common.

The Rifle River Area had not been lumbered for several years, and
had an excessive deer (Odocoileus virginianus) population. Much of
the area contained merchantable or near-merchantable stands of large-

toothed aspen (Populus grandidentata) and trembling aspen (g, tremuloides)

 

 

which could be harvested. Results of vegetation analyses suggested that
plants which are not palatable to deer are the only species reproducing
successfully, and therefore these supply the undergrowth necessary for
the establishment of drumming sites.

Although no quantitative grouse population data are available for
comparison, it is possible that the combined effect of inadequate timber
harvest, and the excessive deer population have reduced the grouse
carrying capacity of this area by reducing the percentage of the area

suitable for grouse territories.

iii

ACKNOWLEDGEMENTS

I thank Mr. H. D. Ruhl, Chief, Game Division, Department of
Conservation, for initiating and urging completion of the long-term
population study. The research was conducted under Pittman-Robertson
Projects W-46-R and W-95-R.

Dr. R. A. MacMullan and Ralph Blouch directly supervised my work
while they were biologists-in-charge of the Houghton Lake Wildlife
Experiment Station. Dr. G. A. Ammann offered suggestions on field
techniques from time to time and he and Dr. C. T. Black kindly edited
the manuscript. Dr. L. L. Eberhardt and R. G. Heath assisted with
statistical methods. Dr. John George, Patuxent Wildlife Research Center,
U. S. Fish and Wildlife Service, helped design the vegetation sampling
plan.

Jeanne Wheeler, Dorothy watling and my wife Patt typed and re-
typed the several versions of the manuscript.

I wish to thank Dr. G. A. Petrides, Professor of Wildlife Manage-
ment, Michigan State University, who constructively criticized and
edited the manuscript and finally to Dr. Peter I. Tack, Head, Department
of Fisheries and Wildlife, Michigan State University who, despite an
already burdensome schedule, urged completion of the report and assisted

in editing the manuscript.

iv

Ii

TABLE OF CONTENTS

Introduction 0 o o o o o o o o o o o o o o o o o o o

The SCUdy Area 0 o o o o o o o o o o o o o o o o o o

Locating and Recording Drumming Log Use

Dimensions, Types and Directions of Drumming Logs .

Analyses of Vegetation . . . . . . . .
Plant Density Analysis . . . . . . . .
Comparison of Plant Densities Near and
Plant Composition . . . . . . . . . .
Discussion . . . . . . . . . . . . . .
Summary . . . . . . . . . . . . . . .
Literature Cited . . . . . . . . . . .

Appendix 0 O O O O O O O O O O O O O 0

Far

Page
. l

. 3
. 4
. 4
.ll
.15
.17
.28
.34
.36
.38

.39

Table

1.

2.

3.

4.

5.

7.

8.

LIST OF TABLES
Page

Cover type, and percentages of the 43l8-acre Rifle
River Area, Michigan . . . . . . . . . . . . . . . . 3
The relationship between the number of years 40 ruffed
grouse drumming logs were used and observed by arbitrary
diameter classes, and the computed probability of use in
any one year by size classes . . . . . . . . . . . . 8
The relationship between the number of years 40 ruffed
grouse drumming logs were used and observed by arbitrary
length classes, and the computed probability of use in
any one year by size classes . . . . . . . . . . . . 9
Number of woody stems tallied by size classes on nine
one-quarter milacre quadrats near 40 ruffed grouse
drumming logs, Rifle River Area, Michigan . . . . . 16
Number of woody stems tallied on "near" and "far" halves
of one-hundreth acre.transects near 40 ruffed grouse
drumming logs, Rifle River Area, Michigan . . . . . 16
Converted average number of woody stems per acre by
size classes encountered on quadrats and transects
located near 40 ruffed grouse drumming logs, Rifle
River Area, Michigan . . . . . . . . . . . . . . . . 17
Comparison of woody plant density in the 0-2 foot tall size
class on matched pairs of quadrats near 40 ruffed grouse
drumming logs, Rifle River Area, Michigan . . . . . . 18
Comparison of woody plant density in the 2-5 foot size
class on matched pairs of quadrats near 40 ruffed grouse

drumming logs, Rifle River Area, Michigan . . . . . . 19

vi

Table

10.

ll.

12.

13.

14.

15.

16.

Comparison of woody plant density in the 5-8 foot size
class on matched pairs of quadrats near 40 ruffed
grouse drumming logs, Rifle River Area, Michigan . . .
Comparison of woody plant density in the 8'-3" dbh
size class on matched pairs of quadrats near 40 ruffed
grouse drumming logs, Rifle River Area, Michigan . . .

Comparison of woody plant density in the 3-9" dbh size

Page

20

21

class on matched pairs of quadrats near 40 ruffed grouse

drumming logs, Rifle River Area, Michigan . . . . . . .
Comparison of woody plant density in the 5-8 foot class
on matched pairs of transects near 40 ruffed grouse
drumming logs, Rifle River Area, Michigan . . . . . . .
Comparison of woody plant density in the 8'-3" dbh
class on matched pairs of transects near 40 ruffed
grouse drumming logs, Rifle River Area, Michigan . . .
Comparison of woody plant density in the 3-9" dbh class
on matched pairs of transects near 40 ruffed grouse
drumming logs, Rifle River Area, Michigan . . . . . . .
Values of t obtained by analyses of stem densities on
quadrats 1-4 compared to quadrats 6-9 and "near" halves
of transects compared to "far" halves surrounding 40
ruffed grouse drumming logs by arbitrary size classes
using the matched pairs technique . . . . . . . . . . .
Numbers of stems of woody deer food plants and non-
deer food plants tallied by size classes on 360 Quadrat
near 40 ruffed grouse drumming logs, Rifle River Area,

MiChigan o o o o o o o o o o o o o o o o o o o o o o o

22

23

24

25

26

S

27

Table

17.

18.

19.

20.

21.

22.

23.

Page
Number of quadrats in which at least one stem of the
37 plant species occurred and the computed relative
frequency of each species by size classes . . . . . . , 29
Total stems and relative density of woody plant
species occurring on 360 quadrats near 40 ruffed grouse
drumming sites . . . . . . . . . . . . . . . . . . . . . 30
Number of drumming logs occupied by male ruffed grouse
during 8 calendar years, Rifle River Area, Michigan . . 31
Analysis of variance of density of woody vegetation in
the 0-2 foot height class on quadrats 1-4 by numbers of
years 40 ruffed grouse drumming logs were occupied,
Rifle River Area, Michigan . . . . . . . . . . . . . . . 32
Analysis of variance of density of woody vegetation in the
2-8 foot height class on quadrats 1-4 by numbers of
years 40 ruffed grouse drumming logs were occupied, Rifle
River Area, Michigan . . . . . . . . . . . . . . . .. . 32
Analysis of variance of density of woody vegetation in
the 8'-3" height class on quadrats 1-4 by numbers of years
40 ruffed grouse drumming logs were occupied, Rifle
River Area, Michigan . . . . . . . . . . . . . . . . . . 33
Analysis of variance of density of woody vegetation in
the 3L9" dbh height class on quadrats 1-4 by numbers of
years 40 ruffed grouse drumming logs were occupied,

Rifle River Area, Michigan . . . . . . . . . . . . . . . 33

LIST OF FIGURES

Figures Page

3.

1.

2.

4.

5.

7.

Record of use of 40 ruffed grouse drumming logs,
1951-1959, Rifle River Area, Michigan . . . . . . . S
Plotted frequency distribution of width, height, and
length of 40 ruffed grouse drumming logs on the

Rifle River Area, Michigan . . . . . . . . . . . . 7
Sketch of quadrat vegetation sampling design used

"near" the drumming log. Each quadrat is one-quarter
milacre in size. Log is hypothetical . . . . . . . 13
Sketch of transect vegetation sampling design with

"far" quadrats. Each transect is one-hundreth acre.

in size. Quadrats 6, 7, 8, and 9 are counterparts

of quadrats l, 2, 3, and 4 . . . . . . . . . . . . 14
The upland aspen type with scanty understory occurs
abundantly in northern lower Michigan . . . . . . . 35
A jack pine forest near Luzerne, Oscoda County,

Michigan . . . . . . . . . . . . . . . . . . . . . 35

The "edge" between upland and lowland aspen types . 35

vii

A STUDY OF RUFFED GROUSE DRUMMING SITES

IN NORTHERN MICHIGAN

BY

Walter Lawrence Palmer

A THESIS

Submitted to the School of Graduate Studies of Michigan
State university of Agriculture and Applied Science
in partial fulfillment of the requirements

for the degree of

MASTER OF SCIENCE

DEPARTMENT OF FISHERIES AND WILDLIFE

1961

Approved

 

INTRODUCTION

The ruffed grouse (Bonasa umbellus L.) is an important game
bird in Michigan, second only to the ring-necked pheasant (Phasianus
colchicus L.) in average annual kill. The kill exceeds a half-million
birds during peaks of abundance, and recently has totalled approximately
a third of a million birds (Ryel and Eberhardt, 1960). Improved highways,
faster cars, shorter work weeks, and competition for hunting areas in
southern Michigan farmland, have been responsible for an increasing
interest in northern Michigan grouse hunting.

Male grouse usually select individual territories during their
first fall and winter, and keep them throughout life (Eng, 1959). Once
these territories are established, males become quite sedentary and
probably spend all their time within an area less than a quarter-mile
in radius (Palmer, 1956; Eng, 1959). Some slightly elevated object within
the territory is selected on which they perform the courtship drumming
act. The elevated object is usually a log, but other objects such as
boulders or hummocks may be used (Edminster, 1957; Frank, 1947; Bump,
Darrow, Edminster and Crissey, 1947). The drumming sound is produced by
moving the wings forcefully downward and forward.

Previous Investigations

 

Edminster (1947) described typical cover used for drumming in New
York State. He found that birds preferred young, second growth woodland,
predominantly hardwood, but where a fair scattering of conifers, especially
young conifers, was present. He also found that mature woodlands, solid

coniferous stands, Open slaihings, and brushland seem to be avoided.

I} lull, I (A

2

Dorney (1959) in Wisconsin noted that males did not select
territories in habitat devoid of forest undergrowth less than 6 feet tall.

Eng (1959) attempted the first detailed analysis of vegetation near
drumming logs, but did not compare this vegetation with that elsewhere
in the habitat.

Since cover, soils, and other ecological factors in northern Michigan
differ considerably from the areas in which Eng's Minnesota study took

place, his data are not readily applicable here.

Present Conditions in Northern Lower Michigan

Virgin forests in northern lower Michigan were cut during the middle
and late nineteenth century. The logging was followed by extensive fires,
which were not adequately controlled until about 1920. Much of the
present-day forest cover dates from this period. As a result, large
areas exist, which have pole-sized or larger trees, where undergrowth
is scanty. This situation is especially apparent on poorer soils. In
many areas excessive numbers of browsing deer (Odocoileus virginianus)
have also helped to reduce plant reproduction. Because of these two
factors, it seemed possible that the availability of suitable drumming
cover is declining in this region. This in turn may be a factor affecting
the abundance of ruffed grouse in Michigan.

Since males are the most sedentary segment of the population,
(Palmer, 1956; Eng, 1959) I thought that a study designed to evaluate
drumming sites would provide valuable information necessary for manage-
ment of areas for ruffed grouse production. Although grouse are polygamous,

and it is possible that cover requirements of cocks and hens differ,

 

 

I‘ 11'

3
especially during the nesting season, generally the two sexes require

similar habitat types.

THE STUDY AREA
During a nine year grouse population study from 1951 to 1959, forty
drumming logs were located and studied on the Rifle River Area, Ogemaw
County, Michigan. This 4318-acre area is largely forest, and several

distinct forest cover types are present. (Table I)

TABLE 1
Cover types, and percentage of the 4318-acre

Rifle River Area, Michigan

 

Typg_ Percentage of total area
Upland Aspen 40.6
Northern swamp hardwoods 15.7
Upland grasses 12.3
Lowland willows and alder 9.4
Lakes 7.9
Northern swamp conifers 7.3
Lowland grass or sedges 4.5
Upland oak 2.1
Upland conifers .2

 

100.0

LOCATING AND RECORDING DRUMMING LOG USE

I located drumming males each spring using the stalking method
(Frank, 1947). The method involved walking slowly through an area
listening for drumming. When I heard a grouse drum, I approached it
carefully. The bird usually flushed at close range, and a search of
the area disclosed the drumming log, which could be identified by the
presence of several fecal droppings. Some males however, did not flush,
but instead hopped down from the log and walked away, often unnoticed
thus making locating of the log much more difficult.

By live-trapping and banding birds, and also by checking areas
repeatedly it was apparent that some birds used more than one log. It
appeared that there was a direct correlation between the birds' tendency
to walk or run away upon being approached and the number of logs on
which it drummed. In other words, grouse which performed on only one
log apparently had a strong attachment for that log and an unwillingness
to leave, and I postulated that this attachment was probably a reflection
of the site's quality. The logs studied to my knowledge, were used by
only one male each spring.

Once drumming logs were located, I attempted to check them in
subsequent years to ascertain whether they were being used. Thus, I had
a continuous record of use for most of the 40 logs for up to 8 years time

(Figure 1).

DIMENSIONS, TYPES AND DIRECTIONS OF DRUMMING LOGS

I revisited each of the forty drumming logs during the summer of 1959

and secured the following information:

PC

1
.f

fx

5
Figure 1. Record of use of 40 ruffed grouse drumming logs,
1951-1959, Rifle River Area, Michigan.

Solid lines represent years logs were definitely used, the left end
being the year each was located. Dashed lines indicate years logs were
not checked for use, and years with no lines indicate logs were not used.

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1951

1952

1953

1954

1955

1956

1957
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1. Log height and width to the nearest one-half inch.

2. Length to the nearest half foot.

3. Species of log or, if very decayed, whether it was a conifer

or hardwood.

4. Azimuth of log axis.

5. Distance of drumming position to nearest end of the log.

Log width varied from 8 to 21 inches with a mean of 13 inches.
Height varied from 7 to 21 inches with a mean of 11.9 inches and length
ranged from 5.5 to 44 feet and average 19.8 feet (Figure 2).

Logs were then tabulated according to the number of years each had
been used and observed, and a ratio was computed between the sum of the
number of years used by the sum of the number of years observed. The
resultant ratio was therefore an estimated probability of use in each
size class for any given year (Tables 2 and 3).

Using this method of analysis, I assumed that if log diameter was
one factor affecting selection the estimated probability of use for each
size class would be lowest in the smaller size classes and approach unity
in the larger log sizes. It appears that this was indeed true, because
the probability of use in the largest diameter class was considerably
larger than in the smallest class.

Thus if logs are at least 7 inches in diameter the probability of
use in any year increases directly with a further increase in log 8123.

Log length showed little correlation with frequency of use, although
I expected that longer logs would tend to be larger in diameter and

therefore should be correlated with use. Although the two larger length

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TABLE 2

The relationship between the number of years 40
ruffed grouse drumming logs were used and observed
by arbitrary diameter classes, and the computed
probability of use in any one year by size classes.

LOG DIAMETER CLASS

 

 

 

 

 

 

 

 

 

 

 

 

No. 7-9.5" Class 10-12.5' Class 13-15.5" Class l6“+ Class
of Years Years Years Years Years Years Years Years
ng_ used observed used ,observed used observed used observed
1 2 5 4 4 7 7 8 9
2 2 7 1 5 4 5 5 9
3 3 6 l 5 4 4 3 3
4 1 4 l 4 4 4 4 4
5 2 2 3 3 4 9
6 3 6 5 5 5 9
7 4 4 1 4
8 4 4 4 8
9 2 3 4 4
10 l 3 5 7
ll 4 4 1 4
12 5 5
13 4 9
l4 8 9
15 7 7
l6 4 5
17 l 4
18 l 3
l9 2 6
Totals 13 3O 62 92 43 65 20 25
Years used 3.433 Years used Years used Years used
Years observed Years observed Years observed Years observed

TABLE 3

The relationship between the number of years 40
ruffed grouse drumming logs were used and observed

by arbitrary length classes, and the computed
probability of use in any one year by size classes.

LOG LENGTH CLASSES

 

 

 

 

 

 

 

 

 

 

 

 

 

No. _5 - 15 feet 15.5-25 feet 25.5-35 feet 35.5 feet +
of Years Years Years Years Years Years Years Years
Logs, used observed used observed used observed used observed
1 l 4 1 3 4 4 3 6
2 2 6 4 5 8 9 4 9
3 1 4 7 7 5 5 8 9
4 5 7 3 6 2 3 3 3
5 2 2 2 7 4 4
6 l 4 4 8 2 5
7 4 4 4 4 5 5
8 1 4 l 5 4 4
9 3 3 l 5 7 7
10 4 4 4 4
11 1 3
l2 5 9
13 5 9
l4 4 9
15 4 4
l6 1 4
l7 4 5
Totals 48 85 27 50 45 50 18 27
Years used Years Used Years used Years used
Years observed Years observed Years observed Years observed

=.565 I".540 =.900 8.667

10

classes were more apt to be used than the smaller classes, the difference
was not consistent. Since none of the 40 logs studied was less than 5.5
feet this appears to be a minimum length requirement, but additional length

apparently is not critical.

Species and types of logs used,gand drumming location selected on logs
Thirty-four of the 40 logs were old, decayed conifers of which at

least 26 were white pine (Pinus strobus). It was sometimeSimpossible to

 

identify species. In most cases logs were either fallen trunks or had been
saw-logs left behind when the virgin stands of timber were cut many years
ago. Freshly fallen, wind-thrown trees, mostly large-toothed aspen

(Populus grandidentata) were sometimes used, and birds invariably drummed

 

near the butt end of these logs, apparently to take advantage of the added
concealment of the exposed roots. On most logs in fact, birds seldom
drummed from near the center or mid-point, but seemed to prefer a position

about 5 feet from the large end.

Logidirections

I recorded azimuth readings to determine whether grouse preferred
logs lying in a particular direction. Twenty-eight or 70 per cent of the
logs lay in a southwest to northeast direction i.e. between 00-900 and 180°-
2700. I did not take a random sample of directions of non-drumming logs
for reasons to be explained in the following section of this report, but I
would expect to find most logs lying in this general direction due to the
effects of the prevailing winds. This is strictly speculation of course,
and therefore the proposition of grouse selecting logs on the basis of

direction cannot be settled at present.

ll

ANALYSES OF VEGETATION

Early in the study I gave some thought to comparing vegetation near
drumming and non-drumming logs. As I have previously mentioned however,
the distinction between these two classes of logs is not always clear-cut.
For example, most cocks select only one log on which to drum and I
speculated that these are prime sites. The 40 logs I studied were of
this type. 0n the other hand, some birds skip from one log to another,
and may use four or five logs. These logs must be classed as drumming
logs. Finally, logs with no evidence of use would be classed as non-
drumming logs, but bias could be introduced if such logs had been used
in the past and were not detected.

The ultimate objective of this study was to ascertain the characteristics
of prime drumming sites so that habitat improvement practices could be directed
toward perpetuation or creation of favorable habitat. Because of the
difficulties and magnitude of a study involving non-drumming as well as
drumming logs outlined above, and also because of the desirability of
learning the requisites of the best sites, I decided to study the
characteristics of the best sites only.

I experimented with several sampling methods which would enable me
to ascertain these characteristics, and at the same time be able to make
comparisons of site characteristics with vegetation adjacently or in the
vicinity of sites. The sampling design had to meet these additional
requirements: (1) vegetation had to be sampled adequately so that the data
could withstand statistical treatment and, (2) the design had to be easily

used in the field.

I'-

12

Eng (1959) used two methods to measure vegetation, the "closest
tree method" described by Cottam and Curtis (1956) to study tree density
and composition, and for smaller vegetation he used four milacre quadrats
surrounding the logs. With these two methods, he was able to specify
characteristics of flora near logs, but was not able to compare this
with vegetation elsewhere in the habitat.

I suspected that vegetation within a few feet of the drumming bird
might be less dense than nearby, because birds must have space in which
to move their wings, and because the presence of the log might decrease
the abundance of stems. Therefore, I used a series of 5 one-quarter
milacre quadrats at the log. One of these (Number 5 in Fig. 3) was used
to measure vegetation immediately surrounding the drumming location, and
quadrats 1-4 were used to measure vegetation adjacently. Four quadrats
of the same size were located 66 feet from the log, one in each cardinal
direction. Quadrats were used primarily to sample the smaller vegetation.

To sample larger vegetation, I used a system of four, hundreth-
acre transects (6.6 x 66 feet) laid out from the drumming location, one
in each cardinal direction. (Fig. 4.) Each transect was later divided
into "near" and'Tar" halves to determine whether plant density and
composition varied between these portions.

In both sampling methods I tallied only woody vegetation, with more
than half the stem diameter in the sampling unit, and stems were grouped
into the following size classes: 0-2 feet tall, 2-5 feet tall, 5-8 feet
tall, between 8 feet tall and 3 inches diameter at breast height, 3-6
inches dbh, 6-9 inches dbh, and 9 inches dbh and larger. All of these

classes were used while tallying stems on quadrats, whereas to save time

l3

 

 

 

 

 

 

 

 

 

“9&4
/

Z

 

 

 

 

 

 

 

Fig. 3 Sketch of quadrat vegetation sampling design used "near”
the drumming log. Each quadrat is one-quarter milacre
in size. Log is hypothetical.

l4

 

 

 

 

 

 

E, N
3 Log
5
[ ffi Near For

 

 

 

 

 

 

 

Er

Fig. 4 Sketch of trensect vegetation sapling design with "fer"
quedrets. lech transect is one-lundredth'ecre in size.
Quedrsts 6, 7, 8, end 9 ere counterparts of quedrsts 1,

2. 3. end 4.

 

15

the first two classes were omitted on the transects.

Working Procedure

I developed an orderly method of securing the needed information
at each drumming location. First, I measured and recorded log size.
After this I noted its general state of decay and attempted to determine
species. When this was not possible, I recorded whether it had been a
deciduous or coniferous species. The azimuth was then recorded and I
measured the distance from the drumming location to the nearest end.

When these properties were recorded, Quadrat 5 was centered over
the drumming location and laid out using a wire frame measuring 3.3 feet
on a side, and with the sides in north-south and east-west directions.
With this quadrat in place, I marked each corner with small sticks which
later served as corners for Quadrats l, 2, 2, and 4. Stems were then
tallied on Quadrats 1-5. When herbaceous cover was very dense, it was
necessary to further divide the quadrats into portions to facilitate
counting.

After completing these quadrats I imbedded a jack knife in the log,
placed a looped end of a white cord over the knife handle, and laid out
each transect, staking the far end of the cord. I then tallied vegetation
along one side and then another of the transect using the same 3.3 foot
wire to delimit the outer margin of the transect that was used to lay out
the quadrats. After completing each transect, its corresponding distal

quadrat was laid out and tallied.

Plant Density Analysis
I converted total numbers of woody stems of all species tallied on

quadrats and transects to equivalent numbers of stems per acre to determine

16

average densities in each size class necessary for establishment of male

courtship territories (Tables 4, 5, and 6).

TABLE 4

Number of woody stems tallied by size classes on nine
one-quarter milacre quadrats near 40 ruffed grouse
drumming logs, Rifle River Area, Michigan

Number of woody stems tallied
Quadrat Number

 

 

 

Size Class 1 2 3 4 5 6 7 8 9 Total
0-2 ft. 80 76 127 97' 91 167 140 168 214 1160
2-5 ft. 60 38 80 61 21 43 42 6o 79 484
5-8 ft. 24 17 24 17 8 22 22 20 20 174
8 ft.-3 in. dbh 62 4o 71 33 9 3o 20 19 22 306
3-6 in. dbh 9 3 4 4 2 3 1 o 6 32
6-9 in. dbh 3 o 1 o o 1 o o o 5
9 in. + dbh 1 o o 1 o 1 1 o o 4

Total 239 174 307 213 131 267 226 267 341 2165

TABLE 5

Number of woody stems tallied on "near" and "far" halves
of one-hundreth acre transects near 40 ruffed grouse
drumming logs, Rifle River Area, Michigan

 

North East South west

, Transect Transect Transect Transect
Size Class Near §§£_ Near Fag Near F§£_ Near Egg Total
5-8 ft. 290 266 277 242 313 199 321 242 2150
8 ft. to 3in.dbh 646 490 624 488 688 424 632 481 4473
3-6 in. dbh 59 60 70 37 68 52 58 44 448
6-9 in. dbh 14 13 10 7 ll 7 ll 11 84
9 in. + dbh 7 12 3 5 3 6 6 12 54

Total 1016 841 984 779 1083 688 1028 790 7209

17

TABLE 6

Converted average number of woody stems per acre by size
classes encountered on quadrats and transects located
near 40 ruffed grouse drumming logs, Rifle River Area,

 

 

 

 

 

 

 

Michigan.
andrats Average number of
Size Class stem§_per acre
Less than 5 feet tall
0-2 ft. tall 12889
2-5 ft. tall 5378
More than 5 feet tall
5-8 ft. tall 1933
8 ft. tall-3 in. dbh 3400
3-6 in. dbh 478
6-9 in. dbh 56
9 in. dbh 44
Transects
Size Class
Less than 5 feet tall
No sample taken
More than 5 feet tall
5-8 ft. tall 1344
8 ft. tall-3 in. dbh 2796
3'6 ine dbh 280
6-9 in. dbh 53
9 in. + dbh 34

Comparison ofgplant densities "near" and "far" from drummingilggs

My first analysis consisted of determining whether vegetation "near"
the log (quadrats 1-4 and "near" halves of transects) differed basically
from vegetation a distance from the log (quadrats 6-9 and "far" halves of
transects). To do this I used the matched pairs method described by
Goulden (1952) whereby numbers of stems on paired groups of sampling units
("near" and "far") were compared for each log. Comparisons were made by
each size class. This method removed the heterogeneitty in plant composition
and density due to inherent differences between log sites. (Tables 7, 8, 9,
10, ll, 12, l3, 14.) Values of t obtained and results of analyses are

presented in Table 15.

18

TABLE 7

Comparison of woody plant density in the 0-2
foot tall size class on matched pairs of
quadrats near hO ruffed grouse drumming logs,
Rifle River Area, Nflchigan.

 

 

 

 

 

 

d
Number of Number of Quadrats l-h
Log Stems Stems minus
Number Quadrats l-h Quadrats 6-9 Quadrats 6-9
1 6 18 -12
2 11 10 h
3 17 11 6
h 9 2o -11
S 9 27 —18
6 5 6 - 1
7 h 21 -17
8 5 12 - 7
9 18 h 14
10 o 11 —11
11 23 7 16
12 22 23 - 1
13 h 6 - 2
1h 11 lb - 3
15 7 137 -130
16 16 11 S
17 4 9 - S
18 10 8 2
19 7 h 3
2o 9 1h - 5
21 6 7 - 1
22 1 11 .10
23 11 68 -37
2h 11 11 - 3
2S 8 6 2
26 3 18 -15
27 12 2o — 8
28 18 2h 28
29 10 10 o
30 2 9 - 7
31 6 3 3
32 10 21 -11
33 7 13 - 6
3h 3 16 -13
35 O 31 -31
36 12 10 2
37 O 15 ~15
38 12 2h -12
39 10 15 - 5
Lo 8 8 0
2d: -316 __ - 22616-211964
(53'; 2261.6h 5‘“ 16x39 " 35939
. 2896. = 7.9 - o
34?? '3T5939‘
:- = 7.9 = 2.198

19

TABLE 8

Comparison of woody plant density in the 2-5
foot size class on matched pairs of quadrats
near hO ruffed grouse drumming logs, Rifle
River Area, Michigan

 

 

 

 

 

d
Log Number of Stems Number of Stems Quadrats 1-h.minus
Number Quadrats l-h Quadrats 6-9 Quadrats 6-9

1 0 S - S

2 1 o 1

3 20 h 16
h 1 1 o
5 S 3 2
6 11 9 2

7 9 6 3

8 8 8 0
9 12 12 o
10 6 9 — 3
11 13 12 1
12 7 2 5
13 3 S - 2
1h 1 16 -12
15 7 S 2
16 18 20 - 2
17 2o 13 7
18 3 0 3
19 l 7 - 6
2o 0 3 - 3
21 1 3 - 2
22 3 o 3
23 1 o 1
2h 1 o 1
25 2 o 2
26 3 h - 1
27 23 8 15
28 10 12 - 2
29 2 2 o
30 1 3 - 2
31 o 1 - 1
32 0 0 0
33 h S - 1
34 1 8 - 7
35 23 3 2o
36 7 16 - 9
37 7 7 0
38 0 1 - 1
39 0 6 - 6
to 1 h - 3

£J=16 I = 0110
ah- 1424 4 = .12
5.. = .9511

(59‘: 6.11 “
4O

20

TABLE 9

Comparison of woody plant density in the 5-8
foot size class on matched pairs of quadrats
near hO ruffed grouse drumming logs, Rifle
River Area, Michigan.

 

 

 

 

d
Log Number of Stems Number of Stems Quadrats l-h minus
Number Quadrats l-h Quadrats 6-9 Quadrats 6-9
1 1 O O
2 O 2 - 2
3 O 1 - 1
11 o h — h
5 7 0 7
6 1 3 - 2
7 3 2 1
8 1 O 1
9 6 S 1
10 O 3 ' 3
11 10 10 O
12 O 2 - 2
13 1 O 1
lb 8 9 - l
15 2 h - 2
16 11 O 11
17 5 10 - 5
18 O O O
19 O O O
20 1 O 1
21 O O O
22 O 1 - 1
23 O O O
211 1 1 o
25 2 0 2
26 6 6 0
27 1 O 1
28 2 2 O
29 1 O 1
30 O O O
31 3 3 0
32 O O O
33 0 l - 1
3h 0 0 0
35 6 1 5
36 3 h - 1
37 O 7 - 7
38 0 O 0
39 0 2 - 2
ho o 1 - 1
IT= —3 3' - .075

I a

0

1.1
ox
w

£4? 33%; 116
(3: a . S: = .

21

TABLE 10

Comparison of woody plant density in the 8'-3"
dbh size class on matched pairs of quadrats
near hO ruffed grouse drumming logs, Rifle
River Area, Michigan.
d
Log Number of Stems Number of Stems Quadrats 1-4 minus
Number Quadrats l-h Quadrats 6-9 Quadrats 6-9

 

 

 

 

\OODK'IOU'IE'UJNH

1.1
p.)

10

UIWE‘WUIOLDHrWUiE’E‘I-‘wmol-‘l-‘OOOU'Lowl-‘C'NNOO‘wNNOE‘Obn-t

12
13
14
15
16
17
18
19
20
21
22
23
24
25
26
27
28
29
30
31
32
33
34
35
36
37
38
39
LO

1.:
1.1

[.1
1.1

H

HWOCDkAWNWOP’C’NMHt’NNOWI—‘w

H

1.1

OOUlep‘ml-Jwt-P'W ONE‘Owle-J
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I
H

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tare 115 .
a} 1627 (- - 3.151.
a: 330.6 s- .912

40 ‘4

‘1
u
M
g1
\n

I.‘ ll!" Illnl

22

TABLE 11

Comparison of woody plant density in the 3-9"
dbh size class on matched pairs of quadrats
near LO ruffed grouse drumming logs, Rifle
River Area, Michigan.
d
Log Number of Stems Number of Stems Quadrats l-h minus
Number Quadrats l-h Quadrats 6-9 Quadrats 6-9

 

 

 

M

1.4
F‘CDF‘F‘CDFJFJFJFJKUFDCDCDC>F‘F’F‘C)CDCDDDFJCDCDCDC)C)FJF‘CDCJCDCDCDFJc>C>C3UJCD
CDF‘C)C)C>CDFJFJCDCDCDCDCDF‘CDCDCDC)FJCDC>C>C>C>FJC>C)AJFJCDFJCDCJCDFJCDCDCDCDCJ
F‘F‘F‘F‘C)FJC>CDFJC1nJCDC>F‘F‘F‘F‘C)FJC)RJFJCDCJF‘CDC)F‘CDC)PJC>C>CDCDCDC>C3u3C>

id. 13 J = 0325
a} 41 4 = 2.117
(571? 4.225 s: :- .154

 

23

TABLE 12

Comparison of woody plant density in the 5-8
foot class on matched pairs of transects near
40 ruffed grouse drumming logs, Rifle River

Area, Michigan.

 

 

 

 

Log d
Number "Near" Transects "Far“ Transects “Near" minus "Far"
1 21 39 -18
2 12 20 - 8
3 5 6 - 1
4 19 20 - 1
S 27 32 - 5
6 8 44 -36
7 40 40 0
8 25 24 1
9 85 71 14
10 41 38 3
11 77 60 17
12 29 34 - 5
13 58 26 32
14 75 57 18
15 41 29 12
16 124 18 106
17 78 52 26
18 17 5 12
19 15 10 5
20 3 2 1
21 S 3 2
22 11 15 - 4
23 7 3 4
24 7 8 - 1
25 26 13 13
26 40 40 0
27 48 3 45
28 32 39 - 7
29 11 24 413
30 8 10 - 2
31 7 7 O
32 3 0 3
33 9 3 6
34 9 20 .11
35 61 47 14
36 36 41 - 5
37 46 25 21
38 26 5 21
39 3 10 - 7
40 6 6 0
(J - 252 2‘ = 6.3
a‘- 19576 4 = 1.855
5.. = 3.396

(241): 1587.6
40

24

TABLE 13

Comparison of woody plant density in the 8‘-3"
dbh class on matched pairs of transects near
40 ruffed grouse drumming logs, Rifle River
Area, Michigan.

 

 

 

 

 

Log d
Number "Near" Transects "Far" Transects "Near" minus "Far“
1 42 45 - 3
2 67 44 23
3 65 24 41
4 45 44 1
5 52 46 6
6 15 42 -27
7 44 42 2
8 80 44 36
9 207 130 77
10 72 81 - 9
11 225 151 74
12 104 61 43
13 136 69 67
14 136 57 79
15 73 4O 33
16 214 68 146
17 49 20 29
18 43 24 19
19 26 34 - 8
20 72 9 63
21 17 24 - 7
22 46 49 - 3
23 6 8 - 2
24 34 22 12
25 56 47 9
26 70 52 18
27 78 38 40
28 92 74 18
29 20 75 -55
30 26 33 - 7
31 15 22 - 7
32 35 21 14
33 35 29 9
34 28 39 .11
35 110 75 35
36 79 94 -15
37 49 34 15
38 36 33 3
39 18 17 1
40 1 20 25 - 5
gr— 754 - =- 18.85
24';- 63756 J 4 - 3.345
(2J)= 14212.9 5... =- 5.635
176 d

25

TABLE 14

Comparison of woody plant density in the 3-9"
class on matched pairs of transects near 40
ruffed grouse drumming logs, Rifle River Area,

 

 

 

 

Michigan.
Log d
Number "Near" Transects "Far" Transects "Near" minus "Far"
1 2 l l
2 l6 l6 0
3 5 l 4
4 9 9 0
5 l 2 - 1
6 0 0 0
7 10 l 9
8 4 3 l
9 0 2 - 2
10 3 2 1
ll 2 0 2
12 8 10 - 2
l3 3 2 l
14 0 2 - 2
15 4 3 1
16 l 9 - 8
l7 4 6 - 2
18 7 6 l
19 10 5 5
20 24 13 ll
21 14 7 7
22 19 12 7
23 2 4 - 2
24 9 4 5
25 13 15 - 2
26 l 4 - 3
27 2 3 - l
28 13 7 6
29 4 3 l
30 5 15 -10
31 19 5 14
32 19 8 ll
33 2 7 - 5
34 12 6 6
35 3 2 l
36 12 11 l
37 8 5 3
38 ll 3 8
39 25 15 10
40 l 2 - 1
2d 3 76 a = 1.9
(.82 = 1166 t = 2.348
(Ed)2= 164.4 sa = .809

40

26

TABLE 15

Values of t obtained by analyses of stem densities on
quadrats 1-4 compared to quadrats 6-9 and "near” halves
of transects compared to "far" halves surrounding 40
ruffed grouse drumming logs by arbitrary size classes
using the matched pairs technique.*

 

 

 

 

Size Class d t value Remarks

andrats

0-2 feet tall -7.9 2.198 Significant difference in stem
density on near and far quadrats.
More stems on far quadrats.

2-5 feet tall .40 .42 No significant difference in stem
density between near and far
quadrats.

5-8 feet tall .075 .163 No significant difference in stem
density between near and far
quadrats.

8 feet tall-3 inches dbh 2.875 3.154 Significant difference in stem
density on near and far quadrats.
More stems tallied on near quadrats.

3-9 inches dbh .325 2.117 Significant difference in stem
density on near and far quadrats.
More stems tallied on near quadrats.

Transects

5-8 feet tall 6.30 1.855 No significant difference in stem
density between near and far halves
of transects.

8 feet tall-3 inches dbh 18.85 3.345 Significant difference in stem
density on near and far transects.
More stems were tallied on near
transects.

3-9 inches dbh 1.90 2.348 Significant difference in stem

density on near and far transects.
More stems were tallied on near
transects.

 

* The t value for 39 degrees of freedom at the 5 per cent level is 2.02.

27

It is apparent from these analyses that drumming ruffed grouse
require a dense layer of undergrowth. The understory stratum in this
study area is typified by alder runs or dogwood clumps. In the two size
classes smaller than the 8 foot-3 inch dbh class I was not able to show
differences in stem densities adjacent to the logs as compared to several
feet from the logs. In the 0-2 foot tall size class I found significantly
fewer stems immediately surrounding the log than elsewhere.

These findings at first appear contradictory, but the analyses are
reasonable when the history of land use and the pressure of a large deer
population are considered. The woody plants in the 0-2 foot tall size
class represent young reproduction. It is probable that deer populations
have little influence on the abundance of stems in this class. The next
two size classes however, are subject to constant browse pressure by deer.
Therefore the understory stratum throughout the Rifle River Area has been
reduced or eliminated in some cases. The 8 foot to 3 inch dbh class is
beyond the reach of deer. I made a comparison of the prevalence of deer
food and non-deer food plants tallied during the study (Table 16) and
the results illustrate how deer have affected plant composition. Non-deer
food plants are those which deer usually refuse to eat even when subjected
to extreme hardship (Duvendeck, 1952).

TABLE 16
Number of stems of woody deer food plants and non-deer

food plants tallied by size classes on 360 quadrats
near 40 ruffed grouse drumming logs, Rifle River Area,

 

 

 

Michigan
Size Class . Deer Food Plants Non-deer Food Plants
0-2 feet tall 656 493
2-8 feet tall 143 526

8 feet to 3 inches dbh 30 274

28

Plant Composition

All 40 logs were located in a lowland vegetative type. Thirty-seven
woody plant species were tallied on the 360 quadrats. These 37, as well
as 7 additional species were found on the transects. A check list is
presented in the Appendix. Nomenclature is from Fernald (1950).

Many species were usually represented on the sampling units i.e. at
least one stem occurred. This was particularly true in the smallest size
class (Table 17). In this size class, 8 different species occurred in
more than 5 per cent of the quadrats. Fewer species were encountered in
the larger classes and these few plants therefore assumed more relative
importance. In fact, in the 8 feet tall to 3 inch dbh class only 4 species
occurred regularly. These included speckled alder (Alnus rugosa), balsam

fir (Abies balsamea), black ash (Fraxinus nigra), and willows (Salix spp.).

 

Alder, moreover, was tallied much more frequently than the others, and it
occurred on 21 per cent of the quadrats.

Alder not only occurred frequently, but it also supplied the necessary
density of stems which perhaps was more important (Table 18). In the very
important tall shrub class (8 feet tall to 3 inches dbh) 73 per cent of the

woody stems tallied were speckled alder.

Plant Densities Compared by Number of Years Logs were Used.
Calendar years when ruffed grouse used the 40 drumming logs were
plotted (Fig. 1), and the number occurring in each years-of-use class

is presented in Table 19.

1‘

PI

29

TABLE 17
Number of quadrats in which at least one stem of the 37
plant species occurred and the computed relative frequency

of each species by size classes.*

Size Class

 

Species 0-2' 2-8' 8'-3" 3-6" 6-9" 9"+
Alder 76(21.1) 138(38.3) 77(21.4)

Red-osier 56(15.6) 29(8.1) 1(.3)

Black ash 38(10.6) 12(3.3) 12(3.3)

Black currant 30(8.3) 6(l.7)

Mountain holly 29(8.1) 27(8)(7.5) 4(l.l)

Balsam fir 29(8.1) 9(2.5) 15(4.2) l4(3.9) 2(.6)
Gray dogwood 30(8.3) ll(3)(3.l)

Trembling aspen 10(2.8) 2(.6) 2(.6) 4(l.l) 1(.3) 1(.3)
Nannyberry l8(5.0) 7(1.9) 3(.8)

Red oak l6(4.4) 2(.6)

Red maple 25(6.9) 4(l.l) 1(.3)

Blueberry ll(3.1)

Willow 6(1.7) l6(4.4) lO(2.8)

Chokeberry ll(3.1) 8(2.2) 2(.6)

Silky dogwood 7(1.9)

White cedar 2(.6) 3(.8) 4(l.l) 2(.6)
Ninebark 1(.3)

Leatherleaf 2(.6) 1(.3)

White spruce 5(1.4) 4(l.l) 2(.6)

White birch 2(.6) 2(.6)
High-bush cranberry 1(.3)

Juneberry .2(.6)

Black cherry 7(1.9)

Hawthorn 5(l)(l.4)

Honeysuckle 3(.8)

Large-tooth aspen 1(.3)

Hazelnut 7(1.9)

Larch 1(.3) 2(.6)
Balsam poplar 1(.3) 1(.3) 1(.3)
Slippery elm 1(.3) 1(.3)

American elm 2(.6) 1(.3)

White ash 1(.3)

Prickly ash 1(.3) 1(.3)

Meadow-sweet 1(.3) 1(.3)

White pine 1(.3)

White oak 1(.3)

Hemlock 1(.3)
Unknown 3(.8) 1(.3)

* Figures in parenthesis are percentages.
computed by dividing the number of quadrats in which the species was
tallied by the total number of quadrats (360).

The relative frequency is

30

TABLE 18

sites.1

 

Total stems and relative density of woody plant species
occurring on 360 quadrats near 40 ruffed grouse drumming

 

total stems recorded by species by size classes.

_Size Class Total
(Species 0-2' 2-5' 5-8' 8'-3" 3-6" 9"+ stems
Alder 186(l6.0)234(47.4)106(60.6) 223(73.l) 750
Red-osier l60(l3.7) 52(10.5) 1(.6) 213
Blueberry 13l(ll.2) 131
Balsam fir 91(7.8) 3(.6) 6(3.4) 26(8.5) 17(54.8) 2(40) 145
Gray dogwood 88(7.6) 21(4.2) 3(l.7) 1(.3) 113
Black ash 73(6.3) l6(3.2) 2(1.l) 12(3.9) 5(16.l) 1(25) 109
Mountain holly 60(5.2) 79(15.9) 16(9.1) 7(2.3) 162
Black currant 55(4.7) 8(l.6) 63
Nannyberry 52(4.5) 9(l.8) 12(6.9) 4(l.3) 77
Red maple 48(4.1) 4(.8) 1(.3) 53
Leatherleaf 32(2.7) 30(6.0) 62
Meadow-sweet 26(2.2) 5(l.0) 31
Chokeberry 21(1.8) 7(1.4) 5(2.9) 2(.7) 35
Silky dogwood 17(l.5) 17
Red oak 18(1.5) 3(.6) 21
Trembling aspen l4(l.2) 1(.2) 1(.6) 2(.7) 4(12.9) 1(20) 1(25) 24
Black cherry l6(l.4) 1(.3) 17
Willow 12(1.0) 10(2.0) l7(9.7) 15(4.9) 54
Hawthorn 8(.7) 8
White spruce 7(.6) 3(.6) 3(l.7) 2(.7) 1(25) l6
Hazelnut 7(.6) 7
Honeysuckle 4(.3) 4
Large-tooth aspen 4(.3) 4
Prickly ash 4(.3) 2(.4) 6
Juneberry 3(.3) 3
American elm 3(.3) 1(.3) 4
White cedar 2(.2) 1(.2) 3(l.7) 5(l.6) 2(6.5) 13
Ninebark 1(.1) l
Highbush cranberry 1(.1) l
Slippery elm 2(.2) 1(.2) 3
White ash 1(.1) 1
White pine 1(.1) 1
White oak 1(.1) l
Hemlock 1(25) 1
Larch 1(.6) 2(6.5) 3
Balsam poplar 2(.4) 1(.3) 1(3.2) 4
White birch 2(.7) 2(6.5) 4
Unknown l6(l.4) 3(.6) 19
Totals 1165 496 175 305 31 4 2181
1

Figures in parentheses represent relative density, which is the percentage of

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'x

31

TABLE 19

Number of drumming logs occupied by male ruffed grouse
during 8 calendar years, Rifle River Area, Michigan

Number of years occupied

Number of logs 9 5 4 l3 5 0 2 2

In addition to size being a factor responsible for perennial
selection in some cases, I suspected that plant densitiés near the logs
might differ between logs used varying lengths of time.

Analysis of variance was used to test whether there was a relationship
between stem density and the frequency of log use (by years). Tests were
performed by the various size classes (Tables 20, 21, 22 and 23). F values
obtained in analyses of all size classes were less than the required value
for 5, 34 degrees of freedom at the 5 per cent level of 2.49. Thus the
analyses suggested there was no relationship between stem density and
frequency of log use.

From analyses of log size and vegetation density therefore, it is
concluded that grouse require logs and surrounding vegetation of certain
specifications. If these specifications are met, the preference for some
logs from year to year cannot be explained at this time. Some other factors
such as population size, or perhaps a spatial relationship exists between
one site to another in the total environment which overshadows minor

vegetational differences at the different sites.

v“

32

TABLE 20

Analysis of variance of density of woody vegetation in

the 0-2 foot height class on quadrats 1-4 by numbers of

years 40 ruffed grouse drumming logs were occupied, Rifle
River Area, Michigan

Number of years logs were used by grouse

 

 

 

 

 

 

 

 

 

l 2 3 4 5 7 and 8
8
6 23 16 11 10 0
14 22 4 ll 7 12
Number of 17 4 10 0 3 10
woody stems 9 ll 0 7 0 8
tallied in 9 7 6 12
each year class 5 9
4 48
5 2
l8 6
3
12
10
Ave. No. Stems 9.7 13.4 7.5 10.2 6.4 7.5
Estimate of "between" year class variance = 32.66 a 32.7 a 43
Estimate of "within" year class variance = 75.56 75.6 '
TABLE 21
Analysis of variance of density of woody vegetation in
the 2-8 foot height class on quadrats 1-4 by numbers of
years 40 ruffed grouse drumming logs were occupied,
Rifle River Area, Michigan
Number of years logs were used byggrouse
l 2 3 4 5 7 and 8
12 7 6 l 23 1
Number of 12 3 4 12 l l
woody stems l8 1 3 9 l 4
tallied in l 10 l 9 3 29
each year 9 12 29 7
class 1 25
0 3
20 2
l 24
0
0
12
4
Ave._No. Stems 8.2 6.7 3.5 10.0 7.0 8.8
Estimate of ”between? year class variance = 30.10 F _ 30.10 = .33

Estimate of "within" year class variance 3 79.54 79.54

33

TABLE 22

Analysis of variance of density of woody vegetation in

the 8'-3" dbh class on quadrats 1-4 by numbers of years

40 ruffed grouse drumming logs were occupied, Rifle
River Area, Michigan.

Number of years logs were used by grouse

 

 

 

 

l 2 3 4 5 7 and 8
Number of 4 5 8 l 23 7
stems tallied 3 4 5 ll 4 0
by year classes 9 0 2 4 l 0
0 3 3 3 5 15
l 12 19 3
1 6
2 4
12 4
2 9
6
3
0
3
Ave. No. Stems 3.8 4.8 4.5 5.6 7.2 5.5
Estimate of ”between" year class variance = 8.7 F = 8.7
Estimate of "within" year class variance =29.6 29.6
TABLE 23

Analysis of variance of density of woody vegetation in

the 3-9” dbh class on quadrats 1-4 by numbers of years

40 ruffed grouse drumming logs were occupied, Rifle
River Area, Michigan.

Number of years logs were used by grouse

 

 

 

l 2 3 4 5 7 and 8
Number of 1 1 0 0 0 l
woody stems l l 0 0 1 1
tallied by 1 0 0 0 l 0
year classes 3 0 0 1 l
0 0 0 l
0 0
0 3 1
0 2
0 0
l
2
l
1
Ave. No. Stems .67 .4 0 .85 .8 .75

 

Estimate of "between” year class variance = 2.65 F a .12
Estimate of "within" year class variance =22.45

'34

DISCUSSION

During this population study various investigations of grouse
ecology enabled me to become familiar with habitat quality in many parts
of northern lower Michigan.

Although foresters are guided by multiple-use policies, in many
areas mature or near-mature stands of aspen, oak, and jack pine are not
being lumbered adequately, at least from a game standpoint. In many
cases stands of pole-size or larger trees are characterized by scanty
undergrowth, and in some areas the situation has become critical because
of excessive browse removal by deer (Fig. 5, 6, 7).

In some counties over-numerous deer have been a problem for as long
as 30 years (Jenkins and Bartlett, 1959). Lowland flora is composed
largely of woody plants, which are unpalatable to deer, such as speckled
alder, white spruce, balsam fir, and a few other species.

Upland areas where these conditions exist such as on the Rifle River
Area probably are not supporting maximum numbers of ruffed grouse.

In most of Michigan ruffed grouse are underhunted. Beaause of this
and also because habitat improvement specifically for ruffed grouse is
expensive, we must rely on standard forestry practices to maintain or
create ideal grouse habitat. Only an intensively used lands, such as our
game areas in southern Michigan, can intensively directed management be
justified. Thus it is doubly important to reduce or maintain deer
populations at a sufficiently low level so that habitat quality is main-
tained at a level suitable for maximum grouse production concomitant with

other land uses.

35

 

 

Fig. 5
Fig. 6
Fig. 7

The upland aspen
type with scanty
understory occurs
abundantly in
northern lower
Michigan.

A jack pine forest
near Luzerne, Oscoda
County, Michigan.
This vegetation type
is also abundant in
northern lower
Michigan. Both types
illustrated represent
poor ruffed grouse
habitat and could be
noticeably improved.

The "edge" between
upland and lowland
sepen types. Note
the development of
the lowland under-
story, composed al-
most entirely of
speckled alder.
Forest age and deer
browsing pressure
have eradicated the
understory stratum
in the upland.

36

SUMMARY

Forty ruffed grouse drumming logs and the nearby vegetation were
studied in 1959. Minimum size of logs and plant density requirements
were ascertained.

Logs were occupied by grouse for varying numbers of years. Logs
used several years tended to be larger in diameter than those used in-
frequently, but there was little correlation between use and length of
logs. Log directions occurred most often in a sourthwesterly - north-
easterly direction, but it was not determined whether this was due to
prevailing winds or whether grouse prefer this direction. Birds selected
drumming locations near one end of the log, and usually this was the
'large or "butt" end. More than three-fourths of the logs were old
conifers, of which most were white pine.

No consistent differences in plant densities were noted near logs
used almost perennially as compared to logs used infrequently. Vegetation
surrounding logs was also compared to vegetation elsewhere in the habitat,
and comparisons were made by arbitrary size classes. Plants in the
smallest size class (0-2 feet tall) were less numerous near the logs
than elsewhere. In the larger size classes, particularly the 8 feet tall
to 3 inch dbh class, plants were much more numerous near the log.

All logs were located in lowland vegetation types. The presence of
all logs in lowland types suggests that the upland types are not suitable
for the establishment of courtship sites. This unsuitability was probably
due to a paucity of undergrowth, resulting from an advanced forest age

and also perhaps from an overabundance of deer. Lowland types were

M—v‘-—‘

37

satisfactory as courtship sites, because plants growing there were
unpalatable to deer and therefore escape serious browsing and can
reproduce. Very few plant species supplied practically all of the
necessary density, although 44 different species were tallied. Speckled
alder (Alnus rugosa) was undoubtedly the most important single plant
species.

Ecological conditions on the Rifle River Area are believed to be
representative of many areas in Michigan, and if maximum ruffed grouse
production is desired, deer numbers must be reduced and lumbering
rotations shortened. In some areas, more intensive habitat improvement
measures may be necessary than can be supplied by the removal of mature
trees, but the least expensive means to create or maintain habitat

quality is through deer herd reduction.

38
LITERATURE CITED

Bump, Gardiner, Robert W. Darrow, Frank C. Edminster and Walter F. Crissey,
1947. The ruffed grouse-life history, propagation management. New
York State Cons. Dept., Albany, 915 pp.

Cottam, Grant and J. T. Curtis, 1956. The use of distance measures in
phytosociological sampling. Ecology 37:451-460.

Dorney, Robert S. 1959. Relationship of ruffed grouse to forest cover
types in Wisconsin. Technical Bulletin No. 18. Wisconsin Cons. Dept.,
Madison, 32 pp.

Duvendeck, Jerry Paul 1952. Some effects of deer browsing on northern
Michigan forest plants. Unpublished Master's thesis, Michigan State
University, East Lansing, Mich. 25 pp.

Edminster, Frank C. 1947. The ruffed grouse, its life story, ecology and
management. The Macmillan Co., New York. 385 pp.

Eng, Robert L. 1959. A study of the ecology of male ruffed grouse

(Bonasa umbellus L.) on the Cloquet Forest Research Center, Minnesota.

 

Unpublished Ph.D. thesis, University Minn., Minneapolis, 107 pp.

Fernald, Merritt Lyndon. 1950. Gray's manual of botany. American Book
Company, New York. 1632 pp.

Frank, William J. 1947. Ruffed grouse drumming site counts. J. Wildl.
Mgmt., 11:307-316.

Goulden, Cyril H. 1952. Methods of statistical analysis. John Wiley and
Sons, Inc., New York, Longon. 467 pp.

Jenkins, David H. and 110 H. Bartlett. 1959. Michigan Whitetails. Mich.
Dept. Conservation, Lansing, 80 pp.

Palmer, Walter L. 1956. Ruffed grouse population studies on hunted and
unhunted areas. Trans. let North American Wildlife Conference.

Ryel, L. A. and L. L. Eberhardt. 1960. Unpublished Game Division report

No. 2288, Michigan Dept. Conservation, Lansing, 8 pp.

39

APPENDIX

40

Check list of woody plants tallied near 40 grouse drum-
ming logs. Nomenclature from Fernald, 1950.

Balsam fir (Abies balsamea)

Hemlock (Tsuga canadensis)

White spruce (Picea glauca)

Larch (Larix laricina)

White pine (Pinus Strobus)

Red pine (Pinus resinosa)

White cedar (Thuja occidentalis)
Willow (Salix spp.)

Trembling aspen (Populus tremuloides)
Large-tooth aspen (Populus grandidentata)
Balsam poplar (Populus balsamifera)
American hazel (Corylus americana)
White birch (Betula papyrifera)
Yellow birch (Betula lutea)

Swamp birch (Betula pumila)

Speckled alder (Alnus rugosa)

White oak (Quercus alba)

Red oak (Quercus rubra)

Slippery elm (Ulmus rubra)

American elm (Ulmus americanum)

Rock elm (Ulmus Thomasi)

Black currant (Ribes americanum)
Ninebark (Physocarpus opulifolia)
Meadow-sweet (Spiraea alba)
Chokeberry (Pyrus melanocarpa)
Juneberry (Amelanchier huronensis)
Hawthorn (Crataegus sp. )

Pin cherry (Prunus pennsylvanica)
Black cherry (Prunus serotina)

Choke cherry (Prunus virginiana)
Prickly ash (Xanthoxylum americanum)
Mountain holly (Nemopanthus mucronata)
Red maple (Acer rubrum)

Red-osier dogwood (Cornus stolonifera)
Gray dogwood (Cornus racemosa)

Silky dogwood (Cornus obliqua)
Leatherleaf (Chamaedaphne calyculata)
Blueberry (Vaccinium sp.)

White ash (Fraxinus americana)

Black ash (Fraxinus nigra)
Honeysuckle (Lonicera canadensis)
Nannyberry (Viburnum lentago)
High-bush cranberry (Viburnum trilobum)
Elderberry (Sambucus canadensis)

 

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